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Western Hunter-Gatherer

April 08, 2024

In archaeogenetics, the term Western Hunter-Gatherer (WHG), West European Hunter-Gatherer, Western European Hunter-Gatherer, Villabruna cluster, or Oberkassel cluster (c. 15,000~5,000 BP) is the name given to a distinct ancestral component of modern Europeans, representing descent from a population of Mesolithic hunter-gatherers who scattered over Western, Southern and Central Europe, from the British Isles in the west to the Carpathians in the east, following the retreat of the ice sheet of the Last Glacial Maximum.[2]

Western Hunter-Gatherer
Genetic ancestry of hunter-gatherers in Europe between 14 ka and 9 ka, with the main area of Western Hunter-Gatherers (WHG) in blue. Individual numbers correspond to calibrated sample dates.[1]

Along with the Scandinavian Hunter-Gatherers (SHG) and Eastern Hunter-Gatherers (EHG), the WHGs constituted one of the three main genetic groups in the postglacial period of early Holocene Europe.[3] The border between WHGs and EHGs ran roughly from the lower Danube, northward along the western forests of the Dnieper towards the western Baltic Sea.[2]

SHGs were in turn a nearly equal mix of WHGs and EHGs. Once the main population throughout Europe, the WHGs were largely displaced by successive expansions of Early European Farmers (EEFs) during the early Neolithic, but experienced a resurgence during the Middle Neolithic. During the Late Neolithic and Early Bronze Age, Western Steppe Herders (WSHs) from the Pontic–Caspian steppe embarked on a massive expansion, which further displaced the WHGs. Among modern-day populations, WHG ancestry is most common among populations of the eastern Baltic.[4]

Research edit

Western Hunter-Gatherers (WHG) are recognised as a distinct ancestral component contributing to the ancestry of most modern Europeans.[5] Most Europeans can be modeled as a mixture of WHG, EEF, and WSH from the Pontic–Caspian steppe.[6] WHGs also contributed ancestry to other ancient groups such as Early European Farmers (EEF), who were, however, mostly of Anatolian descent.[5] With the Neolithic expansion, EEF came to dominate the gene pool in most parts of Europe, although WHG ancestry had a resurgence in Western Europe from the Early Neolithic to the Middle Neolithic.[7]

Expansion into continental Europe (14,000 BP) edit

WHGs themselves are believed to have formed around 14,000 years ago, during the Bølling-Allerød interstadial, at the time of the first major warming period after the Ice Age. They represent a major population shift within Europe at the end of the Ice Age, probably a population expansion into continental Europe, from Southeastern European or West Asian refugia.[8] It is thought that their ancestors separated from eastern Eurasians around 40,000 BP, and from Ancient North Eurasians (ANE) prior to 24,000 BP (the estimated age date of the Mal'ta boy). This date was subsequently put further back in time by the findings of the Yana Rhinoceros Horn Site to around 38kya, shortly after the divergence of West-Eurasian and East-Eurasian lineages.[5][9] Vallini et al. 2022 argues that the dispersal and split patterns of West Eurasian lineages was not earlier than c. 38,000 years ago, with older Initial Upper Paleolithic European specimens, such as Zlaty Kun, Peștera cu Oase and Bacho Kiro, being unrelated to Western Hunter-gatherers but closer to Ancient East Eurasians or basal to both.[10] The WHG displayed higher affinity for ancient and modern Middle Eastern populations when compared against earlier Paleolithic Europeans such as Gravettians. The affinity for ancient Middle Eastern populations in Europe increased after the Last Glacial Maximum, correlating with the expansion of WHG (Villabruna or Oberkassel) ancestry. There is also evidence for bi-directional geneflow between WHG and Middle Eastern populations as early as 15,000 years ago. WHG associated remains belonged primarily to the human Y-chromosome haplogroups I-M170 with a lower frequency of C-F3393 (specifically the clade C-V20/C1a2), which has been found commonly among earlier Paleolithic European remains such as Kostenki-14 and Sungir. The paternal haplogroup C-V20 can still be found in men living in modern Spain, attesting to this lineage's longstanding presence in Western Europe. Their mitochondrial chromosomes belonged primarily to haplogroup U5.[11][12]

 
Last Glacial Maximum refugia, c. 20,000 years ago
  Solutrean culture
  Epigravettian culture[11]

In a genetic study published in Nature in March 2023, the authors found that the ancestors of the WHGs were populations associated with the Epigravettian culture, which largely replaced populations associated with the Magdalenian culture about 14,000 years ago (the ancestors of the Magdalenian-associated individuals were the populations associated with the western Gravettian, Solutrean and Aurignacian cultures).[11][13] In the study, WHG ancestry is renamed 'Oberkassel ancestry', first found north of the Alps in two 14,000 year-old individuals at the eponymous site at Oberkassel, who can be modeled as an admixture of Villabruna ancestry (itself modeled as an admixture between a lineage related to the Věstonice cluster and a lineage ancestral to the Kostenki-14 and Goyet Q116-1 individuals), and Goyet-Q2 ancestry related to individuals found in Europe prior to the Last Glacial Maximum. The study states that all of the individuals of the Oberkassel cluster could be modeled as c. 75% Villabruna and 25% Goyet-Q2 ancestry or, alternatively, as c. 90% Villabruna and 10% Fournol ancestry, a newly identified cluster described as a sister lineage of the Goyet Q116-1 ancestry found in individuals associated with the Gravettian culture of southwestern Europe.[11] The study suggests that Oberkassel ancestry was mostly already formed before expanding, possibly around the west side of the Alps, to Western and Central Europe and Britain, where sampled WHG individuals are genetically homogeneous. This is in contrast to the arrival of Villabruna and Oberkassel ancestry to Iberia, which seems to have involved repeated admixture events with local populations carrying high levels of Goyet-Q2 ancestry. This, and the survival of specific Y-DNA haplogroup C1 clades previously observed among early European hunter-gatherers, suggests relatively higher genetic continuity in southwest Europe during this period.[11]

There are indications that the WHG carried "risk alleles for diabetes and Alzheimer's disease".[14]

 
Transition from Magdalenian Goyet ancestry (green  , Goyet Q2) to Western Hunter Gatherer (WHG) Villabruna ancestry (orange  ) in European sites, according to timeline and climate evolution.[15]

Interaction with other populations edit

 
Cheddar Man, found in Great Britain, had a similar genotype to other Western Hunter-Gatherers.

The WHG were also found to have contributed ancestry to populations on the borders of Europe such as early Anatolian farmers and Ancient Northwestern Africans,[16] as well as other European groups such as Eastern Hunter-Gatherers.[17] The relationship of WHGs to the EHGs remains inconclusive.[17] EHGs are modeled to derive varying degrees of ancestry from a WHG-related lineage, ranging from merely 25% to up to 91%, with the remainder being linked to geneflow from Paleolithic Siberians (ANE) and perhaps Caucasus hunter-gatherers. Another lineage known as the Scandinavian Hunter-Gatherers (SHGs) were found to be a mix of EHGs and WHGs.[a][19][20]

People of the Mesolithic Kunda culture and the Narva culture of the eastern Baltic were a mix of WHG and EHG,[21] showing the closest affinity with WHG. Samples from the Ukrainian Mesolithic and Neolithic were found to cluster tightly together between WHG and EHG, suggesting genetic continuity in the Dnieper Rapids for a period of 4,000 years. The Ukrainian samples belonged exclusively to the maternal haplogroup U, which is found in around 80% of all European hunter-gatherer samples.[22]

