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Genetic Relations to Ancient North Eurasians (ANE):
Eastern hunter-gatherer (EHG)
Caucasus hunter-gatherer (CHG)
Zagros/Iranian Hunter-Gatherer (IHG)
Iranian Neolithic Farmers (INF)
Anatolian hunter-gatherer (AHG)
Anatolian Neolithic Farmers (ANF)
Early European Farmers (EEF)
Yamnaya/Steppe Herders (WSH)
Villabruna 1 (burial)/Ripari Villabruna rock shelter in northern Italy (14,000 years old)
Satsurblia Cave (burial) in the Country of Georgia (13,000 years old)
Motala (burial) (8,000 years old)
A 23,000-year-old southern Iberian individual links human groups that lived in Western Europe before and after the Last Glacial Maximum
“Following the Bølling/Allerød warming interstadial (14,000 years ago), the Goyet Q2 cluster was replaced by the Villabruna cluster in central Europe, named for its oldest Epigravettian-associated individual from northern Italy, but which also includes most of the Epipalaeolithic- and Mesolithic-associated groups from central and western Europe, all of which are also known as western hunter-gatherers (WHG). In this genetic landscape, Iberian hunter-gatherers (HG) stood out as they retained higher proportions of the Goyet Q2-like ancestry during the Epipalaeolithic and Mesolithic periods and thus are often considered separate.” ref
Human populations underwent range contractions during the Last Glacial Maximum (LGM) which had lasting and dramatic effects on their genetic variation. The genetic ancestry of individuals associated with the post-LGM Magdalenian technocomplex has been interpreted as being derived from groups associated with the pre-LGM Aurignacian. However, both these ancestries differ from that of central European individuals associated with the chronologically intermediate Gravettian. Thus, the genomic transition from pre- to post-LGM remains unclear also in western Europe, where we lack genomic data associated with the intermediate Solutrean, which spans the height of the LGM. Here we present genome-wide data from sites in Andalusia in southern Spain, including from a Solutrean-associated individual from Cueva del Malalmuerzo, directly dated to ~23,000 years ago.” ref
“The Malalmuerzo individual carried genetic ancestry that directly connects earlier Aurignacian-associated individuals with post-LGM Magdalenian-associated ancestry in western Europe. This scenario differs from Italy, where individuals associated with the transition from pre- and post-LGM carry different genetic ancestries. This suggests different dynamics in the proposed southern refugia of Ice Age Europe and posits Iberia as a potential refugium for western European pre-LGM ancestry. Moreover, individuals from Cueva Ardales, which were thought to be of Palaeolithic origin, date younger than expected and, together with individuals from the Andalusian sites Caserones and Aguilillas, fall within the genetic variation of the Neolithic, Chalcolithic, and Bronze Age individuals from southern Iberia.” ref
“Abstract: The concept of grief, the metamorphosis of the deceased into the departed, a subject recreated and rethought by the psyche, is crucial for understanding the significance of the grave and funeral rites. We can divide the funeral rites into three phases: seeing the dead person presented socialized, hiding him to begin the mourning process, and finally metamorphosing him into the deceased. Moreover, these three phases typically require the involvement of several community members, some of whom may be less affected by sorrow — a factor that hinders action — compared to close relatives. Considering these factors, it becomes apparent that grief and, consequently, the tomb are more fundamentally social phenomena than cultural ones. The cultural aspect is an overlay, as beliefs and religions facilitate the mourning process by providing guidelines for conduct and contemplation. An evolutionary perspective on the recognition of death and griefs considers these definitions, cognitive developments during human growth, and the cognitive evolution of hominids. Recognizing another’s death without integrating the concept of one’s mortality could have emerged early in human evolution and been a factor in developing consciousness in a feedback loop. Moreover, the funerary rites and tombs are probably older than is commonly accepted by many researchers to date.” ref
Ancient North Eurasian
“In archaeogenetics, the term Ancient North Eurasian (ANE) is the name given to an ancestral component that represents the lineage of the people of the Mal’ta–Buret’ culture (c. 24,000 years ago) and populations closely related to them, such as the Upper Paleolithic individuals from Afontova Gora in Siberia. Genetic studies also revealed that the ANE are closely related to the remains of the preceding Yana Culture (c. 32,000 years ago), which were dubbed as ‘Ancient North Siberians‘ (ANS). Ancient North Eurasians are predominantly of West Eurasian ancestry (related to European Cro-Magnons and ancient and modern peoples in West Asia) who arrived in Siberia via the “northern route,” but also derive a significant amount of their ancestry (c. 1/3) from an East Eurasian source, having arrived to Siberia via the “southern route.” ref
“Around 20,000 to 25,000 years ago, a branch of Ancient North Eurasian people mixed with Ancient East Asians, which led to the emergence of Ancestral Native American, Ancient Beringian, and Ancient Paleo-Siberian populations. It is unknown exactly where this population admixture took place, and two opposing theories have put forth different migratory scenarios that united the Ancient North Eurasians with ancient East Asian populations. Later, ANE populations migrated westward into Europe and admixed with European Western hunter-gatherer (WHG)-related groups to form the Eastern Hunter-Gatherer (EHG) group, which later admixed with Caucasus hunter-gatherers to form the Western Steppe Herder group, which became widely dispersed across Eurasia during the Bronze Age.” ref
“ANE ancestry has spread throughout Eurasia and the Americas in various migrations since the Upper Paleolithic, and more than half of the world’s population today derives between 5 and 42% of their genomes from the Ancient North Eurasians. Significant ANE ancestry can be found in Native Americans, as well as in Europe, South Asia, Central Asia, and Siberia. It has been suggested that their mythology may have featured narratives shared by both Indo-European and some Native American cultures, such as the existence of a metaphysical world tree and a fable in which a dog guards the path to the afterlife.” ref
Eastern hunter-gatherer
“In archaeogenetics, eastern hunter-gatherer (EHG), sometimes east European hunter-gatherer or eastern European hunter-gatherer, is a distinct ancestral component that represents Mesolithic hunter-gatherers of Eastern Europe. The eastern hunter-gatherer genetic profile is mainly derived from Ancient North Eurasian (ANE) ancestry, which was introduced from Siberia, with a secondary and smaller admixture of European western hunter-gatherers (WHG). 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. During the Mesolithic, the EHGs inhabited an area stretching from the Baltic Sea to the Urals and downwards to the Pontic–Caspian steppe. Along with Scandinavian hunter-gatherers (SHG) and western hunter-gatherers (WHG), the EHGs constituted one of the three main genetic groups in the postglacial period of early Holocene Europe.” ref
“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. During the Neolithic and early Eneolithic, likely during the 4th millennium BC EHGs on the Pontic–Caspian steppe mixed with Caucasus hunter-gatherers (CHGs) with the resulting population, almost half-EHG and half-CHG, forming the genetic cluster known as Western Steppe Herder (WSH). WSH populations closely related to the people of the Yamnaya culture are supposed to have embarked on a massive migration leading to the spread of Indo-European languages throughout large parts of Eurasia.” ref
Caucasus hunter-gatherer
“Caucasus hunter-gatherer (CHG), also called Satsurblia cluster, is an anatomically modern human genetic lineage, first identified in a 2015 study, based on the population genetics of several modern Western Eurasian (European, Caucasian, and Near Eastern) populations. It represents an ancestry maximized in some Upper Paleolithic and Mesolithic hunter-gatherer groups in the Caucasus. These groups are also very closely related to Mesolithic hunter-gatherers and Neolithic farmers in the Iranian Plateau, who are sometimes included within the CHG group. Ancestry that is closely related to CHG-Iranian Neolithic farmers is also known from further east, including from the Bactria–Margiana Archaeological Complex and the Harappan/Indus Valley Civilisation. Caucasus hunter-gatherers and Eastern hunter-gatherers are ancestral in roughly equal proportions to the Western Steppe Herders (WSH), who were widely spread across Europe and Asia beginning during the Chalcolithic.” ref
“The CHG lineage is suggested to have diverged from the ancestor of Western Hunter-Gatherers (WHGs) probably during the Last Glacial Maximum (sometime between 45,000 to 26,000 years ago). They further separated from the Anatolian hunter-gatherer (AHG) lineage later, suggested to around 25,000 years ago during the late LGM period. The Caucasus hunter-gatherers managed to survive in isolation since the late LGM period as a distinct population, and display high genetic affinities to Mesolithic and Neolithic populations on the Iranian plateau, such as Neolithic specimens found in Ganj Dareh. The CHG display higher genetic affinities to European and Anatolian groups than Iranian hunter-gatherers do, suggesting a possible cline and geneflow into the CHG and less into Mesolithic and Neolithic Iranian groups.” ref
“Lazaridis et. al (2016) models the CHG as a mixture of Neolithic Iranians, Western Hunter-Gatherers, and Eastern Hunter-Gatherers. In addition, the CHG cluster with early Iranian farmers, who significantly do not share alleles with early Levantine farmers. An alternative model without the need of significant amounts of ANE ancestry has been presented by Vallini et al. (2024), suggesting that the initial Iranian hunter-gatherer-like population, which is basal to the CHG formed primarily from a deep Ancient West Eurasian lineage (‘WEC2’, c. 72%), and from varying degrees of Ancient East Eurasian (c. 10%) and Basal Eurasian (c. 18%) components. The Ancient West Eurasian component associated with Iranian hunter-gatherers (WEC2) is inferred to have diverged from the West Eurasian Core lineage (represented by Kostenki-14; WEC), with the WEC2 component staying in the region of the Iranian Plateau, while the proper WEC component expanded into Europe.” ref
“Irving-Pease et. al (2024) models CHG as being derived from an Out of Africa population that split into basal Northern Europeans and West Asians. The latter was where CHG originated from. At the beginning of the Neolithic, at c. 8000 BC, they were probably distributed across western Iran and the Caucasus, and people similar to northern Caucasus and Iranian plateau hunter-gatherers arrived before 6000 BC in Pakistan and north-west India. A roughly equal merger between the CHG and Eastern Hunter-Gatherers in the Pontic–Caspian steppe resulted in the formation of the Western Steppe Herders (WSHs). The WSHs formed the Yamnaya culture and subsequently expanded massively throughout Europe during the Late Neolithic and Early Bronze Age c. 3000—2000 BCE. Caucasus hunter-gatherer/Iranian-like ancestry, was first reported as maximized in hunter-gatherers from the South Caucasus and early herders/farmers in northwestern Iran, particularly the Zagros, hence the label “CHG/Iranian.” ref
Epigravettian
“The Epigravettian (Greek: epi “above, on top of,” and Gravettian ) was one of the last archaeological industries and cultures of the European Upper Paleolithic . It emerged after the Last Glacial Maximum around ~21,000 years ago or 19,050 BCE. It succeeds the Gravettian culture in Italy. Initially named Tardigravettian (Late Gravettian) in 1964 by Georges Laplace in reference to several lithic industries found in Italy, it was later renamed in order to better emphasize its independent character. Three subphases, the Early Epigravettian (20,000 to 16,000 years ago), the Evolved Epigravettian (16,000 to 14,000 years ago), and the Final Epigravettian (14,000 to 8,000 years ago) have been established, which were further subdivided and reclassified. In this sense, the Epigravettian is simply the Gravettian after ~21,000 years ago, when the Solutrean had replaced the Gravettian in most of France and Spain. Epigravettian Venus like artifacts, maybe 24,000 – 18,000 years ago. An Epigravettian ceramic figurine of a horse or deer, Vela Spila, Croatia, is dated to 15,400-14,600 years ago.” ref
“Several Epigravettian cultural centers have developed contemporaneously after 22,000 years years ago in Europe. These range across southern, central, and most of eastern Europe, including southwestern France, Italy, Southeast Europe , the Caucasus, Ukraine, and Western Russia to the banks of the Volga River. Its lithic complex was first documented at numerous sites in Italy. Great geographical and local variability of the facies is present, however all sites are characterized by the predominance of microliths, such as backed blades, backed points, and bladelets with retouched end. The Epigravettian is the last stage of the Upper Paleolithic succeeded by Mesolithic cultures after 10,000 years ago. In a genetic study published in Nature in May 2016, the remains of an Epigravettian male from Ripari Villabruna in Italy were examined.” ref
“He carried the paternal haplogroup R1b1 and the maternal haplogroup U5b . An Epigravettian from the Satsurblia Cave in Georgia , who was examined in a previous study, has been found to be carrying the paternal haplogroup J1 and the maternal haplogroup K3. Analysis of Epigravettian producing individuals in Italy indicates that they not closely related to earlier Gravettian-producing populations of the peninsula, and instead belong to the Villabruna genetic cluster . This group is more closely related to ancient and modern peoples in the Middle East and the Caucasus than earlier European Cro-Magnons . Epigravettian peoples belonging to the Western Hunter Gatherer genetic cluster expanded across Western Europe at the end of the Pleistocene, largely replacing the producers of the Magdalenian culture that previously dominated the region.” ref
“Magdalenian cultures (Pre-R1b Y-DNA Europe) are later cultures of the Upper Paleolithic and Mesolithic in western Europe. They date from around 17,000 to 12,000 years ago. It is named after the type site of La Madeleine, a rock shelter located in the Vézère valley, commune of Tursac, in France’s Dordogne department. The Magdalenian is associated with reindeer hunters, although Magdalenian sites contain extensive evidence for the hunting of red deer, horses, and other large mammals present in Europe toward the end of the last glacial period. The culture was geographically widespread, and later Magdalenian sites stretched from Portugal in the west to Poland in the east, and as far north as France, the Channel Islands, England, and Wales. Besides La Madeleine, the chief stations of the Magdalenian are Les Eyzies, Laugerie-Basse, and Gorges d’Enfer in the Dordogne; Grotte du Placard in Charente and others in south-west France.” ref
“Magdalenian peoples produced a wide variety of art, including figurines and cave paintings. Evidence has been found suggesting that Magdalenian peoples regularly engaged in (probably ritualistic) cannibalism along with producing skull cups. Genetic studies indicate that the Magdalenian peoples were largely descended from earlier Western European Cro-Magnon groups like the Gravettians that were present in Western Europe over 30,000 years ago prior to the Last Glacial Maximum, who had retreated to southwestern Europe during the LGM. Madgalenian peoples were largely replaced and, in some areas, absorbed by Epigravettian producing groups of Villabruna/Western Hunter Gatherer ancestry at the end of the Pleistocene.” ref
“The genes of seven Magdalenians, the El Miron Cluster in Iberia, have shown a close relationship to a population who had lived in Northern Europe some 20,000 years previously. The analyses suggested that 70-80% of the ancestry of these individuals was from the population represented by Goyet Q116-1, associated with the Aurignacian culture of about 35,000 BP, from the Goyet Caves in modern Belgium. It has also been found that Magdalenians are also closely related to western Gravettians who inhabited France and Spain prior to the Last Glacial Maximum. The 15,000-year-old GoyetQ2 individual from Goyet Caves is often used as a proxy for Magdalenian ancestry.” ref
“Analysis of genomes of GoyetQ2-related Magdalenians suggest that, like earlier Cro-Magnon groups, they probably had a relatively dark skin tone compared to modern Europeans. A 2023 study proposed that relative to earlier Western European Cro-Magnon-related groups like Goyet Q116-1-related Aurignacian and the Western Gravettian-associated Fournol cluster, the Goyet-Q2-related Magdalenians appear to have carried significant (~30% ancestry) from the Villabruna cluster (thought to be of southeastern European origin, and sharing affinities to West Asian peoples not found in earlier European hunter-gatherers) associated with the Epigravettian.” ref
“The three samples of Y-DNA included two samples of haplogroup I and one sample of HIJK. All samples of mtDNA belonged to U, including five samples of U8b and one sample of U5b. Around 14-12,000 years ago, the Western Hunter-Gatherer cluster (which predominantly descended from the Villabruna cluster, with possible ancestry related to the Goyet-Q2 cluster), expanded northwards across the Alps, largely replacing the Goyet-Q2 cluster associated Magdalenian groups in Western Europe. In France and Spain, significant GoyetQ2-related ancestry persisted into the Mesolithic and Neolithic, with some Neolithic individuals in France and Spain largely of Early European Farmer descent showing significant GoyetQ2 ancestry.” ref
Western hunter-gatherer
“In archaeogenetics, western hunter-gatherer (WHG, also known as west European hunter-gatherer, western European hunter-gatherer or Oberkassel cluster) (c. 15,000~5,000 years ago) is 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. It is closely associated and sometimes considered synonymous with the concept of the Villabruna cluster, named after Ripari Villabruna cave in Italy, known from the terminal Pleistocene of Europe, which is largely ancestral to later WHG populations.” ref
“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 their origin likely somewhere in southeast Europe or West Asia. They had expanded into the Italian and Iberian Peninsulas by approximately 19,000 years ago, and subsequently expanded across Western Europe at the end of the Pleistocene around 14-12,000 years ago, largely replacing the Magdalenians who previously dominated the region. The Magdalenians largely descended from earlier Western European Cro-Magnon groups that had arrived in the region over 30,000 years ago, prior to the Last Glacial Maximum.” ref
“WHGs constituted one of the main genetic groups in the postglacial period of early Holocene Europe, along with eastern hunter-gatherers (EHG) in Eastern Europe. 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. EHGs primarily consisted of a mixture of WHG-related and Ancient North Eurasian (ANE) ancestry. Scandinavia was inhabited by Scandinavian hunter-gatherers (SHGs), which were a mixture between WHG and EHG. In the Iberian Peninsula, early Holocene hunter-gathers consisted of a mixture of WHG and Magdalenian Cro-Magnon (GoyetQ2) ancestry.” ref
“Once the main population throughout Europe, the WHGs were largely replaced by successive expansions of Early European Farmers (EEFs) of Anatolian origin during the early Neolithic, who generally carried a minor amount of WHG ancestry due to admixture with WHG groups during their European expansion. Among modern-day populations, WHG ancestry is most common among populations of the eastern Baltic region. Western hunter-gatherers (WHG) are recognised as a distinct ancestral component contributing to the ancestry of most modern Europeans. Most Europeans can be modeled as a mixture of WHG, EEF, and WSH from the Pontic–Caspian steppe. WHGs also contributed ancestry to other ancient groups such as Early European Farmers (EEF), who were, however, mostly of Anatolian descent. 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.” ref
“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 Asian refugia. It is thought that their ancestors separated from eastern Eurasians around 40,000 years ago, and from Ancient North Eurasians (ANE) prior to 24,000 years ago (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 38,000 years ago, shortly after the divergence of West-Eurasian and East-Eurasian lineages. 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 those found in the Zlaty Kun, Peștera cu Oase and Bacho Kiro caves, being unrelated to Western hunter-gatherers but closer to Ancient East Eurasians or basal to both.” ref
“The relationships of the WHG/Villabruna cluster to other Paleolithic human groups in Europe and West Asia are obscure and subject to conflicting intepretations. 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 the Last Glacial Maximum. WHG genomes display 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.” ref
“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. 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 eastern Gravettian producing Cro-Magnon groups, which may reflect shared ancestry in the Balkan region.” ref
“The earliest known individuals of predominantly WHG/Villabruna ancestry in Europe are known from Italy, dating to around 17,000 years ago, though an individual from El Mirón cave in northern Spain with 43% Villabruna ancestry is known from 19,000 years ago. Early WHG/Villabruna populations are associated with the Epigravettian archaeological culture, which largely replaced populations associated with the Magdalenian culture about 14,000 years ago (the ancestry of Magdalenian-associated Goyet-Q2 cluster primarily descended from the earlier Solutrean, and western Gravettian-producing groups in France and Spain). 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.” ref
“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 the Oberkassel cluster, after one of the oldest WHG individuals found north of the Alps. 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.” ref
“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, as well as other European groups such as eastern hunter-gatherers. The relationship of WHGs to the EHGs remains inconclusive. 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. In the Iberian Peninsula early Holocene hunter-gathers consisted of populations with a mixture of WHG and Magdalenian Cro-Magnon (GoyetQ2) ancestry.” ref
