.png) 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] |
Inarchaeogenetics,western hunter-gatherer (WHG, also known aswest European hunter-gatherer,western European hunter-gatherer orOberkassel cluster) (c. 15,000~5,000 BP) is a distinct ancestral component of modern Europeans, representing descent from a population ofMesolithichunter-gatherers who scattered overwestern,southern andcentral Europe, from theBritish Isles in the west to theCarpathians in the east, following the retreat of theice sheet of theLast Glacial Maximum.[2] It is closely associated and sometimes considered synonymous with the concept of theVillabruna cluster, named after theRipari Villabruna cave specimen in Italy, known from the terminal Pleistocene of Europe, which is largely ancestral to later WHG populations.
WHGs share a closer genetic relationship to ancient and modern peoples in the Middle East and the Caucasus than earlier European hunter-gatherers. Their precise relationships to other groups are somewhat obscure, with the origin of the Villabruna cluster likely somewhere in the vicinity of theBalkans. The Villabruna cluster (which is associated with theEpigravettian and other related archaeological cultures) had expanded into the Italian and Iberian Peninsulas by approximately 19,000 years ago, with the WHG cluster subsequently expanding across Western Europe at the end of the Pleistocene around 14-12,000 years ago, largely replacing theMagdalenian peoples who previously dominated the region.[3] These Magdalenian peoples largely descended from earlier Western EuropeanCro-Magnon groups that had arrived in the region over 30,000 years ago, prior to the Last Glacial Maximum.
WHGs constituted one of the main genetic groups in the postglacial period of earlyHolocene Europe, along witheastern hunter-gatherers (EHG) in Eastern Europe. The border between WHGs and EHGs ran roughly from the lowerDanube, northward along the western forests of theDnieper towards the westernBaltic Sea.[2] EHGs primarily consisted of a mixture of WHG-related andAncient North Eurasian (ANE) ancestry.[3] Scandinavia was inhabited byScandinavian hunter-gatherers (SHGs), which were a mixture between WHG and EHG.[4] In the Iberian Peninsula, early Holocene hunter-gathers consisted of a mixture of WHG and Magdalenian Cro-Magnon (GoyetQ2) ancestry.[5]
Once the main population throughout Europe, the WHGs were largely replaced by successive expansions ofEarly European Farmers (EEFs) ofAnatolian origin during the earlyNeolithic, who generally carried a minor amount of WHG ancestry due toadmixture with WHG groups during their European expansion. Among modern-day populations, WHG ancestry is most common among populations of the eastern Baltic region.[6]
WHGs lacked the light skin genes found in modern Europeans, and it has been suggested they were dark skinned and had light coloured eyes.[7][3]
Western hunter-gatherers (WHG) are recognised as a distinct ancestral component contributing to the ancestry of most modernEuropeans.[8] Most Europeans can be modeled as a mixture of WHG, EEF, andWestern Steppe Herders (WSHs) from thePontic–Caspian steppe.[9] WHGs also contributed ancestry to other ancient groups such as Early European Farmers (EEF), who were, however, mostly of Anatolian descent.[8] With theNeolithic expansion, EEF came to dominate the gene pool in most parts of Europe, although WHG ancestry had a resurgence in Western Europe from the EarlyNeolithic to the Middle Neolithic.[10]
WHGs 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 Asianrefugia.[11] It is thought that their ancestors separated from eastern Eurasians around 40,000 BP, and fromAncient North Eurasians (ANE) prior to 24,000 BP (the estimated age date of theMal'ta boy). This date was subsequently put further back in time by the findings of theYana Rhinoceros Horn Site to around 38kya, shortly after the divergence of West-Eurasian and East-Eurasian lineages.[8][12] 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 olderInitial Upper Paleolithic European specimens, such as those found in theZlaty Kun,Peștera cu Oase andBacho Kiro caves, being unrelated to Western hunter-gatherers but closer toAncient East Eurasians or basal to both.[13]