People of the Pit–Comb Ware culture (CCC) of the eastern Baltic were closely related to EHG.[23] Unlike most WHGs, the WHGs of the eastern Baltic did not receive European farmer admixture during the Neolithic. Modern populations of the eastern Baltic thus harbor a larger amount of WHG ancestry than any other population in Europe.[21]

SHGs have been found to contain a mix of WHG components who had likely migrated into Scandinavia from the south, and EHGs who had later migrated into Scandinavia from the northeast along the Norwegian coast. This hypothesis is supported by evidence that SHGs from western and northern Scandinavia had less WHG ancestry (ca 51%) than individuals from eastern Scandinavia (ca. 62%). The WHGs who entered Scandinavia are believed to have belonged to the Ahrensburg culture. EHGs and WHGs displayed lower allele frequencies of SLC45A2 and SLC24A5, which cause depigmentation, and OCA/Herc2, which causes light eye color, than SHGs.[24]

 
The rock shelter where the skeleton of the Loschbour man (c. 8,000 BP) was found

The DNA of eleven WHGs from the Upper Palaeolithic and Mesolithic in Western Europe, Central Europe and the Balkans was analyzed, with regards to their Y-DNA haplogroups and mtDNA haplogroups. The analysis suggested that WHGs were once widely distributed from the Atlantic coast in the West, to Sicily in the South, to the Balkans in the Southeast, for more than six thousand years.[25] The study also included an analysis of a large number of individuals of prehistoric Eastern Europe. Thirty-seven samples were collected from Mesolithic and Neolithic Ukraine (9500-6000 BC). These were determined to be an intermediate between EHG and SHG, although WHG ancestry in this population increased during the Neolithic. Samples of Y-DNA extracted from these individuals belonged exclusively to R haplotypes (particularly subclades of R1b1) and I haplotypes (particularly subclades of I2). mtDNA belonged almost exclusively to U (particularly subclades of U5 and U4).[25] A large number of individuals from the Zvejnieki 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 of haplogroup R1b1a1a and I2a1. The mtDNA belonged exclusively to haplogroup U (particularly subclades of U2, U4 and U5).[25] Forty individuals from three sites of the Iron Gates Mesolithic in the Balkans were also analyzed. These individuals were estimated to be of 85% WHG and 15% EHG descent. The males at these sites carried exclusively haplogroup R1b1a and I (mostly subclades of I2a) haplotypes. mtDNA belonged mostly to U (particularly subclades of U5 and U4).[25] People of the Balkan Neolithic were found to harbor 98% Anatolian ancestry and 2% WHG ancestry. By the Chalcolithic, people of the Cucuteni–Trypillia culture were found to harbor about 20% hunter-gatherer ancestry, which was intermediate between EHG and WHG. People of the Globular Amphora culture were found to harbor ca. 25% WHG ancestry, which is significantly higher than Middle Neolithic groups of Central Europe.[25]

Replacement by Neolithic farmers edit

Further information: Neolithic Europe and Genetic history of Europe
 
Residual genetic ancestry of European hunter-gatherers during the European Neolithic, between 7.5 ka and 5 ka BP (c. 5,500~3,000 BC)
 
Simplified model for the demographic history of Europeans during the Neolithic period and the Neolithic Revolution which introduced agriculture[26]

A seminal 2014 study first identified the contribution of three main components to modern European lineages: the Western Hunter Gatherers (WHG, in proportions of up to 50% in Northern Europeans), the Ancient North Eurasians (ANE, Upper Palaeolithic Siberians later associated with the later Indo-European expansion, present in proportions up to 20%), and finally the Early European Farmers (EEF, agriculturists of mainly Near Eastern origin who migrated to Europe from circa 8,000 BP, now present in proportions from around 30% in the Baltic region to around 90% in the Mediterranean). The Early European Farmer (EEF) component was identified based on the genome of a woman buried c. 7,000 years ago in a Linear Pottery culture grave in Stuttgart, Germany.[27]

This 2014 study found evidence for genetic mixing between WHG and EEF throughout Europe, with the largest contribution of EEF in Mediterranean Europe (especially in Sardinia, Sicily, Malta and among Ashkenazi Jews), and the largest contribution of WHG in Northern Europe and among Basque people.[28]

Since 2014, further studies have refined the picture of interbreeding between EEF and WHG. In a 2017 analysis of 180 ancient DNA datasets of the Chalcolithic and Neolithic periods from Hungary, Germany and Spain, evidence was found of a prolonged period of interbreeding. Admixture took place regionally, from local hunter-gatherer populations, so that populations from the three regions (Germany, Iberia and Hungary) were genetically distinguishable at all stages of the Neolithic period, with a gradually increasing ratio of WHG ancestry of farming populations over time. This suggests that after the initial expansion of early farmers, there were no further long-range migrations substantial enough to homogenize the farming population, and that farming and hunter-gatherer populations existed side by side for many centuries, with ongoing gradual admixture throughout the 5th to 4th millennia BC (rather than a single admixture event on initial contact).[29] Admixture rates varied geographically; in the late Neolithic, WHG ancestry in farmers in Hungary was at around 10%, in Germany around 25% and in Iberia as high as 50%.[30]

Analysis of remains from the Grotta Continenza in Italy showed that out of six remains, three buried between c. 10,000 BC and 7000 BC belonged to I2a-P214; and two-times the maternal haplogroups U5b1 and one U5b3.[31][32] Around 6000 BC, the WHGs of Italy were almost completely genetically replaced by EEFs (two G2a2) and one Haplogroup R1b, although WHG ancestry slightly increased in subsequent millennia.[33]

Neolithic individuals in the British Isles were close to Iberian and Central European Early and Middle Neolithic populations, modeled as having about 75% ancestry from EEF with the rest coming from WHG in continental Europe. They subsequently replaced most of the WHG population in the British Isles without mixing much with them.[34]

The WHG are estimated to have contributed between 20-30% ancestry to Neolithic EEF groups throughout Europe. Specific adaptions against local pathogens may have been introduced via the Mesolithic WHG admixture into Neolithic EEF populations.[35]

A study on Mesolithic hunter-gatherers from Denmark found that they were related to contemporary Western hunter-gatherers, and are associated with the Maglemose, Kongemose and Ertebølle cultures. They displayed "genetic homogeneity from around 10,500 to 5,900 calibrated years before present", until "Neolithic farmers with Anatolian-derived ancestry arrived". The transition to the Neolithic period was "very abrupt and resulted in a population turnover with limited genetic contribution from local hunter-gatherers. The succeeding Neolithic population has been associated with the Funnelbeaker culture.[36]

Physical appearance edit

 
Reconstructed head of the Cheddar Man, found in England (carbon-dated c. 8540-8230 BC), based on the shape of his skull and DNA analysis, shown at the National History Museum in London (2019).[37][38]

According to David Reich, DNA analysis has shown that Western Hunter Gatherers were typically dark skinned, dark haired, and blue eyed.[39] The dark skin was due to their Out-of-Africa origin (all Homo sapiens populations having had initially dark skin), while the blue eyes were the result of a variation in their OCA2 gene, which caused iris depigmentation.[40]

Archaeologist Graeme Warren has said that their skin color ranged from olive to black, and speculated that they may have had some regional variety of eye and hair colors.[41] This is strikingly different from the distantly related Eastern Hunter-Gatherers (EHG)—who have been suggested to be light-skinned, brown-eyed or blue eyed and dark-haired or light-haired.[42]