“People of the Mesolithic Kunda culture and the Narva culture of the eastern Baltic were a mix of WHG and EHG, 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. People of the Pit–Comb Ware culture (CCC) of the eastern Baltic were closely related to EHG. 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.” ref
“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.” ref
“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. 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 BCE). 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).” ref
“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). 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). 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.” ref
Scandinavian hunter-gatherer
“In archaeogenetics, the term Scandinavian hunter-gatherer (SHG) is the name given to a distinct ancestral component that represents descent from Mesolithic hunter-gatherers of Scandinavia. Genetic studies suggest that the SHGs were a mix of western hunter-gatherers (WHGs) initially populating Scandinavia from the south during the Holocene, and eastern hunter-gatherers (EHGs), who later entered Scandinavia from the north along the Norwegian coast. During the Neolithic, they admixed further with Early European Farmers (EEFs) and Western Steppe Herders (WSHs). Genetic continuity has been detected between the SHGs and members of the Pitted Ware culture (PWC), and to a certain degree, between SHGs and modern northern Europeans. The Sámi, on the other hand, have been found to be completely unrelated to the PWC.” ref
“Scandinavian hunter-gatherers (SHG) were identified as a distinct ancestral component by Lazaridis et al. (2014). A number of remains examined at Motala, Sweden, and a separate group of remains from 5,000 year-old hunter-gatherers of the Pitted Ware culture (PWC), were identified as belonging to SHG. The study found that an SHG individual from Motala (‘Motala12’) could be successfully modelled as being of c. 81% western hunter-gatherer (WHG) ancestry, and c. 19% Ancient North Eurasian (ANE) ancestry. Haak et al. (2015) examined the remains of six SHGs buried at Motala between ca. 6000 BCE and 5700 BCE. Of the four males surveyed, three carried the paternal haplogroup I2a1 or various subclades of it, while the other carried I2c. With regard to mtDNA, four individuals carried subclades of U5a, while two carried U2e1.” ref
“The study found SHGs to constitute one of the three main hunter-gatherer populations of Europe during the Mesolithic. The two other groups were WHGs and eastern hunter-gatherers (EHG). EHGs were found to be an ANE-derived population with significant admixture from a WHG-like source. SHGs formed a distinct cluster between WHG and EHG, and the admixture model proposed by Lazaridis et al. could be successfully replaced with a model that takes EHG as the source population for the ANE-like ancestry, with an admixture ratio of ~65% (WHG): ~35% (EHG). SHGs living between 6000 BCE and 3000 BCE were found to be largely genetically homogeneous, with little admixture occurring among them during this period. EHGs were found to be more closely related to SHGs than WHGs.” ref
“Mathieson et al. (2015) subjected the six SHGs from Motala to further analysis. SHGs appeared to have persisted in Scandinavia until after 5,000 years ago. The Motala SHGs were found to be closely related to WHGs. Lazaridis et al. (2016) confirmed SHGs to be a mix of EHGs (~43%) and WHGs (~57%). WHGs were modeled as descendants of the Upper Paleolithic people (Cro-Magnon) of the Grotte du Bichon in Switzerland with minor additional EHG admixture (~7%). EHGs derived c. 75% of their ancestry from ANEs. Günther et al. (2018) examined the remains of seven SHGs. All three samples of Y-DNA extracted belonged to subclades of I2. With respects to mtDNA, four samples belonged to U5a1 haplotypes, while three samples belonged U4a2 haplotypes. All samples from western and northern Scandinavia carried U5a1 haplotypes, while all the samples from eastern Scandinavia except from one carried U4a2 haplotypes.” ref
“The authors of the study suggested that SHGs were descended from a WHG population that had entered Scandinavia from the south, and an EHG population which had entered Scandinavia from the northeast along the coast. The WHGs who entered Scandinavia are believed to have belonged to the Ahrensburg culture. These WHGs and EHGs had subsequently mixed, and the SHGs gradually developed their distinct character. The SHGs from western and northern Scandinavia had more EHG ancestry (c. 49%) than individuals from eastern Scandinavia (c. 38%). The SHGs were found to have a genetic adaptation to high latitude environments, including high frequencies of low pigmentation variants and genes designed for adaptation to the cold and physical performance. SHGs displayed a high frequency of the depigmentation alleles SLC45A2 and SLC24A5, and the OCA/Herc2, which affects eye pigmentation. These genes were much less common among WHGs and EHGs. A surprising continuity was displayed between SHGs and modern populations of Northern Europe in certain respects. Most notably, the presence of the protein TMEM131 among SHGs and modern Northern Europeans was detected. This protein may be involved in long-term adaptation to the cold.” ref
“In a genetic study published in Nature Communications in January 2018, the remains of an SHG female at Motala, Sweden, between 5750 BCE and 5650 BCE was analyzed. She was found to be carrying U5a2d and “substantial ANE ancestry.” The study found that Mesolithic hunter-gatherers of the eastern Baltic also carried high frequencies of the HERC2 allele, and increased frequencies of the SLC45A2 and SLC24A5 alleles. They, however, harbored less EHG ancestry than SHGs. Genetic continuity between the SHGs and the Pitted Ware culture of the Neolithic was detected. The results further underpinned the previous suggestion that SHGs were descended from the northward migration of WHGs and a subsequent southward migration of EHGs. A certain degree of continuity between SHGs and northern Europeans was detected.” ref
“A study published in Nature in February 2018 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 BCE). These were determined to be an intermediate between EHG and SHG. Samples of Y-DNA extracted from these individuals belonged exclusively to R haplotypes (particularly subclades of R1b1 and R1a)) and I haplotypes (particularly subclades of I2). mtDNA belonged almost exclusively to U (particularly subclades of U5 and U4). According to Mathieson et al. (2015), 50% of Scandinavian Hunter Gatherers from Motala carried the derived variant of EDAR-V370A. This variant is typical of modern East Asian populations, and is known to affect dental morphology and hair texture, and also chin protrusion and ear morphology, as well as other facial features.” ref
“The authors did not detect East Asian ancestry in the Scandinavian Hunter Gatherers, and speculated that this gene might not have originated in East Asia, as is commonly believed. However, more recent research incorporating ancient Northeast Asian samples has confirmed that EDAR-V370A originated in Northeast Asia, and spread to West Eurasian populations such as Motala in the Holocene period. Mathieson et al. (2015) also reported: “A second surprise is that, unlike closely related western hunter-gatherers, the Motala samples have predominantly derived pigmentation alleles at SLC45A2 and SLC24A5. The study by Günther et al. (2018) further discovered that SHGs “show a combination of eye color varying from blue to light brown and light skin pigmentation. This is strikingly different from the WHGs—who have been suggested to have the specific combination of blue eyes and dark skin and EHGs—who have been suggested to be brown-eyed and light-skinned.” ref
“Four SHGs from the study yielded diverse eye and hair pigmentation predictions: one individual (SF12) was predicted to be most likely to have had dark hair and blue eyes; a second individual (Hum2) most likely had dark hair and brown eyes; a third (SF9) was predicted to have had light hair and brown eyes; and a fourth individual (SBj) was predicted to have had light hair, with the most likely hair colour being blonde, and blue eyes. Of the SHGs from Motala, four were probably dark-haired, and two others were probably light-haired, and may have been blond. In addition, all of the six SHGs from Motala had high probabilities of being blue-eyed. Both light and dark skin pigmentation alleles are found at intermediate frequencies in the Scandinavian Hunter Gatherers sampled, but only one individual had exclusively light-skin variants of two different SNPs.” ref
“The study found that depigmentation variants of genes for light skin pigmentation (SLC24A5, SLC45A2) and blue eye pigmentation (OCA2/HERC2) are found at high frequency in SHGs relative to WHGs and EHGs, which the study suggests cannot be explained simply as a result of the admixture of WHGs and EHGs. The study argues that these allele frequencies must have continued to increase in SHGs after admixture, which was probably caused by environmental adaptation to high latitudes. On the basis of archaeological and genetic evidence, the Swedish archaeologist Oscar D. Nilsson has made forensic reconstructions of both male and female SHGs.” ref
“Light skin is a human skin color that has a low level of eumelanin pigmentation as an adaptation to environments of low UV radiation. Due to the migrations of people in recent centuries, light-skinned populations today are found all over the world. Light skin is most commonly found amongst the native populations of Europe, East Asia, West Asia, Central Asia, Siberia, and North Africa as measured through skin reflectance. People with light skin pigmentation are often referred to as “white“ although these usages can be ambiguous in some countries where they are used to refer specifically to certain ethnic groups or populations.” ref
“Humans with light skin pigmentation have skin with low amounts of eumelanin, and possess fewer melanosomes than humans with dark skin pigmentation. Light skin provides better absorption qualities of ultraviolet radiation, which helps the body to synthesize higher amounts of vitamin D for bodily processes such as calcium development. On the other hand, light-skinned people who live near the equator, where there is abundant sunlight, are at an increased risk of folate depletion. As a consequence of folate depletion, they are at a higher risk of DNA damage, birth defects, and numerous types of cancers, especially skin cancer.” ref
“Humans with darker skin who live further from the tropics may have lower vitamin D levels, which can also lead to health complications, both physical and mental, including a greater risk of developing schizophrenia. These two observations form the “vitamin D–folate hypothesis,” which attempts to explain why populations that migrated away from the tropics into areas of low UV radiation evolved to have light skin pigmentation. The distribution of light-skinned populations is highly correlated with the low ultraviolet radiation levels of the regions inhabited by them. Historically, light-skinned populations almost exclusively lived far from the equator, in high-latitude areas with low sunlight intensity.” ref
“It is generally accepted that dark skin evolved as a protection against the effect of UV radiation; eumelanin protects against both folate depletion and direct damage to DNA. This accounts for the dark skin pigmentation of Homo sapiens during their development in Africa; the major migrations out of Africa to colonize the rest of the world were also dark-skinned. It is widely supposed that light skin pigmentation developed due to the importance of maintaining vitamin D3 production in the skin. Strong selective pressure would be expected for the evolution of light skin in areas of low UV radiation. After the ancestors of West Eurasians and East Eurasians diverged more than 40,000 years ago, lighter skin tones evolved independently in a subset of each of the two populations. In West Eurasians, the A111T allele of the rs1426654 polymorphism in the pigmentation gene SLC24A5 has the largest skin-lightening effect and is widespread in Europe, South Asia, Central Asia, the Near East, and North Africa.” ref
“In a 2013 study, Canfield et al. established that SLC24A5 sits in a block of haplotypes, one of which (C11) is shared by virtually all chromosomes that bear the A111T variant. This “equivalence” between C11 and A111T indicates that all people who carry this skin-lightening allele descend from a common origin: a single carrier who lived most likely “between the Middle East and the Indian subcontinent.” Canfield et al. attempted to date the A111T mutation but only constrained the age range to before the Neolithic. However, a second study from the same year (Basu Mallick et al.) estimated the coalescent age (split date) for this allele to be between ~28,000 and ~22,000 years ago.” ref
“The second most important skin-lightening factor in West Eurasians is the depigmenting allele F374 of the rs16891982 polymorphism located in the melanin-synthesis gene SLC45A2. From its low haplotype diversity, Yuasa et al. (2006) likewise concluded that this mutation (L374F) “occurred only once in the ancestry of Caucasians.” Summarising these studies, Hanel and Carlberg (2020) decided that the alleles of the two genes SLC24A5 and SLC45A2, which are most associated with lighter skin color in modern Europeans, originated in West Asia about 22,000 to 28,000 years ago and these two mutations each arose in a single carrier. This is consistent with Jones et al. (2015), who reconstructed the relationship between Near Eastern Neolithic farmers and Caucasus Hunter-Gatherers: two populations that carried the light-skin variant of SLC24A5.” ref
“Analyzing newly sequenced ancient genomes, Jones et al. estimated the split date at ~24,000 years ago and localized the separation to somewhere south of the Caucasus. However, a coalescent analysis of this allele by Crawford et al. (2017) gave a more narrowly constrained, and earlier, split date of ~29,000 years ago (with a 95% confidence window from 28,000 to 31,000 years ago). The light-skin variants of SLC24A5 and SLC45A2 were present in Anatolia by 9,000 years ago, where they became associated with the Neolithic Revolution. From here, their carriers spread Neolithic farming across Europe. Lighter skin and blond hair also evolved in the Ancient North Eurasian population.” ref
“A further wave of lighter-skinned populations across Europe (and elsewhere) is associated with the Yamnaya culture and the Indo-European migrations bearing Ancient North Eurasian ancestry and the KITLG allele for blond hair. Furthermore, the SLC24A5 gene linked with light pigmentation in Europeans was introduced into East Africa from Europe over five thousand years ago. These alleles can now be found in the San, Ethiopians, and Tanzanian populations with Afro-Asiatic ancestry. The SLC24A5 in Ethiopia maintains a substantial frequency with Semitic and Cushitic speaking populations, compared with Omotic, Nilotic, or Niger-Congo speaking groups. It is inferred that it may have arrived into the region via migration from the Levant, which is also supported by linguistic evidence. In the San people, it was acquired from interactions with Eastern African pastoralists. Meanwhile, in the case of East Asia and the Americas, a variation of the MFSD12 gene is responsible for lighter skin color. The modern association between skin tone and latitude is thus a relatively recent development.” ref
“According to Crawford et al. (2017), most of the genetic variants associated with light and dark pigmentation appear to have originated more than 300,000 years ago. African, South Asian, and Australo-Melanesian populations also carry derived alleles for dark skin pigmentation that are not found in Europeans or East Asians. Huang et al. (2021) found the existence of “selective pressure on light pigmentation in the ancestral population of Europeans and East Asians”, prior to their divergence from each other. Skin pigmentation was also found to be affected by directional selection towards darker skin among Africans, as well as lighter skin among Eurasians.” ref
“Crawford et al. (2017) similarly found evidence for selection towards light pigmentation prior to the divergence of West Eurasians and East Asians. The A111T mutation in the SLC24A5 gene predominates in populations with Western Eurasian ancestry. The geographical distribution shows that it is nearly fixed in all of Europe and most of the Middle East, extending east to some populations in present-day Pakistan and Northern India. It shows a latitudinal decline toward the Equator, with high frequencies in North Africa (80%), and intermediate (40−60%) in Ethiopia and Somalia.” ref
“Some authors have expressed caution regarding the skin pigmentation SNP predictions in early Paleolithic groups. According to Ju et al. (2021): “Relatively dark skin pigmentation in Early Upper Paleolithic Europe would be consistent with those populations being relatively poorly adapted to high-latitude conditions as a result of having recently migrated from lower latitudes. On the other hand, although we have shown that these populations carried few of the light pigmentation alleles that are segregating in present-day Europe, they may have carried different alleles that we cannot now detect. As an extreme example, Neanderthals and the Altai Denisovan individual show genetic scores that are in a similar range to Early Upper Paleolithic individuals, but it is highly plausible that these populations, who lived at high latitudes for hundreds of thousands of years, would have adapted independently to low UV levels. For this reason, we cannot confidently make statements about the skin pigmentation of ancient populations.” ref
“In 2015, it was discovered that 13,000-year-old samples of Caucasus Hunter-Gatherers (CHG) from Georgia carried the mutation and derived alleles for very fair-skinned pigmentation similar to Early Farmers (EF). This trait was said to have a relatively long history in Eurasia and risen to high frequency during the Neolithic expansion, with its origin probably predating the Last Glacial Maximum (LGM).” ref
“In the same year, a study found that genes contributing to fair skin were nearly fixed in the Anatolian Neolithic Farmers: “The second strongest signal in our analysis is at the derived allele of rs16891982 in SLC45A2, which contributes to light skin pigmentation and is almost fixed in present-day Europeans but occurred at much lower frequency in ancient populations. In contrast, the derived allele of SLC24A5 that is the other major determinant of light skin pigmentation in modern Europe, appears fixed in the Anatolian Neolithic, suggesting that its rapid increase in frequency to around 0.9 (90%) in Early Neolithic Europe was mostly due to migration.” ref
“In 2018, a study was released showing many late Mesolithic Scandinavians from 9,500 years ago in Northern Europe had blonde hair and light skin, which was in contrast to some of their contemporaries, the darker Western Hunter Gatherers (WHG). However, a 2024 paper found that phenotypically most of their studied WHG individuals carried the dark skin and blue eyes characteristic of WHGs, but some other WHGs in France they sequenced also had pale to intermediate skin pigmentation. Another entry in 2018, showed that the Eastern Hunter Gatherers (EHG), Scandinavian Hunter Gatherers (SHG), and the Baltic foragers, all had the derived alleles for light skin pigmentation.” ref
“A study on the populations of the Chalcolithic Levant (6,000-7,000 years ago), found that an allele rs1426654 in the SLC24A5 gene which is one of the most important determinants of light pigmentation in West Eurasians, was fixed for the derived variants in all Levant Chalcolithic samples, suggesting that the light skinned phenotype may have been common in the community. The individuals also had a high incidence of genomic markers associated with blue-eye color. A paper conducted by Fregel, Rosa et al. (2018) showed that in North Africa, Late Neolithic Moroccans had the European/Caucasus derived SLC24A5 mutation and other alleles and genes that predispose individuals to lighter skin and eye colours.” ref
Iberian hunter-gatherers
“The hunter-gatherers from the Iberian Peninsula carry a mix of two older types of genetic ancestry: one that dates back to the Last Glacial Maximum and was once maximized in individuals attributed to Magdalenian culture and another one that is found everywhere in western and central Europe and had replaced the Magdalenian lineage during the Early Holocene everywhere except the Iberian Peninsula,” explains Vanessa Villalba-Mouco of the Max Planck Institute for the Science of Human History, first author of the study. The researchers hope that ongoing efforts to decipher the genetic structure of late hunter-gatherer groups across Europe will help to even better understand Europe’s past and, in particular, the assimilation of a Neolithic way of life brought about by expanding farmers from the Near East during the Holocene.” ref
Ancient DNA from individuals spanning the last 8000 years helps clarify the history and prehistory of the Iberian Peninsula
“The paper published in Science focuses on slightly later time periods, and traces the population history of Iberia over the last 8000 years by analyzing ancient DNA from a huge number of individuals. The study, led by Harvard Medical School and the Broad Institute and including Haak and Villalba-Mouco, analyzed 271 ancient Iberians from the Mesolithic, Neolithic, Copper Age, Bronze Age, Iron Age, and historical periods. The large number of individuals allowed the team to make more detailed inferences about each time period than previously possible. The researchers found that during the transition to a sedentary farming life-style, hunter-gatherers in Iberia contributed subtly to the genetic make-up of newly arriving farmers from the Near East. “We can see that there must have been local mixture as the Iberian farmers also carry this dual signature of hunter-gatherer ancestry unique to Iberia,” explains Villalba-Mouco.” ref
“Between about 2500-2000 BCE, the researchers observed the replacement of 40% of Iberia’s ancestry and nearly 100% of its Y-chromosomes by people with ancestry from the Pontic Steppe, a region in what is today Ukraine and Russia. Interestingly, the findings show that in the Iron Age, “Steppe ancestry” had spread not only into Indo-European-speaking regions of Iberia but also into non-Indo-European-speaking ones, such as the region inhabited by the Basque. The researchers’ analysis suggests that present-day Basques most closely resemble a typical Iberian Iron Age population, including the influx of “Steppe ancestry,” but that they were not affected by subsequent genetic contributions that affected the rest of Iberia. This suggests that Basque speakers were equally affected genetically as other groups by the arrival of Steppe populations, but retained their language in any case. It was only after that time that they became relatively isolated genetically from the rest of the Iberian Peninsula.” ref