The relationships of the WHG/Villabruna cluster to other Paleolithic human groups in Europe and West Asia are obscure and subject to conflicting interpretations.[3] A 2022 study proposed that the WHG/Villabruna population genetically diverged from hunter-gatherers in the Middle East and the Caucasus around 26,000 years ago, during theLast Glacial Maximum.[14] WHG genomes display higher affinity for ancient and modern Middle Eastern populations when compared against earlier Paleolithic Europeans such asGravettians. The affinity for ancient Middle Eastern populations in Europe increased after theLast 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 thehuman Y-chromosome haplogroupsI-M170 with a lower frequency ofC-F3393 (specifically the cladeC-V20/C1a2), which has been found commonly among earlier Paleolithic European remains such asKostenki-14 andSungir. The paternal haplogroup C-V20 can still be found in men living in modernSpain, attesting to this lineage's longstanding presence in Western Europe. The Villabruna cluster also carried theY-haplogroup R1b (R1b-L754), derived from the Ancient North Eurasianhaplogroup R*, indicating "an early link between Europe and the western edge of theSteppe Belt of Eurasia."[15] Theirmitochondrial chromosomes belonged primarily tohaplogroup U5.[3][16] A 2023 study proposed that the Villabruna cluster emerged from the mixing in roughly equal proportions of a divergent West Eurasian ancestry with a West Eurasian ancestry closely related to the 35,000 year old BK1653 individual from Bacho Kiro Cave in Bulgaria, with this BK1653-related ancestry also significantly (~59%) ancestral to the Věstonice cluster characteristic of easternGravettian producingCro-Magnon groups (which have additional ancestry precluding them from being the direct ancestors of the Villabruna cluster), which may reflect shared ancestry in the Balkan region.[3](see supplemental material)
Another genetic study models them as a mix of Upper Paleolithic hunter-gatherers from the Caucasus, Europe and Siberia.[17]
The earliest known individuals of predominantly WHG/Villabruna ancestry in Europe are known from Italy, dating to around 17,000 years ago,[18] though an individual fromEl Mirón cave in northern Spain with 43% Villabruna ancestry is known from 19,000 years ago. While not confirmed, the Villabruna cluster was probably present earlier than in the Balkans region than elsewhere in Southern Europe.[3] Early WHG/Villabruna populations are associated with theEpigravettian archaeological culture, which largely replaced populations associated with theMagdalenian culture about 14,000 years ago (the ancestry of Magdalenian-associated Goyet-Q2 cluster primarily descended from the earlierSolutrean, and western Gravettian-producing groups in France and Spain).[3][19] A 2023 study found that relative to earlier Western European Cro-Magnon populations like the Gravettians, that Magdalenian-associated Goyet-Q2 cluster carried significant (~30%) Villabruna ancestry even prior to the major expansion of WHG-related groups north of the Alps. This study also found that relative to earlier members of the Villabruna cluster from Italy, WHG-related groups which appeared north of the Alps beginning around 14,000 years ago carried around 25% ancestry from the Goyet-Q2 cluster (or alternatively 10% from the western Gravettian associated Fournol cluster). This paper proposed that WHG should be named theOberkassel cluster, after one of the oldest WHG individuals found north of the Alps.[3] 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.[3]
_to_Villabruna_(orange)_genetic_types_in_European_sites%2c_according_to_timeline_and_climate_evolution.jpg)
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,[21] as well as other European groups such aseastern hunter-gatherers.[22] The relationship of WHGs to the EHGs remains inconclusive.[22] 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 perhapsCaucasus hunter-gatherers. Another lineage known as theScandinavian hunter-gatherers (SHGs) were found to be a mix of EHGs and WHGs.[a][24][25]
In the Iberian Peninsula early Holocene hunter-gathers consisted of populations with a mixture of WHG and Magdalenian Cro-Magnon (GoyetQ2) ancestry.[5]
People of the MesolithicKunda culture and theNarva culture of the easternBaltic were a mix of WHG and EHG,[26] showing the closest affinity with WHG. Samples from theUkrainian Mesolithic andNeolithic were found to cluster tightly together between WHG and EHG, suggesting genetic continuity in theDnieper Rapids for 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.[27]
People of thePit–Comb Ware culture (CCC) of the eastern Baltic were closely related to EHG.[28] 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.[26]
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 theNorwegian 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 theAhrensburg culture. EHGs and WHGs displayed lower allele frequencies ofSLC45A2 andSLC24A5, which cause depigmentation, andOCA/Herc2, which causes lighteye color, than SHGs.[29]