Two WHG skeletons with incomplete SNPs, La Braña and Cheddar Man, are predicted to have had dark or dark to black skin, whereas two other WHG skeletons with complete SNPs, "Sven" and Loschbour man, are predicted to have had dark or intermediate-to-dark and intermediate skin, respectively.[43][24][b] Spanish biologist Carles Lalueza-Fox said the La Braña-1 individual had dark skin, "although we cannot know the exact shade."[45]

According to a 2020 study, the arrival of Early European Farmers (EEFs) from western Anatolia from 8500 to 5000 years ago, along with Western Steppe Herders during the Bronze Age, caused a rapid evolution of European populations towards lighter skin and hair.[40] Admixture between hunter-gatherer and agriculturist populations was apparently occasional, but not extensive.[46]

 
Evolution of Upper Paleolithic and Neolithic phenotypes in Eurasia. Dark-skinned Western Hunter-Gatherers resided in Western Europe, and expanded to some extent towards north and eastern Europe.[40]

Some authors have expressed caution regarding skin pigmentation reconstructions: Quillen et al. (2019) acknowledge studies that generally show that "lighter skin color was uncommon across much of Europe during the Mesolithic", including studies regarding the “dark or dark to black” predictions for the Cheddar Man, but warn that "reconstructions of Mesolithic and Neolithic pigmentation phenotype using loci common in modern populations should be interpreted with some caution, as it is possible that other as yet unexamined loci may have also influenced phenotype."[47]

Geneticist Susan Walsh at Indiana University–Purdue University Indianapolis, who worked on Cheddar Man project, said that "we simply don't know his skin colour".[48] German biochemist Johannes Krause stated that we do not know whether the skin color of Western European hunter-gatherers was more similar to the skin color of people from present-day Central Africa or people from the Arab region. It is only certain that they did not carry any known mutation responsible for the light skin in subsequent populations of Europeans.[49]

A 2024 research into the genomic ancestry and social dynamics of the last hunter-gatherers of Atlantic France has stated that "phenotypically, we find some diversity during the Late Mesolithic in France", at which two of the WHG's sequenced in the study "likely had pale to intermediate skin pigmentation", but "most individuals carry the dark skin and blue eyes characteristic of WHGs" of the studied samples.[50]

Notes edit

  1. ^ 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.[18]
  2. ^ These predictions were obtained using a multinomial logistic regression model based on a panel of 36 carefully selected SNPs with a low sensitivity of 0.26 for classifying intermediate skin (compared to 0.99 and 0.90 for white and black skin, respectively). The accuracy of the model used could be further improved with "additional (but currently unknown) SNP predictors once identified via future GWAS".[44]

References edit

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  38. ^ Details on the reconstruction from the Natural History Museum: "Cheddar Man: Mesolithic Britain's blue-eyed boy". www.nhm.ac.uk. Natural History Museum.
  39. ^ Reich, David (2018). Who We Are and How We Hot Here : Ancient DNA and the New Science of the Human Past (First ed.). New York: Knopf Doubleday Publishing Group. ISBN 978-1101870334. "Analysis of ancient DNA data shows that the hunter-gatherers of Western Europe some eight thousand years ago had blue eyes but dark skin and dark hair, a combination that is rare today."
  40. ^ a b c Hanel, Andrea; Carlberg, Carsten (September 2020). "Skin colour and vitamin D: An update". Experimental Dermatology. 29 (9): 867. doi:10.1111/exd.14142. ISSN 0906-6705. PMID 32621306. S2CID 220335539. Homo sapiens arrrived in Europe from Near East some 42 000 years ago. Like in their African origin, these humans had dark skin but due to variations of their OCA2 gene (causing iris depigmentation) many of them had blue eyes" (...) "southern and central Europe, where they [light skin alleles] were introduced by farmers from western Anatolia expanding 8500 to 5000 years ago. This was the start of the Neolithic revolution in these regions, characterized by a more sedentary lifestyle and the domestication of certain animal and plant species. (...) "The rapid increase in population size due to the Neolithic revolution, such as the use of milk products as food source for adults and the rise of agriculture, as well as the massive spread of Yamnaya pastoralists likely caused the rapid selective sweep in European populations towards light skin and hair
  41. ^ Warren, Graeme (2021). Hunter-Gatherer Ireland: making connections in an island world. Oxbow Books. ISBN 978-1789256840. "WHGs for example, had skin pigmentation ranging from olive to brown to black, with blue or blue-green eyes. In some parts of Europe this may also have been associated with blond hair."
  42. ^ Population 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
  43. ^ Brace, Selina; Diekmann, Yoan; Booth, Thomas J.; Faltyskova, Zuzana; Rohland, Nadin; Mallick, Swapan; Ferry, Matthew; Michel, Megan; Oppenheimer, Jonas; Broomandkhoshbacht, Nasreen; Stewardson, Kristin; Walsh, Susan; Kayser, Manfred; Schulting, Rick; Craig, Oliver E.; Sheridan, Alison; Pearson, Mike Parker; Stringer, Chris; Reich, David; Thomas, Mark G.; Barnes, Ian (2019), "Population Replacement in Early Neolithic Britain", Nature Ecology & Evolution, 3 (5): 765–771, Bibcode:2019NatEE...3..765B, doi:10.1038/s41559-019-0871-9, PMC 6520225, PMID 30988490 Supplementary Material. Page 22: "Two WHGs (Cheddar Man and La Braña from northern Spain) are predicted to have had dark or dark-to-black skin, whereas one (Loschbour44from Luxembourg) is predicted to have had intermediate skin suggesting but we find potential temporal and/or geographical variation in pigmentation characteristics, suggesting that diverse skin pigmentation levels coexisted in WHGs by at least ca.8 kBP. Sven was predicted to have had dark to intermediate to dark skin in line with the current hypothesis that alleles commonly associated with lighter skin in Europeans were introduced to north-western Europe by ANFs."
  44. ^ Walsh, Susan (2017). "Global skin colour prediction from DNA". Human Genetics. 136 (7): 847–863. doi:10.1007/s00439-017-1808-5. PMC 5487854. PMID 28500464.
  45. ^ "Dark Skin, Blue Eyes: Genes Paint 7,000-Year-Old European's Picture". NBC News. 26 January 2014.
  46. ^ Callaway, Ewen (12 May 2022). "Ancient DNA maps 'dawn of farming'". Nature. doi:10.1038/d41586-022-01322-w. PMID 35552521. S2CID 248765487. Once established in Anatolia, Excoffier's team found, early farming populations moved west into Europe in a stepping-stone-like fashion, beginning around 8,000 years ago. They mixed occasionally — but not extensively — with local hunter-gatherers.
  47. ^ Quillen, Ellen (2019). "Shades of complexity: New perspectives on the evolution and genetic architecture of human skin". American Journal of Biological Anthropology. 168 (S67): 4–26. doi:10.1002/ajpa.23737. PMID 30408154. S2CID 53237190. Their analyses suggest that the skin color of both individuals was likely dark, with that of Mesolithic Cheddar Man predicted to be "dark or dark to black". These findings suggest that lighter skin color was uncommon across much of Europe during the Mesolithic. This is not, however, in conflict with the date estimates of <20 kya above, which addresses the onset of selection and not time of fixation of favored alleles (Beleza et al., 2013; Beleza, Johnson, et al., 2013). While ancient genome studies predict generally darker skin color among Mesolithic Europeans, derived alleles at rs1426654 and rs16891982 were segregating in European populations during the Mesolithic (González-Fortes et al., 2017; Günther et al., 2018; Mittnik et al., 2018), suggesting that phenotypic variation due to these loci was likely present by this time. However, reconstructions of Mesolithic and Neolithic pigmentation phenotype using loci common in modern populations should be interpreted with some caution, as it is possible that other as yet unexamined loci may have also influenced phenotype.
  48. ^ "Ancient 'dark skinned' Cheddar man find may not be true". New Scientist. 21 February 2018.
  49. ^ Krause, Johannes (2021). A Short History of Humanity A New History of Old Europe. l: Random House Publishing Group. ISBN 9780593229446.
  50. ^ Simões, Luciana G.; Peyroteo-Stjerna, Rita; Marchand, Grégor; Bernhardsson, Carolina; Vialet, Amélie; Chetty, Darshan; Alaçamlı, Erkin; Edlund, Hanna; Bouquin, Denis; Dina, Christian; Garmond, Nicolas; Günther, Torsten; Jakobsson, Mattias (5 March 2024). "Genomic ancestry and social dynamics of the last hunter-gatherers of Atlantic France". Proceedings of the National Academy of Sciences. 121 (10): e2310545121. Bibcode:2024PNAS..12110545S. doi:10.1073/pnas.2310545121. ISSN 0027-8424. PMC 10927518. PMID 38408241.