“Additionally, the researchers looked at historical periods, including times when Greek and later Roman settlements existed in Iberia. The researchers found that beginning at least in the Roman period, the ancestry of the peninsula was transformed by gene flow from North Africa and the eastern Mediterranean. They found that Greek and Roman settlements tended to be quite multiethnic, with individuals from the central and eastern Mediterranean and North Africa as well as locals, and that these interactions had lasting demographic as well as cultural impacts. “Beyond the specific insights about Iberia, this study serves as a model for how a high-resolution ancient DNA transect continuing into historical periods can be used to provide a detailed description of the formation of present-day populations,” explains Haak. “We hope that future use of similar strategies will provide equally valuable insights in other regions of the world.” ref
“An international team of researchers have analyzed ancient DNA from almost 300 individuals from the Iberian Peninsula, spanning more than 12,000 years, in two studies published concurrently in Current Biology and Science. The first study looked at hunter-gatherers and early farmers living in Iberia between 13,000 and 6000 years ago. The second looked at individuals from the region during all time periods over the last 8000 years. Together, the two papers greatly increase our knowledge about the population history of this unique region.” ref
“The Iberian Peninsula has long been thought of as an outlier in the population history of Europe, due to its unique climate and position on the far western edge of the continent. During the last Ice Age, Iberia remained relatively warm, allowing plants and animals – and possibly people – who were forced to retreat from much of the rest of Europe to continue living there. Similarly, over the last 8000 years, Iberia’s geographic location, rugged terrain, position on the Mediterranean coast and proximity to North Africa made it unique in comparison to other parts of Europe in its interactions with other regions. Two new studies, published concurrently in Current Biology and Science, analyze a total of almost 300 individuals who lived from about 13,000 to 400 years ago to give unprecedented clarity on the unique population history of the Iberian Peninsula.” ref
Iberian hunter-gatherers show two ancient Paleolithic lineages
“For the paper in Current Biology, led by researchers at the Max Planck Institute for the Science of Human History, researchers analyzed 11 hunter-gatherers and Neolithic individuals from Iberia. The oldest newly analyzed individuals are approximately 12,000 years old and were recovered from Balma Guilanyà in Spain. Earlier evidence had shown that, after the end of the last Ice Age, western and central Europe were dominated by hunter-gatherers with ancestry associated with an approximately 14,000-year-old individual from Villabruna, Italy. Italy is thought to have been a potential refuge for humans during the last Ice Age, like Iberia. The Villabruna-related ancestry largely replaced earlier ancestry in western and central Europe related to 19,000-15,000-year-old individuals associated with what is known as the Magdalenian cultural complex.” ref
“Interestingly, the findings of the current study show that both lineages were present in Iberian individuals dating back as far as 19,000 years ago. “We can confirm the survival of an additional Paleolithic lineage that dates back to the Late Ice Age in Iberia,” says Wolfgang Haak of the Max Planck Institute for the Science of Human History, senior author of the study. “This confirms the role of the Iberian Peninsula as a refuge during the Last Glacial Maximum, not only for fauna and flora but also for human populations.” This suggests that, far from being replaced by Villabruna-related individuals after the last Ice Age, hunter-gatherers in Iberia in fact already had ancestry from Magdalenian- and Villabruna-related sources. The discovery suggests an early connection between two potential refugia, resulting in a genetic ancestry that survived in later Iberian hunter-gatherers.” ref
Goyet-Q2 ancestry
“One of the most notable examples occurred during the Late Glacial, between the end of the LGM (~23,400 years ago) and the start of the Holocene epoch (~11,700 years ago). This shift is reflected in the ancestries associated with the ~15,090-year-old (IntCal20) Goyet Q2 individual, Belgium, and the ~14,010-year-old (IntCal20) Villabruna individual, Italy, in post-LGM Europe. We use these individuals as shorthand for the ancestries associated with them throughout the text. ‘Goyet Q2’ ancestry, which has previously been defined by the ~18,770-year-old (IntCal20) ‘El Mirón’ individual from Spain, has been identified in individuals associated with the Magdalenian culture, dating from ~20,500 to 14,000 years ago. This Goyet Q2/El Mirón ancestry has been suggested to represent a post-LGM expansion from southwestern European glacial refugia.” ref
“The ‘Villabruna’ ancestry, also broadly known as Western hunter-gatherers or WHG, consists of individuals dated from ~14,000 to 7,000 years ago associated with Epigravettian, Azilian/Federmesser, Epipalaeolithic, and Mesolithic cultures. The Villabruna ancestry is also associated with the observation that from ~14,000 years ago, all European individuals show some level of genetic affinity to present-day Near Eastern populations. The expansion in the geographic distribution of this ancestry also correlates with a period of rapid climate warming of the Late Glacial Interstadial (considered broadly equivalent to the onset of Greenland Interstadial 1 (GI-1), ~14,650 years ago) as well as cultural transitions from the Magdalenian/Late Upper Palaeolithic to the Azilian/Federmesser-Gruppen/Final Palaeolithic and has therefore been suggested to represent the movement of people into northwestern Europe after the LGM.” ref
“Interestingly, however, individuals with a mixture of Goyet Q2 and Villabruna ancestry appear in southern Europe from at least ~18,700 years ago—with the individual from El Mirón being the earliest identified thus far. The presence of individuals with admixed Goyet Q2 and Villabruna ancestry in southern Europe from the LGM onwards raises questions related to the fragmentation of populations into isolated refugia during the last Ice Age. It appears that both cultural and gene flow continued across the continent—although the nature of these processes and the mechanisms involved remain unclear. However, the presence of individuals with un-admixed Goyet Q2 ancestry in northern Europe until ~14,000 years ago also suggests some degree of sustained isolation throughout the LGM and into the Late Glacial. There is evidence of populations living in ice-marginal environments within northern Europe at the LGM and of long-distance movement of people from east to west north of the Alps, which has also been linked to the expansion of Magdalenian cultural groups. This evidence raises suggestions of Magdalenian populations with Goyet Q2 ancestry—who appear to have been cold-adapted hunter gatherers—retreating to northern Europe, perhaps due to climatic warming and the movement of prey species such as reindeer and horse. Conversely, more southerly regions such as northern Spain and Italy, where temperate prey species such as red deer persisted throughout the LGM and Late Glacial, may have provided greater ecological opportunities for population admixture.” ref
“Britain lies at the extreme northwest corner of the post-LGM expansion. With approximately two-thirds of the landmass covered by ice at the LGM and rapid deglaciation thereafter, substantial ecological and environmental change took place in the post-LGM landscape. As such, Britain offers a unique environmental context through which Late Upper Palaeolithic populations can be considered. By ~19,000 years ago, the British–Irish Ice Sheet was undergoing widespread melt, and by ~16,000 years ago, ice was absent from virtually all of England and Wales. Reindeer were present in southwest England by ~17,000 years ago, and habitats were dominated by open steppe–tundra vegetation. However, detailed consideration of Late Upper Palaeolithic sites in the United Kingdom and a series of radiocarbon dating programmes suggest that there is no evidence for post-LGM human recolonization of southwestern Britain before ~15,500 years ago.” ref
“As such, some regions of Britain were colonized before the rapid climate warming at the start of the Late Glacial Interstadial (~14,650 years ago). Accelerator mass spectrometry (AMS) dating indicates that Britain was probably recolonized at a slightly later date than adjacent regions such as the Paris Basin and the Belgian Ardennes—thereby suggesting an expansion of people across the European continent1. Interestingly, the British Magdalenian (known locally as the Creswellian) appears to be very similar (both in terms of chronology and cultural expression/typology) to the Classic Hamburgian, found in the northern Netherlands and the lowlands of northern Germany and Poland. However, understanding the expansion of post-LGM populations into and within the British Isles is hindered by a relative paucity of preserved archaeological remains suitable for dating. As such, the exact nature of human occupation of Late Upper Palaeolithic Britain remains unclear and we have relatively little knowledge of the earliest postglacial populations in Britain.” ref
“Whilst the genetics of Mesolithic, Neolithic, and Bronze Age individuals from Britain have recently been explored, no genetic data have yet been generated for British Palaeolithic individuals, due in part to the scarcity of human skeletal material available from Late Pleistocene Britain. To date, modern human skeletal remains have been recovered from only six Upper Palaeolithic sites. Nonetheless, these rare samples are crucial for our understanding of human populations across post-LGM Europe due to Britain’s location on the most northwesterly fringe of the European continent. Mesolithic British populations have been identified genetically as WHGs (Villabruna ancestry), indicating that this genetic ancestry spread to the most northwesterly area of early Holocene Europe by at least ~10,500 years ago.” ref
“What remains unclear, however, is when this ancestry first arrived in Britain and, additionally, what the genetic ancestry of Palaeolithic populations in Britain may have been. Given the previous association of Goyet Q2 ancestry with Magdalenian cultures across Europe and the similarities between the Creswellian and the Classic Hamburgian cultures, it could be hypothesized that British Late Upper Palaeolithic populations would also fall within the Goyet Q2 genetic cluster. To address these questions and expand our knowledge of the genetic makeup of Europe after the LGM, we investigate here the genetic characteristics of Late Upper Palaeolithic Britain through ancient DNA analyses of human remains from two archaeological sites in England and Wales.” ref
Anatolian hunter-gatherers
“Anatolian hunter-gatherer (AHG) is a distinct anatomically modern human archaeogenetic lineage, first identified in a 2019 study based on the remains of a single Epipaleolithic individual found in central Anatolia, radiocarbon dated to around 13,500 BCE. A population related to this individual was the main source of the ancestry of later Anatolian Neolithic Farmers (also known as Early European Farmers), who along with Western Hunter Gatherers (WHG) and Ancient North Eurasians (via Eastern Hunter Gatherers and or Western Steppe Herders) are one of the three currently known ancestral genetic contributors to present-day Europeans. The existence of this ancient population has been inferred through the genetic analysis of the remains of a man from the site of Pınarbaşı (37 ° 29’N, 33 ° 02’E), in central Anatolia, which has been dated at 13,642-13,073 cal BCE. This population is genetically differentiated from the rest of the known Pleistocene populations.” ref
“It has been discovered that populations of the Anatolian Neolithic (Anatolian Neolithic Farmers) derive most of their ancestry from the AHG, with minor gene flow from Iranian/Caucasus and Levantine sources, suggesting that agriculture was adopted in situ by these hunter-gatherers and not spread by demic diffusion into the region. The Anatolian hunter-gatherers began farming around 8300 BCE, at places such as Çayönü. Cows, sheep and goats may have been domesticated first in southern Turkey. These farmers moved into Thrace (now European Turkey) around 7000 BCE. At the autosomal level, in the Principal component analysis (PCA) the analyzed AHG individual turns out to be close to two later Anatolian populations, the Anatolian Aceramic Farmers (AAF) dating from 8300-7800 BCE, and the Anatolian Ceramic Farmers (ACF) dating from 7000-6000 BCE. The individual analyzed belongs to Y-chromosomal haplogroup C1a2 (C-V20), which has been found in some of the early WHGs, and mitochondrial haplogroup K2b. Both paternal and maternal lineages are rare in present-day Eurasian populations.” ref
“These early Anatolian farmers later replaced the European hunter-gatherer populations in Europe to a large extent, ultimately becoming the main genetic contribution to current European populations, especially those of the Mediterranean. In addition, their position in this analysis is intermediate between Natufian farmers and Western Hunter-Gatherers (WHG). This last point is confirmed by the ADMIXTURE and qp-Adm analysis and confirms the presence of hunter-gatherers of both European and Near-Eastern origins in Central Anatolia in the late Pleistocene. Mesolithic individuals from the Balkans, known as Iron Gates Hunter-Gatherers, are the most genetically similar group to the Anatolian Hunter-Gatherer lineage. Feldman et al. suggest that this affinity is not due to a genetic flow from the AHG to the ancestors of the Villabruna cluster, but on the contrary: there was a genetic flow from the ancestors of the Villabruna cluster to the ancestors of the AHG. The AHG diverged from Caucasus hunter-gatherer around 25,000 years ago.” ref
Iranian hunter-gatherers
“Despite the localization of the southern Caucasus at the outskirt of the Fertile Crescent, the Neolithisation process started there only at the beginning of the sixth millennium with the Shomutepe-Shulaveri culture of yet unclear origins. We present here genomic data for three new individuals from Mentesh Tepe in Azerbaijan, dating back to the beginnings of the Shomutepe-Shulaveri culture. Researchers have evidence that two juveniles, buried embracing each other, were brothers. We show that the Mentesh Tepe Neolithic population is the product of a recent gene flow between the Anatolian farmer-related population and the Caucasus/Iranian population, demonstrating that population admixture was at the core of the development of agriculture in the South Caucasus. By comparing Bronze Age individuals from the South Caucasus with Neolithic individuals from the same region, including Mentesh Tepe, we evidence that gene flows between Pontic Steppe populations and Mentesh Tepe-related groups contributed to the makeup of the Late Bronze Age and modern Caucasian populations.” ref
“The researcher’s results show that the high cultural diversity during the Neolithic period of the South Caucasus deserves close genetic analysis. “In the Near East, the Neolithic way of life emerged between 9000 and 7000 BCE. Several centers of Neolithisation have been identified, such as the Levant or Southern China, from which the agropastoral way of life diffused to other regions. The mechanisms of this diffusion have attracted tremendous attention for the last few decades. In some places, the Neolithic gained ground through the acculturation of local hunter-gatherers (for instance, in Anatolia or Iran); but in most regions (Europe, South-East Asia), farmer populations spread, and assimilation processes took place with a degree of admixture.” ref
“The agricultural transition profoundly changed human societies. We sequenced and analyzed the first genome (1.39x) of an early Neolithic woman from Ganj Dareh in the Zagros Mountains of Iran, a site with early evidence for an economy based on goat herding, ca. 10,000 years ago. We show that Western Iran was inhabited by a population genetically most similar to hunter-gatherers from the Caucasus, but distinct from the Neolithic Anatolian people who later brought food production into Europe. The inhabitants of Ganj Dareh made little direct genetic contribution to modern European populations, suggesting those of the Central Zagros were somewhat isolated from other populations of the Fertile Crescent. Runs of homozygosity are of a similar length to those from Neolithic farmers and shorter than those of Caucasus and Western Hunter-Gatherers, suggesting that the inhabitants of Ganj Dareh did not undergo the large population bottleneck suffered by their northern neighbors. While some degree of cultural diffusion between Anatolia, Western Iran, and other neighboring regions is possible, the genetic dissimilarity between early Anatolian farmers and the inhabitants of Ganj Dareh supports a model in which Neolithic societies in these areas were distinct.” ref
“The mechanism of Neolithisation in the South Caucasus, a region located between the Black and Caspian Seas on the southern slope of the Greater Caucasus Mountains, remains poorly understood. Mesolithic sites are known at Damjili Cave, unit 5 (Western Azerbaijan), Kmlo-2 Rock Shelter (Western Armenia), and Kotias Klde Cave (Western Georgia). Paleogenetic analyses of human bones excavated from the Kotias Klde Cave showed a genetic continuity with earlier Upper Palaeolithic (post-Last Glacial Maximum/LGM) sites but a discontinuity with pre-LGM individuals. Their genetic ancestry shares, to a certain extent, a common origin with ancient Iranian populations and differs from that of Anatolian and Levant hunter-gatherer groups, demonstrating a high genetic differentiation at this time between geographically close populations. The first settlements attributed to the Early Neolithic period belong to an aceramic culture, evidenced in several places in Central Georgia, as at Nagutni, in Western Georgia, as at Paluri, and in Western Azerbaijan at Damjili Cave, unit 4.” ref
“However, evidence of agriculture and herding remains scarce, suggesting that these sites represent a transitional phase between the Mesolithic and the Neolithic. In this context, the Shomutepe-Shulaveri culture (SSC) is the most ancient Caucasus culture with a complete Neolithic package. Found in several clusters of settlements in the northern foothills of the Lesser Caucasus, the SSC is characterized by circular mud-brick houses, domestic animals and cereals, handmade pottery, sometimes with incised and relief decoration, and obsidian and bone industries. Variants, such as the Aratashen/Aknashen culture (Ararat Plain), and other Neolithic contemporaneous cultures like the Kültepe Culture (Nakhchivan region) are also found in the South Caucasus. The slightly later Kamiltepe culture (Mil steppe culture), which probably includes the site of Polutepe, differs by its architecture, the use of flint tools instead of obsidian, and pottery-painted patterns that are rather related to Northern Iran and the Zagros.” ref
“The origins of the SSC are still discussed. Due to the rapid transition from the aceramic stage to the SSC, population continuity during the Neolithisation process is possible. However, several cultural and biological features are nonlocal. Domesticated animals, such as cattle, pigs, or goats, originate from Eastern Anatolia and from the Zagros mountains. Similarly, the glume wheat and barley recovered in the SSC sites have been domesticated elsewhere in the Middle East, even if the ancestors of the naked wheat Aegilops tauschii are found in the Caucasus and may have been local. The material culture and architecture evidence technical transfers with neighboring regions such as Southern Anatolia, Pre-Halafian, and Halafian culture of Northern Mesopotamia and Zagros11. Taken together, these data suggest a strong cultural connection, and maybe a degree of admixture, with other groups from the Fertile Crescent. Indeed, the genome-wide data for one individual coming from the same collective grave at Mentesh Tepe as the samples we analyze here, and confirmed by that of other individuals from Polutepe in the Mughan steppe or from Aknashen and Masis Blur in Armenia already showed that southern Caucasian groups are part of a cline that connected Eastern Anatolian and Zagros populations and evidence a gene flow that began around 6500 years BCE.” ref
“One of the oldest sites of the SSC, Mentesh Tepe, is located in the Tovuz district of western Azerbaijan and has been excavated between 2007 and 2015 (Fig. 1a). Several occupations were revealed, the earliest dating back to the Neolithic SSC period. At this time, the botanical assemblage is dominated by cereals, especially barley, naked wheat, and emmer, a common association during the Neolithic Southern Caucasus. Animal remains consist largely of domesticated ones (ovicaprines, cattle, pigs, dogs), and wild animals are rare. The diet of the Neolithic individuals from Mentesh Tepe relied mainly on C3-plants, such as wheat, barley, and lentils, with some evidence of freshwater fishes; the consumption of animal proteins varies between individuals. The pottery differs from classical SSC sites by being vegetal-tempered, a characteristic shared with Kamiltepe or the first occupation of the Nakhchivan site of Kültepe. As observed in many SSC sites, houses are circular and made of mudbricks, with or without the addition of straw or other organic material. Two SSC occupation phases are represented, separated in some places by a thick layer of ashes.” ref
“In a context where Neolithic burials are rare, Mentesh Tepe is exceptional for the discovery of a collective burial containing around 30 individuals (Fig. 1b), which is associated with the end of the first phase of frequentation of the site. Archaeoanthropological analysis has shown that it was a complex funerary gesture with mostly simultaneous deposits and, in contrast, some successive deposits that permitted manipulations on not completely decomposed bodies. The number of individuals in the burial, as well as their sex and age bias, suggest a dramatic event such as an epidemic, a famine, or a sudden episode, but no trace of violence has been evidenced on the bones. There is no specific orientation or position of the corpses, but some intentional arrangements are visible.” ref