The DNA of eleven WHGs from theUpper Palaeolithic andMesolithic inWestern Europe,Central Europe and theBalkans was analyzed, with regards to theirY-DNA haplogroups andmtDNA 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.[30] 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 toR haplotypes (particularly subclades ofR1b1) andI haplotypes (particularly subclades ofI2).mtDNA belonged almost exclusively toU (particularly subclades ofU5 andU4).[30] 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 R1b1a1a andI2a1. The mtDNA belonged exclusively tohaplogroup U (particularly subclades ofU2,U4 andU5).[30] Forty individuals from three sites of theIron Gates Mesolithic in theBalkans were also analyzed. These individuals were estimated to be of 85% WHG and 15% EHG descent. The males at these sites carried exclusivelyhaplogroup R1b1a andI (mostly subclades ofI2a) haplotypes.mtDNA belonged mostly toU (particularly subclades ofU5 andU4).[30] People of the Balkan Neolithic were found to harbor 98% Anatolian ancestry and 2% WHG ancestry. By theChalcolithic, people of theCucuteni–Trypillia culture were found to harbor about 20% hunter-gatherer ancestry, which was intermediate between EHG and WHG. People of theGlobular Amphora culture were found to harbor ca. 25% WHG ancestry, which is significantly higher than Middle Neolithic groups of Central Europe.[30]
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), theAncient North Eurasians (ANE, Upper Paleolithic Siberians later associated with the laterIndo-European expansion, present in proportions up to 20%), and finally theEarly 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). TheEarly European Farmer (EEF) component was identified based on the genome of a woman buried c. 7,000 years ago in aLinear Pottery culture grave inStuttgart, Germany.[32]
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.[33]
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).[34] 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%.[35]
Analysis of remains from the Grotta Continenza inItaly showed that out of six remains, three buried betweenc. 10,000 BC and 7000 BC belonged toI2a-P214; and two-times the maternal haplogroupsU5b1 and oneU5b3.[36][37] Around 6000 BC, the WHGs of Italy were almost completely genetically replaced by EEFs (twoG2a2) and oneHaplogroup R1b, although WHG ancestry slightly increased in subsequent millennia.[38]
Neolithic individuals in theBritish 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.[39]
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.[40]
A study on Mesolithic hunter-gatherers fromDenmark found that they were related to contemporary Western hunter-gatherers, and are associated with theMaglemose,Kongemose andErtebølle 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 theFunnelbeaker culture.[41]
According toDavid Reich, DNA analysis has shown that Western Hunter Gatherers were typically dark skinned, dark haired, and blue eyed.[7] The dark skin was due to theirOut-of-Africa origin (allHomo sapiens populations having had initially dark skin), while the blue eyes were the result of a variation in their OCA2 gene, which causediris depigmentation.[44]
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.[45] 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.[46]
Two WHG skeletons with incompleteSNPs, La Braña andCheddar Man, are predicted to have had dark or dark to black skin, whereas two other WHG skeletons with complete SNPs, "Sven" andLoschbour man, are predicted to have had dark or intermediate-to-dark and intermediate skin, respectively.[47][29][b] Spanish biologistCarles Lalueza-Fox said the La Braña-1 individual had dark skin, "although we cannot know the exact shade."[49]
According to a 2020 study, the arrival ofEarly European Farmers (EEFs) from westernAnatolia from 8500 to 5000 years ago, along withWestern Steppe Herders during theBronze Age, caused a rapid evolution of European populations towards lighter skin and hair.[44] Admixture between hunter-gatherer and agriculturist populations was apparently occasional, but not extensive.[50]
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Some authors have expressed caution regardingskin 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."[51]
Geneticist Susan Walsh atIndiana University–Purdue University Indianapolis, who worked on Cheddar Man project, said that "we simply don't know his skin colour".[52] German biochemistJohannes 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-dayCentral Africa or people from theArab region. It is only certain that they did not carry any known mutation responsible for thelight skin in subsequent populations of Europeans.[53]
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.[54]
{{cite journal}}: CS1 maint: multiple names: authors list (link)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 Bølling-Allerød 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
{{cite journal}}: CS1 maint: numeric names: authors list (link)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).
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.
Homo sapiens arrived 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
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.
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.
{{cite book}}: CS1 maint: publisher location (link)