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Further reading edit

April 08, 2024
western, hunter, gatherer, archaeogenetics, term, west, european, hunter, gatherer, western, european, hunter, gatherer, villabruna, cluster, oberkassel, cluster, name, given, distinct, ancestral, component, modern, europeans, representing, descent, from, popu. In archaeogenetics the term Western Hunter Gatherer WHG West European Hunter Gatherer Western European Hunter Gatherer Villabruna cluster or Oberkassel cluster c 15 000 5 000 BP is the name given to a distinct ancestral component of modern Europeans representing descent from a population of Mesolithic hunter gatherers who scattered over Western Southern and Central Europe from the British Isles in the west to the Carpathians in the east following the retreat of the ice sheet of the Last Glacial Maximum 2 Western Hunter GathererGenetic ancestry of hunter gatherers in Europe between 14 ka and 9 ka with the main area of Western Hunter Gatherers WHG in blue Individual numbers correspond to calibrated sample dates 1 Along with the Scandinavian Hunter Gatherers SHG and Eastern Hunter Gatherers EHG the WHGs constituted one of the three main genetic groups in the postglacial period of early Holocene Europe 3 The border between WHGs and EHGs ran roughly from the lower Danube northward along the western forests of the Dnieper towards the western Baltic Sea 2 SHGs were in turn a nearly equal mix of WHGs and EHGs Once the main population throughout Europe the WHGs were largely displaced by successive expansions of Early European Farmers EEFs during the early Neolithic but experienced a resurgence during the Middle Neolithic During the Late Neolithic and Early Bronze Age Western Steppe Herders WSHs from the Pontic Caspian steppe embarked on a massive expansion which further displaced the WHGs Among modern day populations WHG ancestry is most common among populations of the eastern Baltic 4 Contents 1 Research 1 1 Expansion into continental Europe 14 000 BP 1 2 Interaction with other populations 1 2 1 Replacement by Neolithic farmers 2 Physical appearance 3 Notes 4 References 5 Bibliography 6 Further readingResearch editWestern Hunter Gatherers WHG are recognised as a distinct ancestral component contributing to the ancestry of most modern Europeans 5 Most Europeans can be modeled as a mixture of WHG EEF and WSH from the Pontic Caspian steppe 6 WHGs also contributed ancestry to other ancient groups such as Early European Farmers EEF who were however mostly of Anatolian descent 5 With the Neolithic expansion EEF came to dominate the gene pool in most parts of Europe although WHG ancestry had a resurgence in Western Europe from the Early Neolithic to the Middle Neolithic 7 Expansion into continental Europe 14 000 BP edit WHGs themselves are believed to have formed around 14 000 years ago during the Bolling Allerod interstadial at the time of the first major warming period after the Ice Age They represent a major population shift within Europe at the end of the Ice Age probably a population expansion into continental Europe from Southeastern European or West Asian refugia 8 It is thought that their ancestors separated from eastern Eurasians around 40 000 BP and from Ancient North Eurasians ANE prior to 24 000 BP the estimated age date of the Mal ta boy This date was subsequently put further back in time by the findings of the Yana Rhinoceros Horn Site to around 38kya shortly after the divergence of West Eurasian and East Eurasian lineages 5 9 Vallini et al 2022 argues that the dispersal and split patterns of West Eurasian lineages was not earlier than c 38 000 years ago with older Initial Upper Paleolithic European specimens such as Zlaty Kun Peștera cu Oase and Bacho Kiro being unrelated to Western Hunter gatherers but closer to Ancient East Eurasians or basal to both 10 The WHG displayed higher affinity for ancient and modern Middle Eastern populations when compared against earlier Paleolithic Europeans such as Gravettians The affinity for ancient Middle Eastern populations in Europe increased after the Last Glacial Maximum correlating with the expansion of WHG Villabruna or Oberkassel ancestry There is also evidence for bi directional geneflow between WHG and Middle Eastern populations as early as 15 000 years ago WHG associated remains belonged primarily to the human Y chromosome haplogroups I M170 with a lower frequency of C F3393 specifically the clade C V20 C1a2 which has been found commonly among earlier Paleolithic European remains such as Kostenki 14 and Sungir The paternal haplogroup C V20 can still be found in men living in modern Spain attesting to this lineage s longstanding presence in Western Europe Their mitochondrial chromosomes belonged primarily to haplogroup U5 11 12 nbsp Last Glacial Maximum refugia c 20 000 years ago Solutrean culture Epigravettian culture 11 In a genetic study published in Nature in March 2023 the authors found that the ancestors of the WHGs were populations associated with the Epigravettian culture which largely replaced populations associated with the Magdalenian culture about 14 000 years ago the ancestors of the Magdalenian associated individuals were the populations associated with the western Gravettian Solutrean and Aurignacian cultures 11 13 In the study WHG ancestry is renamed Oberkassel ancestry first found north of the Alps in two 14 000 year old individuals at the eponymous site at Oberkassel who can be modeled as an admixture of Villabruna ancestry itself modeled as an admixture between a lineage related to the Vestonice cluster and a lineage ancestral to the Kostenki 14 and Goyet Q116 1 individuals and Goyet Q2 ancestry related to individuals found in Europe prior to the Last Glacial Maximum The study states that all of the individuals of the Oberkassel cluster could be modeled as c 75 Villabruna and 25 Goyet Q2 ancestry or alternatively as c 90 Villabruna and 10 Fournol ancestry a newly identified cluster described as a sister lineage of the Goyet Q116 1 ancestry found in individuals associated with the Gravettian culture of southwestern Europe 11 The study suggests that Oberkassel ancestry was mostly already formed before expanding possibly around the west side of the Alps to Western and Central Europe and Britain where sampled WHG individuals are genetically homogeneous This is in contrast to the arrival of Villabruna and Oberkassel ancestry to Iberia which seems to have involved repeated admixture events with local populations carrying high levels of Goyet Q2 ancestry This and the survival of specific Y DNA haplogroup C1 clades previously observed among early European hunter gatherers suggests relatively higher genetic continuity in southwest Europe during this period 11 There are indications that the WHG carried risk alleles for diabetes and Alzheimer s disease 14 nbsp Transition from Magdalenian Goyet ancestry green Goyet Q2 to Western Hunter Gatherer WHG Villabruna ancestry orange in European sites according to timeline and climate evolution 15 Interaction with other populations edit nbsp Cheddar Man found in Great Britain had a similar genotype to other Western Hunter Gatherers The WHG were also found to have contributed ancestry to populations on the borders of Europe such as early Anatolian farmers and Ancient Northwestern Africans 16 as well as other European groups such as Eastern Hunter Gatherers 17 The relationship of WHGs to the EHGs remains