“The most striking is formed by two juveniles embracing each other (Fig. 1c). Such an arrangement is rare, but other examples have been found in Neolithic and Protohistoric times, such as in Diyarbakir (Turkey, 6100 BCE) or Valdaro (Italy, 3000 BCE). Double burials are often considered as a lover’s embrace, but arguments for this explanation are often elusive. To better understand the origin of the Shomu-Shulaveri population and the structuration of this community, we performed paleogenetic studies of some of the individuals found in the collective burial. The genetic data obtained are then compared to those of another individual from the structure already published and to contemporaneous southwestern Asian genomes.” ref
Genetic structure of the Neolithic South-Caucasus
“The researchers merged our genome-wide data with the Human Origins dataset (HO-dataset), as well as with 3529 previously unrelated published ancient genomes (Supplementary Data 2). To decipher the genetic relations between the new Mentesh Tepe individuals and other ancient populations from the Caucasus, Anatolia, the Near East, and the Middle East, we performed: (1) a PCA on the modern dataset on which the ancient genomes have been projected and (2) an unsupervised ADMIXTURE analysis with the HO and the ancient dataset (Fig. 2a, b, Supplementary Fig. 2). The PCA shows that the Mentesh individuals overlap with some other previously published Neolithic or Chalcolithic individuals from the South Caucasus, but the individual from Aknashen falls a bit closer to CHG than the main neolithic cluster (Fig. 2a), and fall intermediate between the Iran Neolithic cluster and the Neolithic Anatolian Farmer group. The ADMIXTURE analysis suggests that the Mentesh individuals carry three main components: i.e., ca. 30% Neolithic Iran (Iran_N; green), 15% Levant Neolithic (PPN; pale rose) and 55% blue and pink components shared with Anatolian or European Neolithic populations (Fig. 2b).” ref
“Both analyses show that the new Mentesh Tepe individuals present a similar profile as that already published from individual MTT001 (despite different sequencing strategy) and from other Neolithic (Polutepe, Azerbaijan, Masis Blur and Aknashen, Armenia) or Chalcolithic (Alkhantepe, Azerbaijan) sites in the South Caucasus without true significant variations in percentage (Kruskal–Wallis test, p-value = 0.25), even though the Aknashen individual has the highest IranN/CHG percentage. They are also very similar to the Chalcolithic and Bronze Age Anatolian populations from Arslantepe and with other Chalcolithic and Bronze Age Anatolian populations but are quite distant from the Late Neolithic Anatolian Tell Kurdu individuals (five individuals identified as Tell_Kurdu_EC by the original publication, whom we refer as TellKurdu_LN) who instead clusters with the Neolithic Anatolian populations. Due to the lack of genomic data from South-eastern Mesopotamia, only the ancestry of groups that lived in North-eastern Mesopotamia during the PPN period and the Late Bronze Age could be considered. We observe that the South Caucasus population displays a different profile from the PPN Mesopotamian one, as it has more Anatolian affinity, but that it presents a profile similar to the LBA individual from North-eastern Mesopotamia (Nemrik9_LBA).” ref
“The agricultural transition started in a region comprising the Ancient Near East and Anatolia ~12,000 years ago with the first Pre-Pottery Neolithic villages and the first domestication of cereals and legumes. Archaeological evidence suggests a complex scenario of multiple domestications in a number of areas, coupled with examples of trade. Ancient DNA (aDNA) has revealed that this cultural package was later brought into Europe by dispersing farmers from Anatolia (so-called ‘demic’ diffusion, as opposed to non-demic cultural diffusion) ~8,400 years ago. However, a lack of aDNA from early Neolithic individuals from the Near East leaves a key question unanswered: was the agricultural transition developed by one major population group spanning the Near East, including Anatolia and the Central Zagros Mountains; or was the region inhabited by genetically diverse populations, as is suggested by the heterogeneous mode and timing of the appearance of early domesticates at different localities?” ref
“To answer this question, we sequenced the genome of an early Neolithic female from Ganj Dareh, GD13a, from the Central Zagros (Western Iran), dated to 10000-9700 years ago, a region located at the eastern edge of the Near East. Ganj Dareh is well known for providing the earliest evidence of herd management of goats beginning at 9,900 years ago. It is a classic mound site at an altitude of ~1400 m in the Gamas-Ab Valley of the High Zagros zone in Kermanshah Province, Western Iran. It was discovered in the 1960s during survey work and excavated over four seasons between 1967 and 1974. The mound, ~40 m in diameter, shows 7 to 8 m of early Neolithic cultural deposits. Five major levels were found, labeled A through E from top to bottom. Extended evidence showed a warren of rooms with evidence of under-floor inhumations within what may be burial chambers and/or disused houses. The current Minimum Number of Individuals is 116, with 56 cataloged as skeletons that had four or more bones recovered.” ref
“The individual analyzed here was part of burial 13, which contained three individuals, and was recovered in level C in 1971 from the floor of a brick-walled structure. The individual sampled, 13A (referred to as GD13a throughout the text), was a 30–50-year-old female; the other individuals in the burial unit were a second adult (13B) and an adolescent (13). The site is directly dated to 9650–9950 years ago, and it has been in intense occupation for over two to three centuries. The economy of the population was that of pastoralists with an emphasis on goat herding. Archaeobotanical evidence is limited12, but the evidence present is for two-row barley with no evidence for wheat, rye, or other domesticates. This implies that the overall economy was at a much earlier stage in the development of cereal agriculture than that found in the Levant, Anatolia, and Northern Mesopotamian basins.” ref
“The petrous bone of GD13a yielded sequencing libraries comprising 18.57% alignable human reads that were used to generate 1.39-fold genome coverage. The sequence data showed read lengths and nucleotide misincorporation patterns indicative of post-mortem damage, and had low contamination estimates (<1%, see Supplementary Fig. S3). The mitochondrion of GD13a (91.74X) was assigned to haplogroup X, most likely to the subhaplogroup X2, which has been associated with an early expansion from the Near East and has been found in early Neolithic samples from Anatolia, Hungary, and Germany.” ref
“We compared GD13a with a number of other ancient genomes and modern populations, using principal component analysis (PCA), ADMIXTURE, and outgroup f3 statistics. GD13a did not cluster with any other early Neolithic individual from Eurasia in any of the analyses. ADMIXTURE and outgroup f3 statistics identified Caucasus Hunter-Gatherers of Western Georgia, just north of the Zagros mountains, as the group genetically most similar to GD13a (Fig. 1B,C), whilst PCA also revealed some affinity with modern Central South Asian populations such as Balochi, Makrani and Brahui (Fig. 1A and Fig. S4). Also genetically close to GD13a were ancient samples from Steppe populations (Yamanya & Afanasievo) that were part of one or more Bronze age migrations into Europe, as well as early Bronze age cultures in that continent (Corded Ware), in line with previous relationships observed for the Caucasus Hunter-Gatherers.” ref
“The researchers further investigated the relationship between GD13a and Caucasus Hunter-Gatherers using D-statistics to test whether they formed a clade to the exclusion of other ancient and modern samples (Table S4). A large number of Western Eurasian samples (both modern and ancient) showed significant excess genetic affinity to the Caucasus Hunter-Gatherers, whilst none did with GD13a. Overall, these results point to GD13a having little direct genetic input into later European populations compared to its northern neighbors.” ref
“To better understand the history of the population to which GD13a belonged, we investigated the distribution of lengths of runs of homozygosity (ROH) (Fig. 2A). A bias towards a high frequency of both long and short ROH is indicative of past strong bottlenecks followed by population re-expansion. GD13a has a distribution with few long ROH (>2 Mb), similar to that of the descendants of Anatolian early farmers (represented by the European farmers NE1 and Stuttgart). In contrast, both Western and Caucasus Hunter-Gatherers have relatively more long as well as short ROH. Thus, GD13a is the descendant of a group that had relatively stable demography and did not suffer the bottlenecks that affected more northern populations.” ref
“The phenotypic attributes of GD13a are similar to the neighbouring Anatolian early farmers and Caucasus Hunter-Gatherers. Based on diagnostic SNPs, she had dark, black hair and brown eyes (see Supplementary). She lacked the derived variant (rs16891982) of the SLC45A2 gene associated with light skin pigmentation but likely had at least one copy of the derived SLC24A5 allele (rs1426654) associated with the same trait. The derived SLC24A5 variant has been found in both Neolithic farmer and Caucasus Hunter-Gatherer groups suggesting that it was already at appreciable frequency before these populations diverged. Finally, she did not have the most common European variant of the LCT gene (rs4988235) associated with the ability to digest raw milk, consistent with the later emergence of this adaptation.” ref
“It is possible that farmers related to GD13a contributed to the eastern diffusion of agriculture from the Near East that reached Turkmenistan by the 6th millennium years ago, and continued further east to the Indus Valley. However, detecting such a contribution is complicated by a later influx from Steppe populations with Caucasus Hunter-Gatherer ancestry during the Bronze Age. We tested whether the Western Eurasian component found in Indian populations can be better attributed to either of these two sources, GD13a and Kotias (a Caucasus Hunter Gatherer), using D-statistics to detect gene flow into an ancestral Indian component (represented by the Onge). Overall, for all tests where a difference could be detected, Kotias and GD13a were equally likely sources (Fig. S9 and Table S6). Whilst the attribution of part of the Western Eurasia component seen in India to Bronze Age migrations is supported by dating of last contact based on patterns of Linkage Disequilibrium, our analysis highlights the possibility that part of that component might derive from earlier contact during the eastern diffusion of agriculture.” ref
“GD13a had little direct genetic input into later European populations compared to the Caucasus Hunter-Gatherers (its northern neighbors), as demonstrated using D-statistics. This lack of connectivity with neighbouring regions might have arisen early on, since we also find that hunter-gatherers from the Caucasus show higher affinity to Western Hunter-Gatherers and early Anatolian farmers; this result suggests the possibility of gene flow between the former and these two latter groups to the exclusion of GD13a. An alternative, but not mutually exclusive, explanation for this pattern is that GD13a might have received genetic input from a source equally distant from all other European populations, and thus basal to them. The Last Glacial Maximum (LGM) made entire regions in northern Eurasia uninhabitable, and therefore a number of hunter-gatherer populations likely moved to the south. For Europe there may be a separation between Western and Eastern populations with minimal occupation of the Central European plains.” ref
“For Eastern Europe, Central Asia, and the northern Near East, glaciation itself was less a factor. In these areas, our understanding of how populations weathered the LGM is still vague at best. It has previously been suggested that differences in the frequency of long and short runs of homozygosity in ancient samples may be associated with different demographic experiences during the LGM. Neolithic farmers, with their lower frequency of short ROH, have been argued to have been relatively little affected by the LGM compared to Western and Caucasus Hunter-Gatherers which are characterised by more long ROH (>2 Mb). GD13a has a profile similar to that of the descendants of Anatolian farmers (i.e. early European farmers), suggesting that her ancestors also faced more benign conditions compared to populations further north. Superimposing the sampling locations of these groups onto climatic reconstructions from the LGM (Fig. 2B), however, does not reveal clear climatic differences among the regions. It is possible that the ancestors of the Anatolian and Ganj Dareh farmers spent the LGM in areas further south or east, which experienced milder climate.” ref
“But it is also possible that they exploited local pockets of favorable climate (refugia). Whilst high-elevation sites in the Zagros were abandoned during the LGM, there are a number of sites in the valleys that were occupied during that period and might have experienced more favorable conditions. The archaeological record indicates an eastward Neolithic expansion from the eastern regions of the Near East into Central and South Asia. Our analysis shows that both the Caucasus Hunter Gatherer Kotias and GD13a are plausible sources for the Eurasian Ancestry found in that part of Asia. Even though part of the Western Eurasian component found in India can be linked to Bronze Age migrations by dating the last contact using Linkage Disequilibrium (thus coming from the Kotias lineage), our results highlight the possibility of an older contribution from a source genetically close to GD13a (which would be hard to disentangle from the later gene flow from the Steppe).” ref
“Eventually, ancient DNA from the Indus Valley will be needed to detect conclusively whether any genetic traces were left by the eastward Neolithic expansion from the Near East, or whether this process was mostly cultural. The presence of two distinct lineages (Anatolian-like agriculturalists and Zagros mountain herders) in the Near East at the beginning of the Neolithic transition raises an interesting question regarding the independence of innovations arising at different locations. Even within the Central Zagros, economies vary greatly in their rate and pathway towards Neolithisation. Ganj Dareh, in the high Zagros, has the earliest known evidence for goat domestication, and the foothills of the Zagros mountains have also been argued to have been the site of early farming. In addition, early sites such as Sheikh-e Abad (11.650-9,600 years ago) provide evidence of early stages of barley cultivation. Were these innovations independent of similar achievements that made up the Neolithic package that North West Anatolians brought into Europe? Or were they exchanged culturally? If the latter, it would imply a cultural diffusion in the absence of much genetic interchange.” ref
“Populations of the Anatolian Neolithic derived most of their ancestry from the Anatolian hunter-gatherers (AHG), with a minor geneflow from Iranian/Caucasus and Levantine related sources, suggesting that agriculture was adopted in situ by these hunter-gatherers and not spread by demic diffusion into the region. Ancestors of AHGs and EEFs are believed to have split off from Western Hunter-Gatherers (WHGs) between 45,000 to 26,000 during the Last Glacial Maximum, and to have split from Caucasian Hunter-Gatherers (CHGs) between 25,000 to 14,000.” ref
“Genetic studies demonstrate that the introduction of farming to Europe in the 7th millennium BCE was associated with a mass migration of people from Northwest Anatolia to Southeast Europe, which resulted in the replacement of almost all (c. 98%) of the local Balkan hunter-gatherer gene pool with ancestry from Anatolian farmers. In the Balkans, the EEFs appear to have divided into two wings, who expanded further west into Europe along the Danube (Linear Pottery culture) or the western Mediterranean (Cardial Ware). Large parts of Northern Europe and Eastern Europe nevertheless remained unsettled by EEFs. During the Middle Neolithic there was a largely male-driven resurgence of WHG ancestry among many EEF-derived communities, leading to increasing frequencies of the hunter-gatherer paternal haplogroups among them.” ref
“Around 7,500 years ago. Iberian EEFs migrated into Northwest Africa, bringing farming to the region. The most common paternal haplogroup among EEFs was haplogroup G2a, while haplogroups E1b1 and R1b have also been found. Their maternal haplogroups consisted mainly of West Eurasian lineages including haplogroups H2, I, and T2, however significant numbers of central European farmers belonged to East Asian maternal lineage N9a, which is almost non-existent in modern Europeans, but common in East Asia. During the Chalcolithic and early Bronze Age, the EEF-derived cultures of Europe were overwhelmed by successive migrations of Western Steppe Herders (WSHs) from the Pontic–Caspian steppe, who carried roughly equal amounts of Eastern Hunter-Gatherer (EHG) and Caucasus Hunter-Gatherer (CHG) ancestries.” ref
“These migrations led to EEF paternal DNA lineages in Europe being almost entirely replaced with WSH-derived paternal DNA (mainly subclades of EHG-derived R1b and R1a). EEF maternal DNA (mainly haplogroup N) was also substantially replaced, being supplanted by steppe lineages, suggesting the migrations involved both males and females from the steppe. A 2017 study found that Bronze Age European with steppe ancestry had elevated EEF ancestry on the X chromosome, suggesting a sex bias, in which Steppe ancestry was inherited by more male than female ancestors. However, this study’s results could not be replicated in a follow-up study by Iosif Lazaridis and David Reich, suggesting that the authors had mis-measured the admixture proportions of their sample.” ref
“EEF ancestry remains widespread throughout Europe, ranging from about 60% near the Mediterranean Sea (with a peak of 65% in the island of Sardinia) and diminishing northwards to about 10% in northern Scandinavia. According to more recent studies the highest EEF ancestry found in modern Europeans ranges from 67% to over 80% in modern Sardinians, Italians, and Iberians, with the lowest EEF ancestry found in modern Europeans ranging around 35-40% in modern Finns, Lithuanians and Latvians. EEF ancestry is also prominent in living Northwest Africans like Moroccans and Algerians.” ref
Early European Farmers
“Early European Farmers (EEF) were a group of the Anatolian Neolithic Farmers (ANF) who brought agriculture to Europe and Northwest Africa. The Anatolian Neolithic Farmers were an ancestral component, first identified in farmers from Anatolia (also known as Asia Minor) in the Neolithic, and outside in Europe and Northwest Africa, they also existed in Iranian Plateau, South Caucasus, Mesopotamia, and Levant. Although the spread of agriculture from the Middle East to Europe has long been recognized through archaeology, it is only recent advances in archaeogenetics that have confirmed that this spread was strongly correlated with a migration of these farmers, and was not just a cultural exchange.” ref
“The earliest farmers in Anatolia derived most (80–90%) of their ancestry from the region’s local hunter-gatherers, with minor Levantine and Caucasus-related ancestry. The Early European Farmers moved into Europe from Anatolia through Southeast Europe from around 7,000 BCE, gradually spread north and westwards, and reached Northwest Africa via the Iberian Peninsula. Genetic studies have confirmed that the later Farmers of Europe generally have also a minor contribution from Western Hunter-Gatherers (WHGs), with significant regional variation. European farmer and hunter-gatherer populations coexisted and traded in some locales, although evidence suggests that the relationship was not always peaceful. Over the course of the next 4,000 years or so, Europe was transformed into agricultural communities, with WHGs being effectively replaced across Europe.” ref
“During the Chalcolithic and early Bronze Age, people who had Western Steppe Herder (WSH) ancestry moved into Europe and mingled with the EEF population; these WSH, originating from the Yamnaya culture of the Pontic steppe of Eastern Europe, probably spoke Indo-European languages. EEF ancestry is common in modern European and Northwest African populations, with EEF ancestry highest in Southern Europeans, especially Sardinians and Basque people. A distinct group of the Anatolian Neolithic Farmers spread into the east of Anatolia, and left a considerable genetic legacy in Iranian Plateau, South Caucasus, Levant (during the Pre-Pottery Neolithic B), and Mesopotamia. They also have a minor role in the ethnogenesis of WSHs of Yamnaya culture. The ANF ancestry is found in substantial levels in contemporary European, West Asian, and North African populations, and also found in Central and South Asian populations (through Bactria–Margiana Archaeological Complex and Corded Ware Culture) with more lower levels.” ref
“European hunter-gatherers were much taller than EEFs, and the replacement of European hunter-gatherers by EEFs resulted in a dramatic decrease in genetic height throughout Europe. During the later phases of the Neolithic, height increased among European farmers, probably due to increasing admixture with hunter-gatherers. During the Late Neolithic and Bronze Age, further reductions of EEF ancestry in Europe due to migrations of peoples with steppe-related ancestry is associated with further increases in height. High frequencies of EEF ancestry in Southern Europe might partly explain the shortness of Southern Europeans as compared to Northern Europeans, who carry increased levels of steppe-related ancestry. The Early European Farmers are believed to have been mostly dark-haired, dark-eyed, and light-skinned, although darker than most modern Europeans.” ref
“A study on different EEF remains throughout Europe concluded that they had “intermediate to light skin complexion.” A 2024 study found that risk alleles for mood-related phenotypes are enriched in the ancestry of Neolithic farmers. EEFs and their Anatolian forebears kept taurine cattle, pigs, sheep, and goats, as livestock, and planted cereal crops like wheat. Genetic analysis of individuals found in Neolithic tombs suggests that least some EEF peoples were patrilineal (tracing descent through the male line), with the tombs occupants mostly consisting of the male descendants of a single male common ancestor and their children, as well as their wives which were genetically unrelated to their husbands, suggesting female exogamy.” ref
To me, Animism starts in Southern Africa, then to West Europe, and becomes Totemism. Another split goes near the Russia and Siberia border becoming Shamanism, which heads into Central Europe meeting up with Totemism, which also had moved there, mixing the two which then heads to Lake Baikal in Siberia. From there this Shamanism-Totemism heads to Turkey where it becomes Paganism.