inconclusive 17 EHGs are modeled to derive varying degrees of ancestry from a WHG related lineage ranging from merely 25 to up to 91 with the remainder being linked to geneflow from Paleolithic Siberians ANE and perhaps Caucasus hunter gatherers Another lineage known as the Scandinavian Hunter Gatherers SHGs were found to be a mix of EHGs and WHGs a 19 20 People of the Mesolithic Kunda culture and the Narva culture of the eastern Baltic were a mix of WHG and EHG 21 showing the closest affinity with WHG Samples from the Ukrainian Mesolithic and Neolithic were found to cluster tightly together between WHG and EHG suggesting genetic continuity in the Dnieper Rapids for a period of 4 000 years The Ukrainian samples belonged exclusively to the maternal haplogroup U which is found in around 80 of all European hunter gatherer samples 22 People of the Pit Comb Ware culture CCC of the eastern Baltic were closely related to EHG 23 Unlike most WHGs the WHGs of the eastern Baltic did not receive European farmer admixture during the Neolithic Modern populations of the eastern Baltic thus harbor a larger amount of WHG ancestry than any other population in Europe 21 SHGs have been found to contain a mix of WHG components who had likely migrated into Scandinavia from the south and EHGs who had later migrated into Scandinavia from the northeast along the Norwegian coast This hypothesis is supported by evidence that SHGs from western and northern Scandinavia had less WHG ancestry ca 51 than individuals from eastern Scandinavia ca 62 The WHGs who entered Scandinavia are believed to have belonged to the Ahrensburg culture EHGs and WHGs displayed lower allele frequencies of SLC45A2 and SLC24A5 which cause depigmentation and OCA Herc2 which causes light eye color than SHGs 24 nbsp The rock shelter where the skeleton of the Loschbour man c 8 000 BP was foundThe DNA of eleven WHGs from the Upper Palaeolithic and Mesolithic in Western Europe Central Europe and the Balkans was analyzed with regards to their Y DNA haplogroups and mtDNA haplogroups The analysis suggested that WHGs were once widely distributed from the Atlantic coast in the West to Sicily in the South to the Balkans in the Southeast for more than six thousand years 25 The study also included an analysis of a large number of individuals of prehistoric Eastern Europe Thirty seven samples were collected from Mesolithic and Neolithic Ukraine 9500 6000 BC These were determined to be an intermediate between EHG and SHG although WHG ancestry in this population increased during the Neolithic Samples of Y DNA extracted from these individuals belonged exclusively to R haplotypes particularly subclades of R1b1 and I haplotypes particularly subclades of I2 mtDNA belonged almost exclusively to U particularly subclades of U5 and U4 25 A large number of individuals from the Zvejnieki 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 of haplogroup R1b1a1a and I2a1 The mtDNA belonged exclusively to haplogroup U particularly subclades of U2 U4 and U5 25 Forty individuals from three sites of the Iron Gates Mesolithic in the Balkans were also analyzed These individuals were estimated to be of 85 WHG and 15 EHG descent The males at these sites carried exclusively haplogroup R1b1a and I mostly subclades of I2a haplotypes mtDNA belonged mostly to U particularly subclades of U5 and U4 25 People of the Balkan Neolithic were found to harbor 98 Anatolian ancestry and 2 WHG ancestry By the Chalcolithic people of the Cucuteni Trypillia culture were found to harbor about 20 hunter gatherer ancestry which was intermediate between EHG and WHG People of the Globular Amphora culture were found to harbor ca 25 WHG ancestry which is significantly higher than Middle Neolithic groups of Central Europe 25 Replacement by Neolithic farmers edit Further information Neolithic Europe and Genetic history of Europe nbsp Residual genetic ancestry of European hunter gatherers during the European Neolithic between 7 5 ka and 5 ka BP c 5 500 3 000 BC nbsp Simplified model for the demographic history of Europeans during the Neolithic period and the Neolithic Revolution which introduced agriculture 26 A seminal 2014 study first identified the contribution of three main components to modern European lineages the Western Hunter Gatherers WHG in proportions of up to 50 in Northern Europeans the Ancient North Eurasians ANE Upper Palaeolithic Siberians later associated with the later Indo European expansion present in proportions up to 20 and finally the Early European Farmers EEF agriculturists of mainly Near Eastern origin who migrated to Europe from circa 8 000 BP now present in proportions from around 30 in the Baltic region to around 90 in the Mediterranean The Early European Farmer EEF component was identified based on the genome of a woman buried c 7 000 years ago in a Linear Pottery culture grave in Stuttgart Germany 27 This 2014 study found evidence for genetic mixing between WHG and EEF throughout Europe with the largest contribution of EEF in Mediterranean Europe especially in Sardinia Sicily Malta and among Ashkenazi Jews and the largest contribution of WHG in Northern Europe and among Basque people 28 Since 2014 further studies have refined the picture of interbreeding between EEF and WHG In a 2017 analysis of 180 ancient DNA datasets of the Chalcolithic and Neolithic periods from Hungary Germany and Spain evidence was found of a prolonged period of interbreeding Admixture took place regionally from local hunter gatherer populations so that populations from the three regions Germany Iberia and Hungary were genetically distinguishable at all stages of the Neolithic period with a gradually increasing ratio of WHG ancestry of farming populations over time This suggests that after the initial expansion of early farmers there were no further long range migrations substantial enough to homogenize the farming population and that farming and hunter gatherer populations existed side by side for many centuries with ongoing gradual admixture throughout the 5th to 4th millennia BC rather than a single admixture event on initial contact 29 Admixture rates varied geographically in the late Neolithic WHG ancestry in farmers in Hungary was at around 10 in Germany around 25 and in Iberia as high as 50 30 Analysis of remains from the Grotta Continenza in Italy showed that out of six remains three buried between c 10 000 BC and 7000 BC belonged to I2a P214 and two times the maternal haplogroups U5b1 and one U5b3 31 32 Around 6000 BC the WHGs of Italy were almost completely genetically replaced by EEFs two G2a2 and one Haplogroup R1b although WHG ancestry slightly increased in subsequent millennia 33 Neolithic individuals in the British Isles were close to Iberian and Central European Early and Middle Neolithic populations modeled as having about 75 ancestry from EEF with the rest coming from WHG in continental Europe They subsequently replaced most of the WHG population in the British Isles without mixing much with them 34 The WHG are estimated to have contributed between 20 30 ancestry to Neolithic EEF groups throughout Europe Specific adaptions against local pathogens may have been introduced via the Mesolithic WHG admixture into Neolithic EEF populations 35 A study on Mesolithic hunter gatherers from Denmark found that they were related to contemporary Western hunter gatherers and are associated with the Maglemose Kongemose and Ertebolle cultures They