32,000-21,000 years ago, Yana Culture, at the Yana Woolly Rhinoceros Horn Site in Siberia, with genetic proximity to Ancient North Eurasian populations (Mal’ta, Afontova Gora), but also Ust-Ishim, Sunghir, and to a lesser extent Tianyuan, as well as similarities with the Clovis culture
ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref
“The genetic prehistory of humans in Asia, based on research using sequence data from humans who lived in Asia as early as 45,000 years ago. Genetic studies comparing present-day Australasians and Asians show that they likely derived from a single dispersal out of Africa, rapidly differentiating into three main lineages: one that persists partially in South Asia, one that is primarily found today in Australasia, and one that is widely represented across Siberia, East Asia, and Southeast Asia. Studies of ancient DNA from human remains in Asia dating from as far back as 45,000 years have greatly increased our understanding of the population dynamics leading to the current Asian populations.” ref
Ust’-Ishim man: Y-DNA haplogroupK2 and mt-DNA haplogroupR*
Tianyuan man: Y-DNA haplogroup K2b and mt-DNA haplogroup B
Yana Rhinoceros Horn Site: Y-DNA haplogroup P1 and mt-DNA haplogroup U
Sungir/Gravettian burials: Y-DNA haplogroup C1 and mt-DNA haplogroups U8c & U2
Ancient North Eurasians: Y-chromosome haplogroups P and its subclades R and Q and mt-DNA haplogroups U and R
Mal’ta–Buret’ culture: basalY-DNA haplogroup R* and mt-DNA haplogroup U
“MA-1 is the only known example of basal Y-DNA R* (R-M207*) – that is, the only member of haplogroup R* that did not belong to haplogroups R1, R2 or secondary subclades of these. The mitochondrial DNA of MA-1 belonged to an unresolved subclade of haplogroup U.” ref
“ANE ancestry has spread throughout Eurasia and the Americas in various migrations since the Upper Paleolithic, and more than half of the world’s population today derives between 5 and 42% of their genomes from the Ancient North Eurasians. Significant ANE ancestry can be found in Native Americans, as well as in Europe, South Asia, Central Asia, and Siberia. It has been suggested that their mythology may have featured narratives shared by both Indo-European and some Native American cultures, such as the existence of a metaphysical world tree and a fable in which a dog guards the path to the afterlife.” ref
Afontova Gora culture: Y-DNA haplogroup R and mt-DNA haplogroup R1b
“The bodies of two individuals, known as Afontova Gora 2 (AG2) and Afontova Gora 3 (AG3) were discovered within the complex (the name Afontova Gora 1 refers to the remains of a candid).” ref
Afontova Gora 2 “human remains”
“The human fossil remains of Afontova Gora 2 were discovered in the 1920s at Afontova Gora II and stored at the Hermitage Museum. The remains are dated to around 17,000 years ago (16,930-16,490 years ago). In 2009, researchers visited the Hermitage Museum and extracted DNA from the humerus of Afontova Gora 2. Despite significant contamination, researchers succeeded in extracting low coverage genomes. DNA analysis confirmed that the individual was male. The individual showed close genetic affinities to Mal’ta 1 (Mal’ta boy). Afontova Gora 2 also showed greater genetic affinity for the Karitiana people than for the Han Chinese. Around 1.9-2.7% of the genome was Neanderthal in origin.” ref
Afontova Gora 3 “human remains”
“In 2014, more human fossil remains were discovered at Afontova Gora II during salvage excavation before the construction of a new bridge over the Yenesei River. The remains belonged to two different females: the atlas of an adult female and the mandible and five lower teeth of a teenage girl (Afontova Gora 3), estimated to be around 14–15 years old. Initially, the new findings were presumed to be roughly contemporaneous with Afontova Gora 2. In 2017, direct AMS dating revealed that Afontova Gora 3 is dated to around 16,090 cal BCE. The mandible of Afontova Gora 3 was described as being gracile. Researchers analyzing the dental morphology of Afontova Gora 3 concluded that the teeth showed distinct characteristics with most similarities to another fossil (the Listvenka child) from the Altai-Sayan region and were neither western nor eastern. Afontova Gora 3 and Listvenka showed distinct dental characteristics that were also different from other Siberian fossils, including those from Mal’ta.” ref
“DNA was extracted from one of the teeth of Afontova Gora 3 and analyzed. Compared to Afontova Gora 2, researchers were able to obtain higher coverage genomes from Afontova Gora 3. DNA analysis confirmed that the individual was female. mtDNA analysis revealed that Afontova Gora 3 belonged to the mitochondrial Haplogroup R1b. Around 2.9-3.7% of the genome was Neanderthal in origin. In a 2016 study, researchers determined that Afontova Gora 2, Afontova Gora 3, and Mal’ta 1 (Mal’ta boy) shared common descent and were clustered together in a Mal’ta cluster. Genetically, Afontova Gora 3 is not closer to Afontova Gora 2 when compared to Mal’ta 1. When compared to Mal’ta 1, the Afontova Gora 3 lineage apparently contributed more to modern humans and is genetically closer to Native Americans.” ref
Afontova Gora 3 with Blond hair?
“Phenotypic analysis shows that Afontova Gora 3 carries the derived rs12821256 allele associated with, and likely causal for, blond hair color, making Afontova Gora 3 the earliest individual known to carry this derived allele. The allele was found in three later members of the largely ANE-derived Eastern Hunter-Gatherers populations from Samara, Motala and Ukraine c. 10,000 years ago, suggesting that it originated in the Ancient North Eurasian population before spreading to western Eurasia. The hundreds of millions of copies of this mutated alelle (a single-nucleotide polymorphism) are at the root of the classic European blond hair mutation, as massive population migrations from the Eurasian steppe, by a people who had substantial Ancient North Eurasian ancestry, entered continental Europe.” ref
The genetic proximity of Afontova Gora 3 with the Tarim mummies?
“A 2021 genetic study on the Tarim mummies found that they were primarily descended from a population represented by the Afontova Gora 3 specimen (AG3), genetically displaying “high affinity” with it. The genetic profile of the Afontova Gora 3 individual represented about 72% of the ancestry of the Tarim mummies, while the remaining 28% of their ancestry was derived from Baikal EBA (Early Bronze Age Baikal populations). The Tarim mummies are thus one of the rare Holocene populations who derive most of their ancestry from the Ancient North Eurasians (ANE, specifically the Mal’ta and Afontova Gora populations), despite their distance in time (around 14,000 years). More than any other ancient populations, they can be considered as “the best representatives” of the Ancient North Eurasians.” ref
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Here are my thoughts/speculations on where I believe is the possible origin of shamanism, which may have begun sometime around 35,000 to 30,000 years ago seen in the emergence of the Gravettian culture, just to outline his thinking, on what thousands of years later led to evolved Asian shamanism, in general, and thus WU shamanism as well. In both Europe-related “shamanism-possible burials” and in Gravettian mitochondrial DNA is a seeming connection to Haplogroup U. And the first believed Shaman proposed burial belonged to Eastern Gravettians/Pavlovian culture at Dolní Věstonice in southern Moravia in the Czech Republic, which is the oldest permanent human settlement that has ever been found. It is at Dolní Věstonice where approximately 27,000-25,000 years ago a seeming female shaman was buried and also there was an ivory totem portrait figure, seemingly of her.
And my thoughts on how cultural/ritual aspects were influenced in the area of Göbekli Tepe. I think it relates to a few different cultures starting in the area before the Neolithic. Two different groups of Siberians first from northwest Siberia with U6 haplogroup 40,000 to 30,000 or so. Then R Haplogroup (mainly haplogroup R1b but also some possible R1a both related to the Ancient North Eurasians). This second group added its “R1b” DNA of around 50% to the two cultures Natufian and Trialetian. To me, it is likely both of these cultures helped create Göbekli Tepe. Then I think the female art or graffiti seen at Göbekli Tepe to me possibly relates to the Epigravettians that made it into Turkey and have similar art in North Italy. I speculate that possibly the Totem pole figurines seen first at Kostenki, next went to Mal’ta in Siberia as seen in their figurines that also seem “Totem-pole-like”, and then with the migrations of R1a it may have inspired the Shigir idol in Russia and the migrations of R1b may have inspired Göbekli Tepe.
Göbekli Tepe Shamanism
From the journal Praehistorische Zeitschrift
“Abstract: The term shamanism is widely used in archaeology to describe early belief systems. Sometimes, this has taken the form of a one-size-fits-all-explanation, without a discussion of the concept or the cultural contexts it was applied to. Recently, the Early Neolithic (9600–7000 BCE) of southwestern Asia has become a focal point of this discussion. Sites like Nevalı Çori, Göbekli Tepe, Jerf el Ahmar, Körtik Tepe, Tell Abr’3, Tell Qaramel, Wadi Faynan 16, Karahantepe and Sayburç have produced rich evidence, mostly of an iconographical nature, that seems to offer direct insights into early belief systems. The current contribution uses one of the best-researched sites, Göbekli Tepe, as a case study to develop criteria for the identification of shamanism in the archaeological record.” ref
“R1b is the most common haplogroup in Western Europe, reaching over 80% of the population in Ireland, the Scottish Highlands, western Wales, the Atlantic fringe of France, the Basque country, and Catalonia. It is also common in Anatolia and around the Caucasus, in parts of Russia, and in Central and South Asia. Besides the Atlantic and North Sea coast of Europe, hotspots include the Po valley in north-central Italy (over 70%), Armenia (35%), the Bashkirs of the Urals region of Russia (50%), Turkmenistan (over 35%), the Hazara people of Afghanistan (35%), the Uyghurs of North-West China (20%) and the Newars of Nepal (11%). R1b-V88, a subclade specific to sub-Saharan Africa, is found in 60 to 95% of men in northern Cameroon.” ref
R1b Origins & History
R1b and Paleolithic mammoth hunters
“Haplogroup R* originated in North Asia just before the Last Glacial Maximum (26,500-19,000 years ago). This haplogroup has been identified in the remains of a 24,000 year-old boy from the Altai region, in south-central Siberia (Raghavan et al. 2013). This individual belonged to a tribe of mammoth hunters that may have roamed across Siberia and parts of Europe during the Paleolithic. Autosomally this Paleolithic population appears to have contributed mostly to the ancestry of modern Europeans and South Asians, the two regions where haplogroup R also happens to be the most common nowadays (R1b in Western Europe, R1a in Eastern Europe, Central and South Asia, and R2 in South Asia).” ref
“The oldest forms of R1b (M343, P25, L389) are found dispersed at very low frequencies from Western Europe to India, a vast region where could have roamed the nomadic R1b hunter-gatherers during the Ice Age. The three main branches of R1b1 (R1b1a, R1b1b, R1b1c) all seem to have stemmed from the Middle East. The southern branch, R1b1c (V88), is found mostly in the Levant and Africa. The northern branch, R1b1a (P297), seems to have originated around the Caucasus, eastern Anatolia or northern Mesopotamia, then to have crossed over the Caucasus, from where they would have invaded Europe and Central Asia. R1b1b (M335) has only been found in Anatolia.” ref
R1b and Neolithic cattle herders
“It has been hypothesised that R1b people (perhaps alongside neighboring J2 tribes) were the first to domesticate cattle in northern Mesopotamia some 10,500 years ago. R1b tribes descended from mammoth hunters, and when mammoths went extinct, they started hunting other large game such as bisons and aurochs. With the increase of the human population in the Fertile Crescent from the beginning of the Neolithic (starting 12,000 years ago), selective hunting and culling of herds started replacing the indiscriminate killing of wild animals. The increased involvement of humans in the life of aurochs, wild boars, and goats led to their progressive taming. Cattle herders probably maintained a nomadic or semi-nomadic existence, while other people in the Fertile Crescent (presumably represented by haplogroups E1b1b, G, and T) settled down to cultivate the land or keep smaller domesticates.” ref
“The analysis of bovine DNA has revealed that all the taurine cattle (Bos taurus) alive today descend from a population of only 80 aurochs. The earliest evidence of cattle domestication dates from circa 8,500 BCE in the Pre-Pottery Neolithic cultures in the Taurus Mountains. The two oldest archaeological sites showing signs of cattle domestication are the villages of Çayönü Tepesi in southeastern Turkey and Dja’de el-Mughara in northern Iraq, two sites only 250 km away from each others. This is presumably the area from which R1b lineages started expanding – or in other words the “original homeland” of R1b.” ref
“The early R1b cattle herders would have split in at least three groups. One branch (M335) remained in Anatolia, but judging from its extreme rarity today wasn’t very successful, perhaps due to the heavy competition with other Neolithic populations in Anatolia, or to the scarcity of pastures in this mountainous environment. A second branch migrated south to the Levant, where it became the V88 branch. Some of them searched for new lands south in Africa, first in Egypt, then colonizing most of northern Africa, from the Mediterranean coast to the Sahel.” ref
“The third branch (P297), crossed the Caucasus into the vast Pontic-Caspian Steppe, which provided ideal grazing grounds for cattle. They split into two factions: R1b1a1 (M73), which went east along the Caspian Sea to Central Asia, and R1b1a2 (M269), which at first remained in the North Caucasus and the Pontic Steppe between the Dnieper and the Volga. It is not yet clear whether M73 actually migrated across the Caucasus and reached Central Asia via Kazakhstan, or if it went south through Iran and Turkmenistan. In any case, M73 would be a pre-Indo-European branch of R1b, just like V88 and M335.” ref
“R1b-M269 (the most common form in Europe) is closely associated with the diffusion of Indo-European languages, as attested by its presence in all regions of the world where Indo-European languages were spoken in ancient times, from the Atlantic coast of Europe to the Indian subcontinent, which comprised almost all Europe (except Finland, Sardinia, and Bosnia-Herzegovina), Anatolia, Armenia, European Russia, southern Siberia, many pockets around Central Asia (notably in Xinjiang, Turkmenistan, Tajikistan, and Afghanistan), without forgetting Iran, Pakistan, northern India, and Nepal. The history of R1b and R1a are intricately connected to each others.” ref
The Levantine & African branch of R1b (V88)
“Like its northern counterpart (R1b-M269), R1b-V88 is associated with the domestication of cattle in northern Mesopotamia. Both branches of R1b probably split soon after cattle were domesticated, approximately 10,500 years ago (8,500 BCE). R1b-V88 migrated south towards the Levant and Egypt. The migration of R1b people can be followed archeologically through the presence of domesticated cattle, which appear in central Syria around 8,000-7,500 BCE (late Mureybet period), then in the Southern Levant and Egypt around 7,000-6,500 BCE (e.g. at Nabta Playa and Bir Kiseiba). Cattle herders subsequently spread across most of northern and eastern Africa. The Sahara desert would have been more humid during the Neolithic Subpluvial period (c. 7250-3250 BCE), and would have been a vast savannah full of grass, an ideal environment for cattle herding.” ref
“Evidence of cow herding during the Neolithic has shown up at Uan Muhuggiag in central Libya around 5500 BCE, at the Capeletti Cave in northern Algeria around 4500 BCE. But the most compelling evidence that R1b people related to modern Europeans once roamed the Sahara is to be found at Tassili n’Ajjer in southern Algeria, a site famous pyroglyphs (rock art) dating from the Neolithic era. Some painting dating from around 3000 BCE depict fair-skinned and blond or auburn haired women riding on cows. The oldest known R1b-V88 sample in Europe is a 6,200 year-old farmer/herder from Catalonia tested by Haak et al. (2015). Autosomally this individual was a typical Near Eastern farmer, possessing just a little bit of Mesolithic West European admixture.” ref
“After reaching the Maghreb, R1b-V88 cattle herders could have crossed the Strait of Gibraltar to Iberia, probably accompanied by G2 farmers, J1 and T1a goat herders. These North African Neolithic farmers/herders could have been the ones who established the Almagra Pottery culture in Andalusia in the 6th millennium BCE.” ref
“Nowadays small percentages (1 to 4%) of R1b-V88 are found in the Levant, among the Lebanese, the Druze, and the Jews, and almost in every country in Africa north of the equator. Higher frequency in Egypt (5%), among Berbers from the Egypt-Libya border (23%), among the Sudanese Copts (15%), the Hausa people of Sudan (40%), the the Fulani people of the Sahel (54% in Niger and Cameroon), and Chadic tribes of northern Nigeria and northern Cameroon (especially among the Kirdi), where it is observed at a frequency ranging from 30% to 95% of men.” ref
“According to Cruciani et al. (2010) R1b-V88 would have crossed the Sahara between 9,200 and 5,600 years ago, and is most probably associated with the diffusion of Chadic languages, a branch of the Afroasiatic languages. V88 would have migrated from Egypt to Sudan, then expanded along the Sahel until northern Cameroon and Nigeria. However, R1b-V88 is not only present among Chadic speakers, but also among Senegambian speakers (Fula-Hausa) and Semitic speakers (Berbers, Arabs).” ref
“R1b-V88 is found among the native populations of Rwanda, South Africa, Namibia, Angola, Congo, Gabon, Equatorial Guinea, Ivory Coast, Guinea-Bissau. The wide distribution of V88 in all parts of Africa, its incidence among herding tribes, and the coalescence age of the haplogroup all support a Neolithic dispersal. In any case, a later migration out of Egypt would be improbable since it would have brought haplogroups that came to Egypt during the Bronze Age, such as J1, J2, R1a, or R1b-L23. The maternal lineages associated with the spread of R1b-V88 in Africa are mtDNA haplogroups J1b, U5, and V, and perhaps also U3 and some H subclades (=> see Retracing the mtDNA haplogroups of the original R1b people).” ref
The North Caucasus and the Pontic-Caspian steppe : the Indo-European link
“Modern linguists have placed the Proto-Indo-European homeland in the Pontic-Caspian Steppe, a distinct geographic and archeological region extending from the Danube estuary to the Ural mountains to the east and North Caucasus to the south. The Neolithic, Eneolithic, and early Bronze Age cultures in Pontic-Caspian steppe has been called the Kurgan culture (4200-2200 BCE) by Marija Gimbutas, due to the lasting practice of burying the deads under mounds (“kurgan”) among the succession of cultures in that region. It is now known that kurgan-type burials only date from the 4th millenium BCE and almost certainly originated south of the Caucasus. The genetic diversity of R1b being greater around eastern Anatolia, it is hard to deny that R1b evolved there before entering the steppe world.” ref
“Horses were first domesticated around 4600 BCE in the Caspian Steppe, perhaps somewhere around the Don or the lower Volga, and soon became a defining element of steppe culture. Nevertheless, it is unlikely that R1b was already present in the eastern steppes at the time, so the domestication of the horse should be attributed to the indigenous R1a people, or tribes belonging to the older R1b-P297 branch, which settled in eastern Europe during the Late Paleolithic or Mesolithic period. Samples from Mesolithic Samara (Haak 2015) and Latvia (Jones 2017) all belonged to R1b-P297. Autosomally these Mesolithic R1a and R1b individuals were nearly pure Mesolithic East European, sometimes with a bit of Siberian admixture, but lacked the additional Caucasian admixture found in the Chalcolithic Afanasevo, Yamna and Corded Ware samples.” ref
“It is not yet entirely clear when R1b-M269 crossed over from the South Caucasus to the Pontic-Caspian steppe. This might have happened with the appearance of the Dnieper-Donets culture (c. 5100-4300 BCE). This was the first truly Neolithic society in the Pontic-Caspian Steppe. Domesticated animals (cattle, sheep, and goats) were herded throughout the steppes and funeral rituals were elaborate. Sheep wool would play an important role in Indo-European society, notably in the Celtic and Germanic (R1b branches of the Indo-Europeans) clothing traditions up to this day.” ref
“However, many elements indicate a continuity in the Dnieper-Donets culture with the previous Mesolithic hunter-gatherers, and at the same time an influence from the Balkans and Carpathians, with regular imports of pottery and copper objects. It is, therefore, more likely that Dnieper-Donets marked the transition of indigenous R1a and/or I2a1b people to early agriculture, perhaps with an influx of Near Eastern farmers from ‘Old Europe’. Over 30 DNA samples from Neolithic Ukraine (5500-4800 BCE) were tested by Mathieson et al. (2017).” ref
“They belonged to Y-haplogroups I, I2a2, R1a, R1b1a (L754), and one R1b1a2 (L388). None of them belonged to R1b-M269 or R1b-L23 clades, which dominated during the Yamna period. Mitochondrial lineages were also exclusively of Mesolithic European origin (U4a, U4b, U4d, U5a1, U5a2, U5b2, as well as one J2b1 and one U2e1). None of those maternal lineages include typical Indo-European haplogroups, like H2a1, H6, H8, H15, I1a1, J1b1a, W3, W4 or W5 that would later show up in the Yamna, Corded Ware, and Unetice cultures. Indeed, autosomally genomes from Neolithic Ukraine were purely Mesolithic European (about 90% EHG and 10% WHG) and completely lacked the Caucasian (CHG) admxiture later found in Yamna and subsequent Indo-European cultures during the Bronze Age.” ref
“The first clearly Proto-Indo-European cultures were the Khvalynsk (5200-4500 BCE) and Sredny Stog (4600-3900 BCE) cultures in the Pontic-Caspian Steppe. This is when small kurgan burials begin to appear, with the distinctive posturing of the dead on the back with knees raised and oriented toward the northeast, which would be found in later steppe cultures as well. There is evidence of population blending from the variety of skull shapes.” ref
“Towards the end of the 5th millennium, an elite starts to develop with cattle, horses, and copper used as status symbols. It is at the turn of the Khvalynsk and Sredny Stog periods that R1b-M269’s main subclade, L23, is thought to have appeared, around 4,500 BCE. 99% of Indo-European R1b descends from this L23 clade. The other branch descended from M269 is PF7562, which is found mostly in the Balkans, Turkey, and Armenia today, and may represent an early Steppe migration to the Balkans dating from the Sredny Stog period.” ref
“Another migration across the Caucasus happened shortly before 3700 BCE, when the Maykop culture, the world’s first Bronze Age society, suddenly materialized in the north-west Caucasus, apparently out of nowhere. The origins of Maykop are still uncertain, but archeologists have linked it to contemporary Chalcolithic cultures in Assyria and western Iran. Archeology also shows a clear diffusion of bronze working and kurgan-type burials from the Maykop culture to the Pontic Steppe, where the Yamna culture developed soon afterwards (from 3500 BCE). Kurgan (a.k.a. tumulus) burials would become a dominant feature of ancient Indo-European societies and were widely used by the Celts, Romans, Germanic tribes, and Scythians, among others.” ref
“The Yamna period (3500-2500 BCE) is the most important one in the creation of Indo-European culture and society. Middle Eastern R1b-M269 people had been living and blending to some extent with the local R1a foragers and herders for over a millennium, perhaps even two or three. The close cultural contact and interactions between R1a and R1b people all over the Pontic-Caspian Steppe resulted in the creation of a common vernacular, a new lingua franca, which linguists have called Proto-Indo-European (PIE). It is pointless to try to assign another region of origin to the PIE language. Linguistic similarities exist between PIE and Caucasian and Hurrian languages in the Middle East on the one hand, and Uralic languages in the Volga-Ural region on the other hand, which makes the Pontic Steppe the perfect intermediary region.” ref
“During the Yamna period, cattle and sheep herders adopted wagons to transport their food and tents, which allowed them to move deeper into the steppe, giving rise to a new mobile lifestyle that would eventually lead to the great Indo-European migrations. This type of mass migration in which whole tribes moved with the help of wagons was still common in Gaul at the time of Julius Caesar, and among Germanic peoples in the late Antiquity.” ref
The Yamna horizon was not a single, unified culture. In the south, along the northern shores of the Black Sea coast until the the north-west Caucasus, was a region of open steppe, expanding eastward until the Caspian Sea, Siberia, and Mongolia (the Eurasian Steppe). The western section, between the Don and Dniester Rivers (and later the Danube), was the one most densely settled by R1b people, with only a minority of R1a people (5-10%). The eastern section, in the Volga basin until the Ural mountains, was inhabited by R1a people with a substantial minority of R1b people (whose descendants can be found among the Bashkirs, Turkmans, Uyghurs, and Hazaras, among others).” ref
“The northern part of the Yamna horizon was forest-steppe occupied by R1a people, also joined by a small minority of R1b (judging from Corded Ware samples and from modern Russians and Belarussians, whose frequency of R1b is from seven to nine times lower than R1a). The western branch would migrate to the Balkans and Greece, then to Central and Western Europe, and back to their ancestral Anatolia in successive waves (Hittites, Phrygians, Armenians, etc.). The eastern branch would migrate to Central Asia, Xinjiang, Siberia, and South Asia (Iran, Pakistan, India). The northern branch would evolve into the Corded Ware culture and disperse around the Baltic, Poland, Germany, and Scandinavia.” ref
The Maykop culture, the R1b link to the Steppe?
“The Maykop culture (3700-2500 BCE) in the north-west Caucasus was culturally speaking a sort of southern extension of the Yamna horizon. Although not generally considered part of the Pontic-Caspian steppe culture due to its geography, the North Caucasus had close links with the steppes, as attested by numerous ceramics, gold, copper, and bronze weapons and jewelry in the contemporaneous cultures of Mikhaylovka, Sredny Stog, and Kemi Oba. The link between the northern Black Sea coast and the North Caucasus is older than the Maykop period. Its predecessor, the Svobodnoe culture (4400-3700 BCE), already had links to the Suvorovo-Novodanilovka and early Sredny Stog cultures. The even older Nalchik settlement (5000-4500 BCE) in the North Caucasus displayed a similar culture as Khvalynsk in the Caspian Steppe and Volga region. This may be the period when R1b started interracting and blending with the R1a population of the steppes.” ref
“The Yamna and Maykop people both used kurgan burials, placing their deads in a supine position with raised knees and oriented in a north-east/south-west axis. Graves were sprinkled with red ochre on the floor, and sacrificed domestic animal buried alongside humans. They also had in common horses, wagons, a heavily cattle-based economy with a minority of sheep kept for their wool, use of copper/bronze battle-axes (both hammer-axes and sleeved axes), and tanged daggers. In fact, the oldest wagons and bronze artifacts are found in the North Caucasus, and appear to have spread from there to the steppes.” ref
‘Maykop was an advanced Bronze Age culture, actually one of the very first to develop metalworking, and therefore metal weapons. The world’s oldest sword was found at a late Maykop grave in Klady kurgan 31. Its style is reminiscent of the long Celtic swords, though less elaborated. Horse bones and depictions of horses already appear in early Maykop graves, suggesting that the Maykop culture might have been founded by steppe people or by people who had close link with them. However, the presence of cultural elements radically different from the steppe culture in some sites could mean that Maykop had a hybrid population.” ref
“Without DNA testing it is impossible to say if these two populations were an Anatolian R1b group and a G2a Caucasian group, or whether R1a people had settled there too. The two or three ethnicities might even have cohabited side by side in different settlements. The one typical Caucasian Y-DNA lineage that does follow the pattern of Indo-European migrations is G2a-L13, which is found throughout Europe, Central Asia, and South Asia. In the Balkans, the Danube basin, and Central Europe its frequency is somewhat proportional to the percentage of R1b.” ref
“Maykop people are the ones credited for the introduction of primitive wheeled vehicles (wagons) from Mesopotamia to the Steppe. This would revolutionise the way of life in the steppe, and would later lead to the development of (horse-drawn) war chariots around 2000 BCE. Cavalry and chariots played an vital role in the subsequent Indo-European migrations, allowing them to move quickly and defeat easily anybody they encountered. Combined with advanced bronze weapons and their sea-based culture, the western branch (R1b) of the Indo-Europeans from the Black Sea shores are excellent candidates for being the mysterious Sea Peoples, who raided the eastern shores of the Mediterranean during the second millennium BCE.” ref
“The rise of the IE-speaking Hittites in Central Anatolia happened a few centuries after the disappearance of the Maykop and Yamna cultures. Considering that most Indo-European forms of R1b found in Anatolia today belong to the R1b-Z2103 subclade, it makes little doubt that the Hittites came to Anatolia via the Balkans, after Yamna/Maykop people invaded Southeast Europe. The Maykop and Yamna cultures were succeeded by the Srubna culture (1600-1200 BCE), possibly representing an advance of R1a-Z282 people from the northern steppes towards the Black Sea shores, filling the vacuum left by the R1b tribes who migrated to Southeast Europe and Anatolia.” ref
The Siberian & Central Asian branch
“When R1b crossed the Caucasus in the Late Neolithic, it split into two main groups. The western one (L51) would settle the eastern and northern of the Black Sea. The eastern one (Z2103) migrated to the Don-Volga region, where horses were domesticated circa 4600 BCE. R1b probably mixed with indigenous R1a people and founded the Repin culture (3700-3300 BCE) a bit before the Yamna culture came into existence in the western Pontic Steppe. R1b would then have migrated with horses along the Great Eurasian Steppe until the Altai mountains in East-Central Asia, where they established the Afanasevo culture (c. 3600-2400 BCE). Afanasevo people might be the precursors of the Tocharian branch of Indo-European languages. In 2014, Clément Hollard of Strasbourg University tested three Y-DNA samples from the Afanasevo culture and all three turned out to belong to haplogroup R1b, including two to R1b-M269.” ref
“The R1b people who stayed in the Volga-Ural region were probably the initiators of the Poltavka culture (2700-2100 BCE), then became integrated into the R1a-dominant Sintashta-Petrovka culture (2100-1750 BCE) linked to the Indo-Aryan conquest of Central and South Asia (=> see R1a for more details).” ref
“Nowadays in Russia R1b is found at higher frequencies among ethnic minorities of the Volga-Ural region (Udmurts, Komi, Mordvins, Tatars) than among Slavic Russians. R1b is also present in many Central Asian populations, the highest percentages being observed among the Uyghurs (20%) of Xinjiang in north-west China, the Yaghnobi people of Tajikistan (32%), and the Bashkirs (47%, or 62.5% in the Abzelilovsky district) of Bashkortostan in Russia (border of Kazakhstan).” ref
“R1b-M73, found primarily in North Asia (Altai, Mongolia), Central Asia, and the North Caucasus is thought to have spread during the Neolithic from the Middle East to Central and North Asia, and therefore can be considered to be pre-Indo-European.” ref
The European & Middle Eastern branch
“The Indo-Europeans’s bronze weapons and the extra mobility provided by horses would have given them a tremendous advantage over the autochthonous inhabitants of Europe, namely the native haplogroup C1a2, F and I (descendants of Cro-Magnon) and the early Neolithic herders and farmers (G2a, H2, E1b1b, and T1a). This allowed R1a and R1b to replace most of the native male lineages (=> see How did R1b come to replace most of the older lineages in Western Europe?), although female lineages seem to have been less affected.” ref
“A comparison with the Indo-Iranian invasion of South Asia shows that 40% of the male linages of northern India are R1a, but less than 10% of the female lineages could be of Indo-European origin. The impact of the Indo-Europeans was more severe in Europe because European society 4,000 years ago was less developed in terms of agriculture, technology (no bronze weapons), and population density than that of the Indus Valley civilization.” ref
“This is particularly true of the native Western European cultures where farming arrived much later than in the Balkans or Central Europe. Greece, the Balkans, and the Carpathians were the most advanced of European societies at the time and were the least affected in terms of haplogroup replacement. neolithic lineages survived better in regions that were more difficult to reach or less hospitable to horse breeders, like the Alps, the Dinaric Alps, the Apennines, and Sardinia.” ref
The Conquest of “Old Europe” and Central Europe (4200-2500 BCE)
“The first forays of Steppe people into the Balkans happened between 4200 BCE and 3900 BCE, when cattle herders equipped with horse-drawn wagons crossed the Dniester and Danube and apparently destroyed the towns of the Gumelnița, Varna and Karanovo VI cultures in Eastern Romania and Bulgaria. A climatic change resulting in colder winters during this exact period probably pushed steppe herders to seek milder pastures for their stock, while failed crops would have led to famine and internal disturbance within the Danubian and Balkanic communities. The ensuing Cernavodă culture (Copper Age, 4000-3200 BCE), Coțofeni/Usatovo culture (Copper to Bronze Age, 3500-2500 BCE), Ezero culture (Bronze Age, 3300-2700 BCE), in modern Romania, seems to have had a mixed population of steppe immigrants and people from the old tell settlements. These Steppe immigrants were likely a mixture of both R1a and R1b lineages, with a probably higher percentage of R1a than later Yamna-era invasions.” ref
“The Steppe invaders would have forced many Danubian farmers to migrate to the Cucuteni-Trypillian towns in the eastern Carpathians, causing a population boom and a north-eastward expansion until the Dnieper valley, bringing Y-haplogroups G2a, I2a1 (probably the dominant lineage of the Cucuteni-Trypillian culture), E1b1b, J2a, and T1a in what is now central Ukraine. This precocious Indo-European advance westward was fairly limited, due to the absence of Bronze weapons and an organized army at the time, and was indeed only possible thanks to climatic catastrophes which reduced the defenses of the towns of Old Europe. The Carphatian, Danubian, and Balkanic cultures were too densely populated and technologically advanced to allow for a massive migration.” ref
“In comparison, the forest-steppe R1a people successfully penetrated into the heart of Europe with little hindrance, due to the absence of developed agrarian societies around Poland and the Baltic. The Corded Ware culture (3200-1800 BCE) was a natural northern and western expansion of the Yamna culture, reaching as far west as Germany and as far north as Sweden and Norway. DNA analysis from the Corded Ware confirmed the presence of R1a and R1b in Poland c. 2700 BCE and R1a central Germany around 2600 BCE. The Corded Ware tribes expanded from the northern fringe of the Yamna culture where R1a lineages were prevalent over R1b ones.” ref
“The expansion of R1b people into Old Europe was slower, but proved inevitable. In 2800 BCE, by the time the Corded Ware had already reached Scandinavia, the Bronze Age R1b cultures had barely moved into the Pannonian Steppe. They established major settlements in the Great Hungarian Plain, the most similar habitat to their ancestral Pontic Steppes. Around 2500 BCE, the western branch of Indo-European R1b were poised for their next major expansion into modern Germany and Western Europe. By that time, the R1b immigrants had blended to a great extent with the indigenous Mesolithic and Neolithic populations of the Danubian basin, where they had now lived for 1,700 years.” ref
“The strongly partriarchal Indo-European elite remained almost exclusively R1b on the paternal side, but absorbed a high proportion of non-Indo-European maternal lineages. Hybridised, the new Proto-Indo-European R1b people would have lost most of their remaining Proto-Europoid or Mongolid features inherited from their Caspian origins (which were still clearly visible in numerous individuals from the Yamna period). Their light hair, eye, and skin pigmentation, once interbred with the darker inhabitants of Old Europe, became more like that of modern Southern Europeans.” ref
“The R1a people of the Corded Ware culture would come across far less populous societies in Northern Europe, mostly descended from the lighter Mesolithic population, and therefore retained more of their original pigmentation (although facial traits evolved considerably in Scandinavia, where the I1 inhabitants were strongly dolicocephalic and long-faced, as opposed to the brachycephalic and broad-faced Steppe people).” ref
The Conquest of Western Europe (2500-1200 BCE)