displayed genetic homogeneity from around 10 500 to 5 900 calibrated years before present until Neolithic farmers with Anatolian derived ancestry arrived The transition to the Neolithic period was very abrupt and resulted in a population turnover with limited genetic contribution from local hunter gatherers The succeeding Neolithic population has been associated with the Funnelbeaker culture 36 Physical appearance edit nbsp Reconstructed head of the Cheddar Man found in England carbon dated c 8540 8230 BC based on the shape of his skull and DNA analysis shown at the National History Museum in London 2019 37 38 According to David Reich DNA analysis has shown that Western Hunter Gatherers were typically dark skinned dark haired and blue eyed 39 The dark skin was due to their Out of Africa origin all Homo sapiens populations having had initially dark skin while the blue eyes were the result of a variation in their OCA2 gene which caused iris depigmentation 40 Archaeologist Graeme Warren has said that their skin color ranged from olive to black and speculated that they may have had some regional variety of eye and hair colors 41 This is strikingly different from the distantly related Eastern Hunter Gatherers EHG who have been suggested to be light skinned brown eyed or blue eyed and dark haired or light haired 42 Two WHG skeletons with incomplete SNPs La Brana and Cheddar Man are predicted to have had dark or dark to black skin whereas two other WHG skeletons with complete SNPs Sven and Loschbour man are predicted to have had dark or intermediate to dark and intermediate skin respectively 43 24 b Spanish biologist Carles Lalueza Fox said the La Brana 1 individual had dark skin although we cannot know the exact shade 45 According to a 2020 study the arrival of Early European Farmers EEFs from western Anatolia from 8500 to 5000 years ago along with Western Steppe Herders during the Bronze Age caused a rapid evolution of European populations towards lighter skin and hair 40 Admixture between hunter gatherer and agriculturist populations was apparently occasional but not extensive 46 nbsp Evolution of Upper Paleolithic and Neolithic phenotypes in Eurasia Dark skinned Western Hunter Gatherers resided in Western Europe and expanded to some extent towards north and eastern Europe 40 Some authors have expressed caution regarding skin pigmentation reconstructions Quillen et al 2019 acknowledge studies that generally show that lighter skin color was uncommon across much of Europe during the Mesolithic including studies regarding the dark or dark to black predictions for the Cheddar Man but warn that reconstructions of Mesolithic and Neolithic pigmentation phenotype using loci common in modern populations should be interpreted with some caution as it is possible that other as yet unexamined loci may have also influenced phenotype 47 Geneticist Susan Walsh at Indiana University Purdue University Indianapolis who worked on Cheddar Man project said that we simply don t know his skin colour 48 German biochemist Johannes Krause stated that we do not know whether the skin color of Western European hunter gatherers was more similar to the skin color of people from present day Central Africa or people from the Arab region It is only certain that they did not carry any known mutation responsible for the light skin in subsequent populations of Europeans 49 A 2024 research into the genomic ancestry and social dynamics of the last hunter gatherers of Atlantic France has stated that phenotypically we find some diversity during the Late Mesolithic in France at which two of the WHG s sequenced in the study likely had pale to intermediate skin pigmentation but most individuals carry the dark skin and blue eyes characteristic of WHGs of the studied samples 50 Notes edit 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 18 These predictions were obtained using a multinomial logistic regression model based on a panel of 36 carefully selected SNPs with a low sensitivity of 0 26 for classifying intermediate skin compared to 0 99 and 0 90 for white and black skin respectively The accuracy of the model used could be further improved with additional but currently unknown SNP predictors once identified via future GWAS 44 References edit Posth 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 ISSN 1476 4687 PMC 9977688 PMID 36859578 S2CID 257259969 a b Anthony 2019b p 28 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 Davy Tom Ju Dan Mathieson Iain Skoglund Pontus April 2023 Hunter gatherer admixture facilitated natural selection in Neolithic European farmers Current Biology 33 7 1365 1371 e3 Bibcode 2023CBio 33E1365D doi 10 1016 j cub 2023 02 049 ISSN 0960 9822 PMC 10153476 PMID 36963383 a b c Lazaridis 2014 Mathieson 2015 Haak 2015 Fu Qiaomei 2016 The genetic history of Ice Age Europe Nature 534 7606 200 205 Bibcode 2016Natur 534 200F doi 10 1038 nature17993 PMC 4943878 PMID 27135931 Beginning with the Villabruna Cluster at least 14 000 years ago all European individuals analyzed show an affinity to the Near East This correlates in time to the Bolling Allerod interstadial the first significant warming period after the Ice Age Archaeologically it correlates with cultural transitions within the Epigravettian in Southern Europe and the Magdalenian to Azilian transition in Western Europe Thus the appearance of the Villabruna Cluster may reflect migrations or population shifts within Europe at the end of the Ice Age an observation that is also consistent with the evidence of turnover of mitochondrial DNA sequences at this time One scenario that could explain these patterns is a population expansion from southeastern European or west Asian refugia after the Ice Age drawing together the genetic ancestry of Europe and the Near East Sixth within the Villabruna Cluster some but not all individuals have affinity to East Asians An important direction for future work is to generate similar ancient DNA data from southeastern Europe and the Near East to arrive at a more complete picture of the Upper Paleolithic population history of western Eurasia Sikora Martin Pitulko Vladimir V Sousa Vitor C Allentoft Morten E Vinner Lasse Rasmussen Simon Margaryan Ashot de Barros Damgaard Peter de la Fuente Constanza Renaud Gabriel Yang Melinda A Fu Qiaomei Dupanloup Isabelle Giampoudakis Konstantinos Nogues Bravo David June 2019 The population history of northeastern Siberia since the Pleistocene Nature 570 7760 182 188 Bibcode 2019Natur 570 182S doi 10 1038 s41586 019 1279 z hdl 1887 3198847 ISSN 1476 4687 PMID 31168093 S2CID 174809069 Vallini et al 2022 4 July 2022 Genetics and Material Culture Support Repeated Expansions into Paleolithic Eurasia from a Population Hub Out of Africa Retrieved 16 April 2023 a href Template Cite web html title Template Cite web cite web a CS1 maint numeric names authors list link a b c d e Posth C Yu H Ghalichi A 2023 Palaeogenomics of Upper Palaeolithic to Neolithic European hunter gatherers Nature 615 2 March 2023 117 126 Bibcode 2023Natur 615 117P doi 10 1038 s41586 023 05726 0 PMC 9977688 PMID 36859578 a href Template Cite journal html title Template Cite journal cite journal a CS1 maint multiple names authors list link Scozzari Rosaria Massaia Andrea D Atanasio Eugenia Myres Natalie M Perego Ugo A Trombetta Beniamino Cruciani Fulvio 7 November 2012 Molecular Dissection of the Basal Clades in the Human Y Chromosome Phylogenetic Tree PLOS ONE 7 11 e49170 