“The R1b conquest of Europe happened in two phases. For nearly two millennia, starting from circa 4200 BCE, Steppe people limited their conquest to the rich Chalcolithic civilizations of the Carpathians and the Balkans. These societies possessed the world’s largest towns, notably the tell settlements of the Cucuteni-Tripolye culture. Nothing incited the R1b conquerors to move further into Western Europe at such an early stage, because most of the land north and west of the Alps was still sparsely populated woodland. The Neolithic did not reach the British Isles and Scandinavia before circa 4000 BCE. Even northern France and most of the Alpine region had been farming or herding for less than a millennium and were still quite primitive compared to Southeast Europe and the Middle East.” ref
“North-west Europe remained a tribal society of hunter-gatherers practicing only limited agriculture for centuries after the conquest of the Balkans by the Indo-Europeans. Why would our R1b “conquistadors” leave the comfort of the wealthy and populous Danubian civilizations for the harsh living conditions that lie beyond? Bronze Age people coveted tin, copper, and gold, of which the Balkans had plenty, but that no one had yet discovered in Western Europe.” ref
“R1b-L51 is thought to have arrived in Central Europe (Hungary, Austria, Bohemia) around 2500 BCE, approximately two millennia after the shift to the Neolithic lifestyle in these regions. Agrarian towns had started to develop. Gold and copper had begun to be mined. The prospects of a conquest were now far more appealing.” ref
The archeological and genetic evidence (distribution of R1b subclades) point at several consecutive waves towards eastern and central Germany between 2800 BCE and 2300 BCE. The Unetice culture was probably the first culture in which R1b-L11 lineages played a major role. It is interesting to note that the Unetice period happen to correspond to the end of the Maykop (2500 BCE) and Kemi Oba (2200 BCE) cultures on the northern shores of the Black Sea, and their replacement by cultures descended from the northern steppes.” ref
“It can therefore be envisaged that the (mostly) R1b population from the northern half of the Black Sea migrated westward due to pressure from other Indo-European people (R1a) from the north, for example that of the burgeoning Proto-Indo-Iranian branch, linked to the contemporary Poltavka and Abashevo cultures.” ref
“It is doubtful that the Bell Beaker culture (2900-1800 BCE) in Western Europe was already Indo-European because its attributes are in perfect continuity with the native Megalithic cultures. The Beaker phenomenon started during the Late Neolithic and Early Chalcolithic in Portugal and propagated to the north-east towards Germany. During the same period Bronze Age Steppe cultures spread from Germany in the opposite direction towards Iberia, France, and Britain, progressively bringing R1b lineages into the Bell Beaker territory.” ref
“It is more likely that the beakers and horses found across Western Europe during that period were the result of trade with neighboring Indo-European cultures, including the first wave of R1b into Central Europe. It is equally possible that the Beaker people were R1b merchants or explorers who traveled across Western Europe and brought back tales of riches poorly defended by Stone Age people waiting to be to be conquered. This would have prompted a full-scale Indo-European (R1b) invasion from about 2500 BCE in Germany, reaching the Atlantic (north of the Pyrenees at least) around 2200 BCE.” ref
“Ancient DNA tests conducted by Lee et al. (2012), Haak et al. (2015), and Allentoft et al. (2015) have all confirmed the presence of R1b-L51 (and deeper subclades such as P312 and U152) in Germany from the Bell Beaker period onwards, but none in earlier cultures. German Bell Beaker R1b samples only had about 50% of Yamna autosomal DNA and often possessed Neolithic non-Steppe mtDNA, which confirms that R1b invaders took local wives as they advanced westward. Another study by Olalde et al. (2017) confirmed that Iberian Bell Beakers were genetically distinct from the previously tested German samples.” ref
“None of the Spanish or Portuguese individuals associated with Bell Beaker pottery possessed any Steppe admixture, and none belonged to the Indo-European haplogroup R1b-L23 or its subclades. Instead, they belonged to typical Megalithic lineages like G2a, I2a1, I2a2, and the Neolithic R1b-V88. The paper also confirmed a high frequency of R1b-L51 lineages in central Europe during the Beall Beaker period. In Britain, Megalithic individuals belonged exclusively to Y-haplogroup I2 (mostly I2a2 and I2a1b-L161), but were entirely replaced by R1b-L51 (mostly L21 clade) in the Early Bronze Age.” ref
“This means that the Bell Beaker culture was not associated with one particular ethnic group. Beaker pottery originated in Megalithic Iberia, but then spread to France and central Europe and was used by invading R1b-L51 Steppe people, who brought it with them to the British Isles, while wiping out most of the indigenous Megalithic population. There was, therefore, no ‘Bell Beaker people’, but just various populations trading and using Beaker pots during that period.” ref
“DNA samples from the Unetice culture (2300-1600 BCE) in Germany, which emerged less than two centuries after the appearance of the first R1b-L51 individuals in the late Bell Beaker Germany, had a slightly higher percentage of Yamna ancestry (60~65%) and of Yamna-related mtDNA lineages, which indicates a migration of both Steppe men and women. That would explain why archeological artifacts from the Unetice culture are clearly Yamna-related (i.e. Indo-European), as they abruptly introduced new technologies and a radically different lifestyle, while the Bell Beaker culture was in direct continuity with previous Neolithic or Chalcolithic cultures.” ref
“R1b men may simply have conquered the Bell Beaker people and overthrown the local rulers without obliterating the old culture due to their limited numbers. Taking the analogy of the Germanic migrations in the Late Antiquity, the R1b invasion of the Bell Beaker period was more alike to that of the Goths, Burgunds and Vandals, who all migrated in small numbers, created new kingdoms within the Roman empire, but adopted Latin language and Roman culture. In contrast, the Corded Ware and Unetice culture involved large-scale migrations of Steppe people, who imposed their Indo-European language and culture and conquered people, just like the Anglo-Saxons or the Bavarians did in the 5th century.” ref
“The cultures that succeeded to Unetice in Central Europe, chronologically the Tumulus culture (1600-1200 BCE), Urnfield culture (1300-1200 BCE), and Hallstatt culture (1200-750 BCE) cultures remained typically Indo-European. The Hallstatt culture, centered around the Alps, is considered the first classical Celtic culture in Europe. It quickly expanded to France, Britain, Iberia, northern Italy, and the Danube valley, probably spreading for the first time Celtic languages, although not bronze technology nor R1b lineages, which had both already spread over much of western Europe during the Bell Beaker period. => See also Metal-mining and stockbreeding explain R1b dominance in Atlantic fringe.” ref
“The linguistic gap between pre-IE vernaculars and IE languages was about as big as between modern English and Chinese. English, Greek, Russian, and Hindi are all related IE languages and therefore easier to learn for IE speakers than non-IE languages like Chinese, Arabic, or Hungarian. From a linguistic point of view, only a wide-scale migration of IE speakers could explain the thorough adoption of IE languages in Western Europe – leaving only Basque as a remnant of the Neolithic languages.” ref
“Besides pottery, archaeology provides ample evidence that the early Bronze Age in Central and Western Europe coincides with a radical shift in food production. Agriculture experiences an abrupt reduction in exchange for an increased emphasis on domesticates. This is also a period when horses become more common and cow milk is being consumed regularly. The overall change mimics the Steppic way of life almost perfectly. Even after the introduction of agriculture around 5200 BCE, the Bug-Dniester culture and later Steppe cultures were characterized by an economy dominated by herding, with only limited farming. This pattern expands into Europe exactly at the same time as bronze working.” ref
“Religious beliefs and arts undergo a complete reversal in Bronze Age Europe. Neolithic societies in the Near East and Europe had always worshipped female figurines as a form of fertility cult. The Steppe cultures, on the contrary, did not manufacture female figurines. As bronze technology spreads from the Danube valley to Western Europe, symbols of fertility and fecundity progressively disappear and are replaced by sculptures of domesticated animals.” ref
“Another clue that Indo-European Steppe people came in great number to Central and Western Europe is to be found in burial practices. Neolithic Europeans either cremated their dead (e.g. Cucuteni-Tripolye culture) or buried them in collective graves (this was the case of Megalithic cultures). In the Steppe, each person was buried individually, and high-ranking graves were placed in a funeral chamber and topped by a circular mound. The body was typically accompanied by weapons (maces, axes, daggers), horse bones, and a dismantled wagon (or later chariot).” ref
“These characteristic burial mounds are known as kurgans in the Pontic Steppe. Men were given more sumptuous tombs than women, even among children, and differences in hierarchy are obvious between burials. The Indo-Europeans had a strongly hierarchical and patrilinear society, as opposed to the more egalitarian and matrilinear cultures of Old Europe. The proliferation of ststus-conscious male-dominant kurgans (or tumulus) in Central Europe during the Bronze Age is a clear sign that the ruling elite had now become Indo-European.” ref
“The practice also spread to central Asia and southern Siberia, two regions where R1a and R1b lineages are found nowadays, just like in Central Europe. The ceremony of burial is one of the most emotionally charged and personal aspect of a culture. It is highly doubtful that people would change their ancestral practice “just to do like the neighbours”. In fact, different funerary practices have co-existed side by side during the European Neolithic and Chalcolithic. The ascendancy of yet another constituent of the Pontic Steppe culture in the rest of Europe, and in this case one that does not change easily through contact with neighbours, adds up to the likelihood of a strong Indo-European migration.” ref
“The adoption of some elements of a foreign culture tends to happen when one civilization overawes the adjacent cultures by its superiority. This process is called ‘acculturation’. However, there is nothing that indicates that the Steppe culture was so culturally superior as to motivate a whole continent, even Atlantic cultures over 2000 km away from the Pontic Steppe, to abandon so many fundamental symbols of their own ancestral culture, and even their own language. In fact, Old Europe was far more refined in its pottery and jewelry than the rough Steppe people. The Indo-European superiority was cultural but military, thanks to horses, bronze weapons, and an ethic code valuing individual heroic feats in war (these ethic values are known from the old IE texts, like the Rig Veda, Avesta, or the Mycenaean and Hittite literature).” ref
“After linguistics and archaeology, the third category of evidence comes from genetics itself. It had first been hypothesized that R1b was native to Western Europe, because this is where it was most prevalent. It has since been proven that R1b haplotypes displayed higher microsatellite diversity in Anatolia and in the Caucasus than in Europe. European subclades are also more recent than Middle Eastern or Central Asian ones. The main European subclade, R-P312/S116, only dates back to approximately 3500 to 3000 BCE.” ref
“It does not mean that the oldest common ancestor of this lineage arrived in Western Europe during this period, but that the first person who carried the mutation R-P312/S116 lived at least 5,000 years ago, assumably somewhere in the lower Danube valley or around the Black Sea. In any case, this timeframe is far too recent for a Paleolithic origin or a Neolithic arrival of R1b. The discovery of what was thought to be “European lineages” in Central Asia, Pakistan, and India hit the final nail on the coffin of a Paleolithic origin of R1b in Western Europe, and confirmed the Indo-European link.” ref
“All the elements concur in favor of a large scale migration of Indo-European speakers (possibly riding on horses) to Western Europe between 2500 to 2100 BCE, contributing to the replacement of the Neolithic or Chalcolithic lifestyle by an inherently new Bronze Age culture, with simpler pottery, less farming, more herding, new rituals (single graves) and new values (patrilinear society, warrior heroes) that did not evolve from local predecessors.” ref
Haplogroup migrations related to the Ancient North Eurasians: I added stuff to this map to help explain.
People reached Lake Baikal Siberia around 25,000 years ago. They (to Damien) were likely Animistic Shamanists who were also heavily totemistic as well. Being animistic thinkers they likely viewed amazing things in nature as a part of or related to something supernatural/spiritual (not just natural as explained by science): spirit-filled, a sprit-being relates to or with it, it is a sprit-being, it is a supernatural/spiritual creature, or it is a great spirit/tutelary deity/goddess-god. From there comes mythology and faith in things not seen but are believed to somehow relate or interact with this “real world” we know exists.
Both areas of Lake Baikal, one on the west side with Ancient North Eurasian culture and one on the east side with Ancient Northern East Asian culture (later to become: Ancient Northeast Asian culture) areas are the connected areas that (to Damien) are the origin ancestry religion area for many mythologies and religious ideas of the world by means of a few main migrations and many smaller ones leading to a distribution of religious ideas that even though are vast in distance are commonly related to and centering on Lake Baikal and its surrounding areas like the Amur region and Altai Mountains region.
To an Animistic Thinker: “Things are not just as they seem, they may have a spirit, or spirit energy relates to them”
To a Totemistic Thinker: “Things are not just as they seem, they may have a spirit, or spirit energy relates to them; they may have religio-cultural importance.”
“Ancient North Eurasian population had Haplogroups R, P, U, and Q DNA types: defined by maternal West-Eurasian ancestry components (such as mtDNA haplogroup U) and paternal East-Eurasian ancestry components (such as yDNA haplogroup P1 (R*/Q*).” ref
“Lighter skin and blond hair evolved in the Ancient North Eurasian (ANE) population. The SLC24A5 gene’s derived threonine or Ala111Thr allele (rs1426654) has been shown to be a major factor in the light skin tone of Europeans. Possibly originating as long as 19,000 years ago, it has been the subject of selection in the ancestors of Europeans as recently as within the last 5,000 years, and is fixed in modern European populations.” ref, ref
DNA-researcher: It’s not ‘woke’ to portray prehistoric Europeans with dark skin.
“It’s evolution. Ancient DNA analyses suggest that prehistoric Europeans looked different from modern Europeans today, but some people find that hard to accept. There was an artistic picture of an almost 6,000-year-old, girl who was walking along Lolland’s south coast and spits a piece of birch tar into the reeds. It didn’t taste great, but it helped to soothe her toothache. Fast forward 6,000 years, Danish archaeologists working on the Fehmarnbelt project stumble across the piece and recognize it for what it is: an almost 6,000-year-old piece of chewing gum. This ancient piece of gum is now on display at the Museum Lolland-Falster in southern Denmark among an amazing collection of Stone Age artifacts uncovered during the excavations. If you have not been, it is well worth a visit. In 2019, my research team at the University of Copenhagen managed something quite remarkable: We succeeded in extracting DNA from the gum and used it to reconstruct the girl’s entire genome — the first time anyone had sequenced an ancient human genome from anything other than skeletal remains. As the gum had been found on Lolland, we affectionately nicknamed her ‘Lola’.” ref
Stone-age girl in social media ‘shitstorm’
“The story of Lola and her chewing gum made headlines around the world when we published the genome in 2019 and then, suddenly, in the summer of 2023, Lola was back in the news, caught up in a media ‘shitstorm’. The ‘shitstorm’ first gathered pace on X, the platform formerly known as Twitter, and escalated to the point where the museum had to defend itself on national TV. Even the Danish newspaper ‘Ekstrabladet’ felt they had to comment and gave their opinion in a passionate editorial. So, what happened? These things are difficult to reconstruct, but evidently some people who had seen the image of Lola thought that she looked “way too dark” and accused us—and the museum—of ‘blackwashing’ the past. I suppose this episode says more about our own biases than anything else, and I would like to take this opportunity to explain why we portrayed Lola the way we did and what this tells us about the evolution of skin color in this part of the world.” ref
What we know about Lola
“First a disclaimer, we do not know exactly how old Lola was when she spat that chewing gum into the water. But based on her genome and other DNA trapped in the gum, we learned a lot of other things about her and her world. For example, we learned that she was a hunter-gatherer who lived off wild resources like fish, nuts, and wild game. At the time, small farming communities started to appear in other parts of Europe, but from what we can tell Lola and her kin still lived — as her ancestors had done for thousands of years before her — as hunter-gatherers. We also learned that she likely had dark skin, dark hair, and blue eyes. But how do we know that?” ref
The genetics of human skin pigmentation
“Skin color is a highly heritable and polygenic trait, meaning that it is influenced by multiple genes and their interactions with one another. One of the most well-known genes associated with skin pigmentation is the melanocortin 1 receptor gene (MC1R), but there are dozens more that have been reported to be involved in the pigmentation process. Most of these genes influence skin color by regulating the production of melanin, a dark pigment that protects from the deleterious effects of UV radiation. Basically, the more melanin you have in your skin, the darker it will be, and the more sun your skin can tolerate before you get sunburn. Eye and hair color are determined in a similar way, but the mechanisms that control the production of melanin in the eyes and hair are quite complex and independent processes. That is why it is possible to end up with different combinations of traits, such as the dark hair and blue eyes that are often seen in Europeans today, or the light hair and brown eyes that are common for Solomon Islanders, for example.” ref
How do we know what Lola looked like?