Bibcode 2012PLoSO 749170S doi 10 1371 journal pone 0049170 ISSN 1932 6203 PMC 3492319 PMID 23145109 Through this analysis we identified a chromosome from southern Europe as a new deep branch within haplogroup C C V20 or C7 Figure S1 Previously only a few examples of C chromosomes only defined by the marker RPS4Y711 had been found in southern Europe 32 33 To improve our knowledge regarding the distribution of haplogroup C in Europe we surveyed 1965 European subjects for the mutation RPS4Y711 and identified one additional haplogroup C chromosome from southern Europe which has also been classified as C7 data not shown Scientists Sequence Genomes of Prehistoric Hunter Gatherers from Different Eurasian Cultures Sci News 2 March 2023 Irving Pease Evan K Refoyo Martinez Alba Barrie William Ingason Andres Pearson Alice Fischer Anders Sjogren Karl Goran Halgren Alma S Macleod Ruairidh Demeter Fabrice Henriksen Rasmus A Vimala Tharsika McColl Hugh Vaughn Andrew H Speidel Leo January 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 ISSN 1476 4687 PMC 10781624 PMID 38200293 whereas risk alleles for diabetes and Alzheimer s disease are enriched for Western hunter gatherer ancestry Charlton Sophy Brace Selina November 2022 Dual ancestries and ecologies of the Late Glacial Palaeolithic in Britain Nature Ecology amp Evolution 6 11 1658 1668 Bibcode 2022NatEE 6 1658C doi 10 1038 s41559 022 01883 z ISSN 2397 334X PMC 9630104 PMID 36280785 Simoes Luciana G Gunther Torsten Martinez Sanchez Rafael M Vera Rodriguez Juan Carlos Iriarte Eneko Rodriguez Varela Ricardo Bokbot Youssef Valdiosera Cristina Jakobsson Mattias 7 June 2023 Northwest African Neolithic initiated by migrants from Iberia and Levant Nature 618 7965 550 556 Bibcode 2023Natur 618 550S doi 10 1038 s41586 023 06166 6 ISSN 1476 4687 PMC 10266975 PMID 37286608 a b Lazaridis Iosif 1 December 2018 The evolutionary history of human populations in Europe Current Opinion in Genetics amp Development Genetics of Human Origins 53 21 27 arXiv 1805 01579 doi 10 1016 j gde 2018 06 007 ISSN 0959 437X PMID 29960127 S2CID 19158377 Lazaridis 2016 Haak Wolfgang Lazaridis Iosif Patterson Nick Rohland Nadin Mallick Swapan Llamas Bastien Brandt Guido Nordenfelt Susanne Harney Eadaoin Stewardson Kristin Fu Qiaomei Mittnik Alissa Banffy 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 ISSN 1476 4687 PMC 5048219 PMID 25731166 Sikora Martin Pitulko Vladimir V Sousa Vitor C Allentoft Morten E Vinner Lasse Rasmussen Simon Margaryan Ashot de Barros Damgaard Peter de la Fuente Constanza Renaud Gabriel Yang Melinda A Fu Qiaomei Dupanloup Isabelle Giampoudakis Konstantinos Nogues Bravo David June 2019 The population history of northeastern Siberia since the Pleistocene Nature 570 7760 182 188 Bibcode 2019Natur 570 182S doi 10 1038 s41586 019 1279 z hdl 1887 3198847 ISSN 1476 4687 PMID 31168093 S2CID 174809069 a b Mittnik 2018 Jones 2017 Saag 2017 a b Gunther 2018 a b c d e Mathieson 2018 Sikora M Carpenter ML Moreno Estrada A Henn BM Underhill PA Sanchez Quinto F et al May 2014 Population genomic analysis of ancient and modern genomes yields new insights into the genetic ancestry of the Tyrolean Iceman and the genetic structure of Europe PLOS Genetics 10 5 e1004353 doi 10 1371 journal pgen 1004353 PMC 4014435 PMID 24809476 Lazaridis Iosif Patterson Nick Mittnik Alissa September 2014 Ancient human genomes suggest three ancestral populations for present day Europeans Nature 513 7518 409 413 arXiv 1312 6639 Bibcode 2014Natur 513 409L doi 10 1038 nature13673 ISSN 1476 4687 PMC 4170574 PMID 25230663 Most present day Europeans derive from at least three highly differentiated populations west European hunter gatherers who contributed ancestry to all Europeans but not to Near Easterners ancient north Eurasians related to Upper Palaeolithic Siberians who contributed to both Europeans and Near Easterners and early European farmers who were mainly of Near Eastern origin but also harboured west European hunter gatherer related ancestry Lazaridis et al 2014 Supplementary Information p 113 Lipson et al Parallel palaeogenomic transects reveal complex genetic history of early European farmers Nature 551 368 372 16 November 2017 doi 10 1038 nature24476 Lipson et al 2017 Fig 2 Antonio et al 2019 Table 2 Sample Information Rows 4 6 Antonio et al 2019 p 1 Antonio et al 2019 p 2 Fig 1 Brace Selina Diekmann Yoan Booth Thomas J van Dorp Lucy Faltyskova Zuzana Rohland Nadin Mallick Swapan Olalde Inigo Ferry Matthew Michel Megan Oppenheimer Jonas Broomandkhoshbacht Nasreen Stewardson Kristin Martiniano Rui Walsh Susan Kayser Manfred Charlton Sophy Hellenthal Garrett Armit Ian Schulting Rick Craig Oliver E Sheridan Alison Parker Pearson Mike Stringer Chris Reich David Thomas Mark G Barnes Ian 2019 Ancient genomes indicate population replacement in Early Neolithic Britain Nature Ecology amp Evolution 3 5 765 771 Bibcode 2019NatEE 3 765B doi 10 1038 s41559 019 0871 9 ISSN 2397 334X PMC 6520225 PMID 30988490 Davy Tom Ju Dan Mathieson Iain Skoglund Pontus April 2023 Hunter gatherer admixture facilitated natural selection in Neolithic European farmers Current Biology 33 7 1365 1371 e3 Bibcode 2023CBio 33E1365D doi 10 1016 j cub 2023 02 049 ISSN 0960 9822 PMC 10153476 PMID 36963383 Allentoft Morten E Sikora Martin Fischer Anders Sjogren Karl Goran Ingason Andres Macleod Ruairidh Rosengren Anders Schulz Paulsson Bettina Jorkov Marie Louise Schjellerup Novosolov Maria Stenderup Jesper Price T Douglas Fischer Mortensen Morten Nielsen Anne Birgitte Ulfeldt Hede Mikkel 10 January 2024 100 ancient genomes show repeated population turnovers in Neolithic Denmark Nature 625 7994 329 337 Bibcode 2024Natur 625 329A doi 10 1038 s41586 023 06862 3 ISSN 1476 4687 PMC 10781617 PMID 38200294 Conneller Chantal 29 November 2021 The Mesolithic in Britain Landscape and Society in Times of Change Routledge p 126 ISBN 978 1 000 47515 9 Details on the reconstruction from the Natural History Museum Cheddar Man Mesolithic Britain s blue eyed boy www nhm ac uk Natural History Museum Reich David 2018 Who We Are and How We Hot Here Ancient DNA and the New Science of the Human Past First ed New York Knopf Doubleday Publishing Group ISBN 978 1101870334 Analysis of ancient DNA data shows that the hunter gatherers of Western Europe some eight thousand years ago had blue eyes but dark skin and dark hair a combination that is rare today a b c Hanel Andrea Carlberg Carsten September 2020 Skin colour and vitamin D An update Experimental Dermatology 29 9 867 doi 10 1111 exd 14142 ISSN 0906 6705 PMID 32621306 S2CID 220335539 Homo sapiens arrrived in Europe from Near East some 42 000 years ago Like in their African origin these humans had dark skin but due to variations of their OCA2 gene causing iris depigmentation many of them had blue eyes southern and central Europe where they light skin alleles were introduced by farmers from western Anatolia expanding 8500 to 5000 years ago This was the start of the Neolithic revolution in these regions characterized by a more sedentary lifestyle and the domestication of certain animal and plant species The rapid increase in population size due to the Neolithic revolution such as the use of milk products as food source for adults and the rise of agriculture as well as the massive spread of Yamnaya pastoralists likely caused the rapid selective sweep in European populations towards light skin and hair Warren Graeme 2021 Hunter Gatherer Ireland making connections in an island world Oxbow Books ISBN 978 1789256840 WHGs for example had skin pigmentation ranging from olive to brown to black with blue or blue green eyes In some parts of Europe this may also have been associated with blond hair Population 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 Brace Selina Diekmann Yoan Booth Thomas J Faltyskova Zuzana Rohland Nadin Mallick Swapan Ferry Matthew Michel Megan Oppenheimer Jonas Broomandkhoshbacht Nasreen Stewardson Kristin Walsh Susan Kayser Manfred Schulting Rick Craig Oliver E Sheridan Alison Pearson Mike Parker Stringer Chris Reich David Thomas Mark G Barnes Ian 2019 Population Replacement in Early Neolithic Britain Nature Ecology amp Evolution 3 5 765 771 Bibcode 2019NatEE 3 765B doi 10 1038 s41559 019 0871 9 PMC 6520225 PMID 30988490 Supplementary Material Page 22 Two WHGs Cheddar Man and La Brana from northern Spain are predicted to have had dark or dark to black skin whereas one Loschbour44from Luxembourg is predicted to have had intermediate skin suggesting but we find potential temporal and or geographical variation in pigmentation characteristics suggesting that diverse skin pigmentation levels coexisted in WHGs by at least ca 8 kBP Sven was predicted to have had dark to intermediate to dark skin in line with the current hypothesis that alleles commonly associated with lighter skin in Europeans were introduced to north western Europe by ANFs Walsh Susan 2017 Global skin colour prediction from DNA Human Genetics 136 7 847 863 doi 10 1007 s00439 017 1808 5 PMC 5487854 PMID 28500464 Dark Skin Blue Eyes Genes Paint 7 000 Year Old European s Picture NBC News 26 January 2014 Callaway Ewen 12 May 2022 Ancient DNA maps dawn of farming Nature doi 10 1038 d41586 022 01322 w PMID 35552521 S2CID 248765487 Once established in Anatolia Excoffier s team found early farming populations moved west into Europe in a stepping stone like fashion beginning around 8 000 years ago They mixed occasionally but not extensively with local hunter gatherers Quillen Ellen 2019 Shades of complexity New perspectives on the evolution and genetic architecture of human skin American Journal of Biological Anthropology 168 S67 4 26 doi 10 1002 ajpa 23737 PMID 30408154 S2CID 53237190 Their analyses suggest that the skin color of both individuals was likely dark with that of Mesolithic Cheddar Man predicted to be dark or dark to black These findings suggest that lighter skin color was uncommon across much of Europe during the Mesolithic This is not however in conflict with the date estimates of lt 20 kya above which addresses the onset of selection and not time of fixation of favored alleles Beleza et al 2013 Beleza Johnson et al 2013 While ancient genome studies predict generally darker skin color among Mesolithic Europeans derived alleles at rs1426654 and rs16891982 were segregating in European populations during the Mesolithic Gonzalez Fortes et al 2017 Gunther et al 2018 Mittnik et al 2018 suggesting that phenotypic variation due to these loci was likely present by this time However reconstructions of Mesolithic and Neolithic pigmentation phenotype using loci common in modern populations should be interpreted with some caution as it is possible that other as yet unexamined loci may have also influenced phenotype Ancient dark skinned Cheddar man find may not be true New Scientist 21 February 2018 Krause Johannes 2021 A Short History of Humanity A New History of Old Europe l Random House Publishing Group ISBN 9780593229446 Simoes Luciana G Peyroteo Stjerna Rita Marchand Gregor Bernhardsson Carolina Vialet Amelie Chetty Darshan Alacamli Erkin Edlund Hanna Bouquin Denis Dina Christian Garmond Nicolas Gunther Torsten Jakobsson Mattias 5 March 2024 Genomic ancestry and social dynamics of the last hunter gatherers of Atlantic France Proceedings of the National Academy of Sciences 121 10 e2310545121 Bibcode 2024PNAS 12110545S doi 10 1073 pnas 2310545121 ISSN 0027 8424 PMC 10927518 PMID 38408241 Bibliography editAnthony David W 2019b Ancient DNA Mating Networks and the Anatolian Split In Serangeli Matilde Olander Thomas eds Dispersals and Diversification Linguistic and Archaeological Perspectives on the Early Stages of Indo European BRILL pp 21 54 ISBN 978 9004416192 Antonio Margaret L et al 8 November 2019 Ancient Rome A genetic crossroads of Europe and the Mediterranean Science 366 6466 American Association for the Advancement of Science 708 714 Bibcode 2019Sci 366 708A doi 10 1126 science aay6826 PMC 7093155 PMID 31699931 Gunther Thorsten 1 January 2018 Population genomics of Mesolithic Scandinavia Investigating early postglacial migration routes and high latitude adaptation PLOS Biology 16 1 PLOS e2003703 doi 10 1371 journal pbio 2003703 PMC 5760011 PMID 29315301 Haak Wolfgang 11 June 2015 Massive migration from the steppe was a source for Indo European languages in Europe Nature 522 7555 Nature Research 207 211 arXiv 1502 02783 Bibcode 2015Natur 522 207H doi 10 1038 nature14317 PMC 5048219 PMID 25731166 Jones Eppie R 20 February 2017 The Neolithic Transition in the Baltic Was Not Driven by Admixture with Early European Farmers Current Biology 27 4 Cell Press 576 582 Bibcode 2017CBio 27 576J doi 10 1016 j cub 2016 12 060 PMC 5321670 PMID 28162894 Kashuba Natalija 15 May 2019 Ancient DNA from mastics solidifies connection between material culture and genetics of mesolithic hunter gatherers in Scandinavia Communications Biology 2 105 Nature Research 185 doi 10 1038 s42003 019 0399 1 PMC 6520363 PMID 31123709 Lazaridis Iosif 17 September 2014 Ancient human genomes suggest three ancestral populations for present day Europeans Nature 513 7518 Nature Research 409 413 arXiv 1312 6639 Bibcode 2014Natur 513 409L doi 10 1038 nature13673 hdl 11336 30563 PMC 4170574 PMID 25230663 Lazaridis Iosif 25 July 2016 Genomic insights into the origin of farming in the ancient Near East Nature 536 7617 Nature Research 419 424 Bibcode 2016Natur 536 419L doi 10 1038 nature19310 PMC 5003663 PMID 27459054 Mathieson Iain 23 November 2015 Genome wide patterns of selection in 230 ancient Eurasians Nature 528 7583 Nature Research 499 503 Bibcode 2015Natur 528 499M doi 10 1038 nature16152 PMC 4918750 PMID 26595274 Mathieson Iain 21 February 2018 The Genomic History of Southeastern Europe Nature 555 7695 Nature Research 197 203 Bibcode 2018Natur 555 197M doi 10 1038 nature25778 PMC 6091220 PMID 29466330 Mittnik Alisa 30 January 2018 The genetic prehistory of the Baltic Sea region Nature Communications 16 1 Nature Research 442 Bibcode 2018NatCo 9 442M doi 10 1038 s41467 018 02825 9 PMC 5789860 PMID 29382937 Saag Lehti 24 July 2017 Extensive Farming in Estonia Started through a Sex Biased Migration from the Steppe Current Biology 27 14 Cell Press 2185 2193 Bibcode 2017CBio 27E2185S doi 10 1016 j cub 2017 06 022 PMID 28712569 Further reading editAnthony David Spring Summer 2019 Archaeology Genetics and Language in the Steppes A Comment on Bomhard Journal of Indo European Studies 47 1 2 Retrieved 9 January 2020 Lazaridis Iosif December 2018 The evolutionary history of human populations in Europe Current Opinion in Genetics amp Development 53 Elsevier 21 27 arXiv 1805 01579 doi 10 1016 j gde 2018 06 007 PMID 29960127 S2CID 19158377 Retrieved 15 July 2020 Retrieved from https en wikipedia org w index php title Western Hunter Gatherer amp oldid 1217514040, wikipedia, wiki, book, books, library,

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