“Because the genes involved in pigmentation have been well studied, it is possible to predict the skin, eye, and hair color of an individual based on their genotype with a certain probability, something that is routinely done in forensic investigations. In practice, this works by checking which variants of a gene are present and what phenotype they are associated with. The more genes we can include in this analysis, the more confident we can be that our prediction is correct. In Lola’s case, we studied 41 gene variants across her genome that have been associated with skin, hair, and eye color in humans, and concluded that she likely had this unusual (at least for today) combination of dark skin, dark hair, and blue eyes.” ref
A common look in prehistoric Europe
“It is difficult to know exactly what people looked like 10,000 years ago. But based on ancient DNA studies, it appears that Lola’s ‘look’ was much more common in prehistoric Europe than it is today. Thanks to advances in ancient DNA sequencing, we now have the genomes of dozens of Upper Palaeolithic and Mesolithic (i.e. the period between around 50,000 and 5,000 years before present in Europe) individuals from Western Europe. And interestingly they all seem to lack the skin-lightening variants that are so common in Europeans today, indicating that they had dark skin. This is true for ‘Cheddar Man’ who lived around 10,000 years ago in southern England, as well as dozens of other Upper Palaeolithic and Mesolithic hunter-gatherer individuals from France, northern Italy, Spain, the Baltic, and other parts of Europe. Like skin color, eye color is also a fairly complex trait, involving the interaction of many different genes. Therefore, eye color is fairly difficult to predict, but it looks like Upper Palaeolithic and Mesolithic hunter-gatherers from Western Europe often had blue eyes, just like Lola. Overall, it looks like Lola’s phenotype—the combination of dark skin, dark hair, and blue eyes—was much more common in prehistoric Europe than it is today.” ref
How Europeans got their lighter skin
“So, why did people in prehistoric Europe look so different from northern Europeans today? The answer to this question lies in a complex interplay between our genes, our changing diets, population movements, and the environment. It has been theorized for some time that lighter skin emerged as an adaptive trait to light poor environments as it allows you to absorb sunlight more effectively, which is essential for the production of vitamin D. However, it was unclear when this happened. Early studies suggested that we first may have evolved lighter skin as our ancestors moved out of Africa and into Europe c. 50,000 years ago, but we now believe that this happened much later in European prehistory. In fact, there is evidence that lighter skin only evolved within the last 5,000 years or so, as a result of genetic admixture from Neolithic farming populations (who carried the skin-lightening variant) and strong selection favoring lighter skin.” ref
Our changing diet also played a part
“In addition, it looks like our changing diets also played a part. During most of European prehistory people relied on wild resources like nuts, game, and fish that are all rich in vitamin D, which is essential to our health. That changed dramatically during the Neolithic when people started to rely on a farmer’s diet that was rich in carbohydrates, but poor in vitamin D. Interestingly, this is exactly the period when we see lighter skin tones evolve in Western Europe and we think that the lack of vitamin D in the diet may have increased the selection pressures favouring lighter skin. All in all, there is solid evidence to suggest that lighter skin tones only evolved in Europe within the last 5,000 years or so, and that people who lived in Europe before then typically had darker skin. It is not that surprising, then, that Lola had darker skin. It simply reflects the fact that she lived at a time when Europeans had not yet evolved their lighter skin.” ref
Groups partially derived from the Ancient North Eurasians
“The ANE lineage is defined by association with the MA-1, or “Mal’ta boy”, remains of 24,000 years ago in central Siberia Mal’ta-Buret’ culture 24,000-15,000 years ago. The Ancient North Eurasians (ANE) samples (Afontova Gora 3, Mal’ta 1, and Yana-RHS) show evidence for minor gene flow from an East Asian-related group (simplified by the Amis, Han, or Tianyuan) but no evidence for ANE-related geneflow into East Asians (Amis, Han, Tianyuan), except the Ainu, of North Japan.” ref
“The ANE lineage is defined by association with the MA-1, or “Mal’ta boy”, remains of 24,000 years ago in central Siberia Mal’ta-Buret’ culture 24,000-15,000 years ago “basal to modern-day Europeans”. Some Ancient North Eurasians also carried East Asian populations, such as Tianyuan Man.” ref
“Bronze-age-steppe Yamnaya and Afanasevo cultures were ANE at around 50% and Eastern Hunter-Gatherer (EHG) at around 75% ANE. Karelia culture: Y-DNA R1a-M417 8,400 years ago, Y-DNA J, 7,200 years ago, and Samara, of Y-haplogroup R1b-P297 7,600 years ago is closely related to ANE from Afontova Gora, 18,000 years ago around the time of blond hair first seen there.” ref
Ancient North Eurasian
“In archaeogenetics, the term Ancient North Eurasian (often abbreviated as ANE) is the name given to an ancestral West Eurasian component that represents descent from the people similar to the Mal’ta–Buret’ culture and populations closely related to them, such as from Afontova Gora and the Yana Rhinoceros Horn Site. Significant ANE ancestry are found in some modern populations, including Europeans and Native Americans.” ref
“The ANE lineage is defined by association with the MA-1, or “Mal’ta boy“, the remains of an individual who lived during the Last Glacial Maximum, 24,000 years ago in central Siberia, Ancient North Eurasians are described as a lineage “which is deeply related to Paleolithic/Mesolithic hunter-gatherers in Europe,” meaning that they diverged from Paleolithic Europeans a long time ago.” ref
“The ANE population has also been described as having been “basal to modern-day Europeans” but not especially related to East Asians, and is suggested to have perhaps originated in Europe or Western Asia or the Eurasian Steppe of Central Asia. However, some samples associated with Ancient North Eurasians also carried ancestry from an ancient East Asian population, such as Tianyuan Man. Sikora et al. (2019) found that the Yana RHS sample (31,600 BP) in Northern Siberia “can be modeled as early West Eurasian with an approximately 22% contribution from early East Asians.” ref
“Populations genetically similar to MA-1 were an important genetic contributor to Native Americans, Europeans, Central Asians, South Asians, and some East Asian groups, in order of significance. Lazaridis et al. (2016:10) note “a cline of ANE ancestry across the east-west extent of Eurasia.” The ancient Bronze-age-steppe Yamnaya and Afanasevo cultures were found to have a noteworthy ANE component at ~50%.” ref
“According to Moreno-Mayar et al. 2018 between 14% and 38% of Native American ancestry may originate from gene flow from the Mal’ta–Buret’ people (ANE). This difference is caused by the penetration of posterior Siberian migrations into the Americas, with the lowest percentages of ANE ancestry found in Eskimos and Alaskan Natives, as these groups are the result of migrations into the Americas roughly 5,000 years ago.” ref
“Estimates for ANE ancestry among first wave Native Americans show higher percentages, such as 42% for those belonging to the Andean region in South America. The other gene flow in Native Americans (the remainder of their ancestry) was of East Asian origin. Gene sequencing of another south-central Siberian people (Afontova Gora-2) dating to approximately 17,000 years ago, revealed similar autosomal genetic signatures to that of Mal’ta boy-1, suggesting that the region was continuously occupied by humans throughout the Last Glacial Maximum.” ref
“The earliest known individual with a genetic mutation associated with blonde hair in modern Europeans is an Ancient North Eurasian female dating to around 16000 BCE from the Afontova Gora 3 site in Siberia. It has been suggested that their mythology may have included a narrative, found in both Indo-European and some Native American fables, in which a dog guards the path to the afterlife.” ref
“Genomic studies also indicate that the ANE component was introduced to Western Europe by people related to the Yamnaya culture, long after the Paleolithic. It is reported in modern-day Europeans (7%–25%), but not of Europeans before the Bronze Age. Additional ANE ancestry is found in European populations through paleolithic interactions with Eastern Hunter-Gatherers, which resulted in populations such as Scandinavian Hunter-Gatherers.” ref
“The Ancient North Eurasians (ANE) split from the ancestors of European peoples somewhere in the Middle East or South-central Asia, and used a northern dispersal route through Central Asia into Northern Asia and Siberia. Genetic analyses show that all ANE samples (Afontova Gora 3, Mal’ta 1, and Yana-RHS) show evidence for minor gene flow from an East Asian-related group (simplified by the Amis, Han, or Tianyuan). In contrast, no evidence for ANE-related geneflow into East Asians (Amis, Han, Tianyuan), except the Ainu, was found.” ref
“Genetic data suggests that the ANE formed during the Terminal Upper-Paleolithic (36+-1,5ka) period from a deeply European-related population, which was once widespread in Northern Eurasia, and from an early East Asian-related group, which migrated northwards into Central Asia and Siberia, merging with this deeply European-related population. These population dynamics and constant northwards geneflow of East Asian-related ancestry would later gave rise to the “Ancestral Native Americans” and Paleosiberians, which replaced the ANE as dominant population of Siberia.” ref
Groups partially derived from the Ancient North Eurasians
“Eastern Hunter-Gatherer (EHG) is a lineage derived predominantly (75%) from ANE. It is represented by two individuals from Karelia, one of Y-haplogroup R1a-M417, dated c. 8.4 kya, the other of Y-haplogroup J, dated c. 7.2 kya; and one individual from Samara, of Y-haplogroup R1b-P297, dated c. 7.6 kya. This lineage is closely related to the ANE sample from Afontova Gora, dated c. 18 kya. After the end of the Last Glacial Maximum, the Western Hunter-Gatherers (WHG) and EHG lineages merged in Eastern Europe, accounting for early presence of ANE-derived ancestry in Mesolithic Europe. Evidence suggests that as Ancient North Eurasians migrated West from Eastern Siberia, they absorbed Western Hunter-Gatherers and other West Eurasian populations as well.” ref
“Caucasian Hunter-Gatherer (CHG) is represented by the Satsurblia individual dated ~13 kya (from the Satsurblia cave in Georgia), and carried 36% ANE-derived admixture. While the rest of their ancestry is derived from the Dzudzuana cave individual dated ~26 kya, which lacked ANE-admixture, Dzudzuana affinity in the Caucasus decreased with the arrival of ANE at ~13 kya Satsurblia.” ref
“Scandinavian Hunter-Gatherer (SHG) is represented by several individuals buried at Motala, Sweden ca. 6000 BC. They were descended from Western Hunter-Gatherers who initially settled Scandinavia from the south, and later populations of EHG who entered Scandinavia from the north through the coast of Norway.” ref
“Iran Neolithic (Iran_N) individuals dated ~8.5 kya carried 50% ANE-derived admixture and 50% Dzudzuana-related admixture, marking them as different from other Near-Eastern and Anatolian Neolithics who didn’t have ANE admixture. Iran Neolithics were later replaced by Iran Chalcolithics, who were a mixture of Iran Neolithic and Near Eastern Levant Neolithic.” ref
“Ancient Beringian/Ancestral Native American are specific archaeogenetic lineages, based on the genome of an infant found at the Upward Sun River site (dubbed USR1), dated to 11,500 years ago. The AB lineage diverged from the Ancestral Native American (ANA) lineage about 20,000 years ago.” ref
“West Siberian Hunter-Gatherer (WSHG) are a specific archaeogenetic lineage, first reported in a genetic study published in Science in September 2019. WSGs were found to be of about 30% EHG ancestry, 50% ANE ancestry, and 20% to 38% East Asian ancestry.” ref
“Western Steppe Herders (WSH) is the name given to a distinct ancestral component that represents descent closely related to the Yamnaya culture of the Pontic–Caspian steppe. This ancestry is often referred to as Yamnaya ancestry or Steppe ancestry.” ref
“Late Upper Paeolithic Lake Baikal – Ust’Kyakhta-3 (UKY) 14,050-13,770 BP were mixture of 30% ANE ancestry and 70% East Asian ancestry.” ref
“Lake Baikal Holocene – Baikal Eneolithic (Baikal_EN) and Baikal Early Bronze Age (Baikal_EBA) derived 6.4% to 20.1% ancestry from ANE, while rest of their ancestry was derived from East Asians. Fofonovo_EN near by Lake Baikal were mixture of 12-17% ANE ancestry and 83-87% East Asian ancestry.” ref
“Hokkaido Jōmon people specifically refers to the Jōmon period population of Hokkaido in northernmost Japan. Though the Jōmon people themselves descended mainly from East Asian lineages, one study found an affinity between Hokkaido Jōmon with the Northern Eurasian Yana sample (an ANE-related group, related to Mal’ta), and suggest as an explanation the possibility of minor Yana gene flow into the Hokkaido Jōmon population (as well as other possibilities). A more recent study by Cooke et al. 2021, confirmed ANE-related geneflow among the Jōmon people, partially ancestral to the Ainu people. ANE ancestry among Jōmon people is estimated at 21%, however, there is a North to South cline within the Japanese archipelago, with the highest amount of ANE ancestry in Hokkaido and Tohoku.” ref
ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref
“A 2016 study found that the global maximum of Ancient North Eurasian (ANE) ancestry occurs in modern-day Kets, Mansi, Native Americans, and Selkups. ANE ancestry has spread throughout Eurasia and the Americas in various migrations since the Upper Paleolithic, and more than half of the world’s population today derives between 5 and 42% of their genomes from the Ancient North Eurasians. Significant ANE ancestry can be found in Native Americans, as well as in regions of northern Europe, South Asia, Central Asia, and Siberia. It has been suggested that their mythology may have featured narratives shared by both Indo-European and some Native American cultures, such as the existence of a metaphysical world tree and a fable in which a dog guards the path to the afterlife.” ref
Ancient Northern East Asian/ later became Ancient Northeast Asian
Ancient Paleo-Siberian
Mal’ta–Buret’ culture (Mal’ta boy MA-1)
The Kolyma Shaitans: Legends and Reality (I only use just a small part)
“A unique “shaitan” burial was discovered on the bank of Omuk-Kuel Lake in the Middle-Kolyma ulus in Yakutia. According to the legends, buried in it are mummified remains of a shaman woman who died during a devastating smallpox epidemics in the 18th c. In an attempt to overcome the deadly disease, the shaman’s relatives used her remains as an emeget fetish. The author believes that these legends reflect the real events of those far-away years. The Arabic word “shaitan” came to the Russian language from Turkic languages. According to Islamic tradition, a shaitan is a genie, an evil spirit, a demon. During Russian colonization and Christianization of Siberia, all sacred things used by the aborigines as fetishes, patron spirits of the family, and the tribe, grew to be called “shaitans.” There are various facts, dating to the 18th and 19th cc., confirming that this word also referred to the mummified remains of outstanding shamans.” ref
“In the 1740s, a member of the Second Kamchatka Expedition Yakov Lindenau wrote, “Meat is scratched off the [shaman’s] bones and the bones are put together to form a skeleton, which is dressed in human’s clothes and worshipped as a deity. The Yukagirs place such dressed bones…in their yurts, their number can sometimes reach 10 or 15. If somebody commits even a minor sacrilege with respect to these bones, he stirs up rancor on the part of the Yukagirs… While traveling and hunting, the Yukagirs carry these bones in their sledges, and moreover, in their best sledges pulled by their best deer. When the Yukagirs are going to undertake something really important, they tell fortune using these skeletons: lift a skeleton up, and if it seems light, it means that their enterprise will have a favorable outcome. The Yukagirs call these skeletons stariks (old men), endow them with their best furs, and sit them on beds covered with deer hides, in a circle, as though they are alive.” (Lindenau, 1983, p. 155)” ref
“In the late 19th c., a famous explorer of aboriginal culture V. I. Jochelson noted the changes that occurred in the ritual in the last century and a half. So, the Yukagirs divided among themselves the shaman’s meat dried in the sun and then put it in separate tents. The dead bodies of killed dogs were left there as well. “After that,” V. I. Jochelson writes, “they would divide the shaman’s bones, dry them and wrap in clothes. The skull was an object of worshipping. It was put on top of a trunk (body) cut out of wood. A caftan and two hats – a winter and a summer one – were sewn for the idol. The caftan was all embroidered. On the skull, a special mask was put, with holes for the eyes and the mouth… The figure was placed in the front corner of the home. Before a meal, a piece of food was thrown into the fire and the idol was held above it. This feeding of the idol… was committed before each meal.” (V. I. Jochelson, 2005, pp. 236—237)” ref
“The idol was kept by the children of the dead shaman. One of them was inducted into the shamanism mysteries while his father was still alive. The idol was carried in a wooden box. Sometimes, in line with the air burial ritual, the box was erected on poles or trees, and the idol was taken out only before hunting or a long journey so that the outcome of the enterprise planned could be predicted. With time, the Yukagirs began using wooden idols as charms. V. I. Jochelson notes that by the late 19th c. the Yukagirs had developed a skeptical attitude towards idols and referred to them as “shaitans.” In this way, under the influence of Christianity, the worshipped ancestor’s spirit turned into its opposite – an evil spirit, a devil, a Satan.” ref
Ancestral Native American, Ancient Beringian
14,000-year-old Ust-Kyakhta-3 (UKY) individual found near Lake Baikal
Chertovy Vorota Cave/Devil’s Gate Cave
Bactria–Margiana Archaeological Complex
32,000-21,000 years ago Yana Culture, at the Yana Woolly Rhinoceros Horn Site in Siberia, with genetic proximity to Ancient North Eurasian populations (Mal’ta and Afontova Gora), but also Ust-Ishim, Sunghir, and to a lesser extent Tianyuan, as well as similarities with the Clovis culture
Paleolithic to Bronze Age Siberians Reveal Connections with First Americans and across Eurasia
“An Upper Paleolithic Siberian shows a deep link with the First Peoples of the Americas. A 10,000-year continuum of Ancient North Eurasian ancestry in the Lake Baikal region. The Neolithic to Bronze Age population formation occurred through prolonged local admixture.” ref
“Modern humans have inhabited the Lake Baikal region since the Upper Paleolithic, though the precise history of its peoples over this long time span is still largely unknown. Here, we report genome-wide data from 19 Upper Paleolithic to Early Bronze Age individuals from this Siberian region. An Upper Paleolithic genome shows a direct link with the First Americans by sharing the admixed ancestry that gave rise to all non-Arctic Native Americans. We also demonstrate the formation of Early Neolithic and Bronze Age Baikal populations as the result of prolonged admixture throughout the eighth to sixth millennium years ago. Moreover, we detect genetic interactions with western Eurasian steppe populations and reconstruct Yersinia pestis genomes from two Early Bronze Age individuals without western Eurasian ancestry. Overall, our study demonstrates the most deeply divergent connection between Upper Paleolithic Siberians and the First Americans and reveals human and pathogen mobility across Eurasia during the Bronze Age.” ref
“The Lake Baikal region in Siberia has been inhabited by modern humans since the Upper Paleolithic and has a rich archaeological record. In the past 5 years, ancient genomic studies have revealed multiple genetic turnovers and admixture events in this region. The 24,000-year-old individual (MA1) from the Mal’ta site represents an ancestry referred to as “Ancient North Eurasian (ANE),” which was widespread across Siberia during the Paleolithic and that contributed to the genetic profile of a vast number of present-day Eurasian populations as well as Native Americans. ANE ancestry was suggested to have been largely replaced in the Lake Baikal region during the Early Neolithic by a gene pool related to present-day northeast Asians, with a limited resurgence of ANE ancestry by the Early Bronze Age.” ref
“Siberia has also been proposed as a source for multiple waves of dispersals into the Americas, the first of which was shown to be driven by a founding population estimated to have formed around 25,000–20,000 years ago. The so-called Ancient Beringian ancestry represented by a 11,500-year-old Alaskan individual (USR1) was shown to be part of this founding population, estimated to have split from other Native Americans around 23,000 years ago. In addition, the recently published 9,800-year-old Kolyma genome from northeastern Siberia was suggested to represent the closest relative to Native American populations outside of the Americas. Moreover, the Paleo-Eskimo ancestry represented by a 4,000-year-old Saqqaq individual from Greenland was also estimated to have split from northeastern Siberian groups and migrated to Arctic America around 6,000–5,000 years ago. Although these waves of migration are generally linked to ancient Siberian populations, their origins in the context of the Siberian genetic history remain poorly understood. Further studies of the Siberian population history using ancient genomes are, therefore, critical for the better understanding of the formation of Native American populations.” ref
“Furthermore, the Neolithic to Bronze Age transition in Eurasia was marked by complex cultural and genetic changes facilitated by extensive population movements, though their impact in the Lake Baikal region is still unclear. Looking to the west, the Early Bronze Age groups from the Pontic-Caspian steppe associated with the Yamnaya complex spread both east and west along with their distinct genetic profile often referred to as “Steppe ancestry”. The eastward expansion of this group is considered to be associated with the Early Bronze Age Afanasievo culture. However, the later Middle Bronze Age Okunevo-related population from the central steppe, as well as the Late Bronze Age Khövsgöl-related population from the eastern steppe, harbor only a limited proportion of Steppe ancestry. Therefore, the effect of steppe migrations in eastern Eurasia, particularly the interactions of Bronze Age Baikal hunter-gatherers with the contemporaneous and geographically proximal Afanasievo population, is still largely unexplored.” ref
“In this study, researchers report 19 newly sequenced ancient hunter-gatherers from the Lake Baikal and its surrounding regions, spanning from the Upper Paleolithic to the Early Bronze Age. Their analyses alongside published data reveal the most deeply divergent ancestry that link Upper Paleolithic Siberians and the First Peoples of the Americas, and more clearly delineate the complex transition between Early Neolithic and Early Bronze Age populations in the Lake Baikal region. We also provide both human and pathogen genomic evidence demonstrating the influence of western Eurasian steppe populations in this region during the Early Bronze Age and discuss the genetic contribution of Lake Baikal hunter-gatherers to Siberian populations through time.” ref
“Most of the Lake Baikal individuals occupied the space on a “ANE-NEA” cline running between “Northeast Asian” (NEA) ancestry represented by Neolithic hunter-gathers from the Devil’s Gate in the Russian Far East, and the ANE ancestry represented by Upper Paleolithic Siberian individuals MA1, AfontovaGora 2 (AG2), and AfontovaGora 3 (AG3), which was first described by. Our newly sequenced Upper Paleolithic genome from the Ust-Kyakhta-3 site (UKY) just south to the Lake Baikal is placed close to the Mesolithic northeastern Siberian Kolyma individual and is shifted toward Native American populations compared to the rest of the ancient Baikal individuals along PC2. All four Early Neolithic individuals cluster with published Early Neolithic groups from the same region (Shamanka_EN, Lokomotiv_EN, UstBelaya_EN) designated as the “Baikal_EN” population. The LNBA individuals were divided into four groups. The major “Baikal_LNBA” group included 10 individuals and clustered with published Late Neolithic to Bronze Age Baikal populations (Shamanka_EBA, Kurma_EBA, UstIda_EBA, UstIda_LN, UstBelaya_BA).” ref
“These individuals were positioned in PCA closer to ANE-related individuals compared with the Early Neolithic individuals from the same region, as well as closer to the Paleo-Eskimo Saqqaq individual. Another two individuals (GLZ001 and GLZ002) from the Glazkovskoe predmestie site, unlike the third individual from the same archaeological site (GLZ003), seemed shifted from the main cluster and showed closer genetic affinity to the Devil’s Gate and Early Neolithic Baikal individuals. One of the six individuals from the Kachug site (KPT005) was substantially displaced from the Baikal_LNBA group toward western Eurasians along PC1, not along the ANE-NEA cline but toward later Bronze Age populations, suggesting a potential introgression of the Steppe-related ancestry. Finally, an Early Bronze Age individual (BZK002) from the Bazaikha site in the Yenisei River region further to the west of the Lake Baikal was significantly displaced toward ANE-related individuals and located close to published Bronze Age individuals associated to the Okunevo culture.” ref
“Population clustering with ADMIXTURE based on worldwide populations also showed a similar clustering pattern. When selecting a K value of 16, the published and newly sequenced individuals belonging to main Early Neolithic to Bronze Age Baikal groups all showed genetic profiles composed of a mixture of three major components that were mostly enriched in ANE-related individuals, northeast Asians, and central Siberians represented by the Uralic-speaking Nganasan population. The ANE and central Siberian ancestries were both of higher proportion in most LNBA Baikal individuals than in the Early Neolithic ones, while GLZ001 and GLZ002 showed higher NEA ancestry, similar to the Early Neolithic population. The BZK002 individual presented a profile similar to the published Okunevo group, with a much larger ANE component compared to other Lake Baikal individuals. The KPT005 individual also displayed a substantial contribution derived from European “Western Hunter-Gatherer” (WHG) ancestry, likely acquired through gene flow from the west.” ref
“Researchers estimated the runs of homozygosity (ROH) of selected individuals together with published Baikal individuals and did not identify an inbreeding signal in any individual. The Kolyma individual showed significantly more ROH compared with other individuals, suggesting a smaller population size in Mesolithic northeastern Siberia. The sharing of identity-by-descent (IBD) segments between individuals suggested a close relationship between UKY and Kolyma, supporting our analyses based on genome-wide SNP data, and also revealed that Early Neolithic and LNBA Baikal individuals shared genetic affinity with each other as well as with the older UKY and Kolyma genomes.” ref
World’s oldest known fort was constructed by hunter-gatherers 8,000 years ago in Siberia
“The fact that this Stone Age fort was built by hunter-gatherers is transforming our understanding of ancient human societies. Hunter-gatherers built the oldest known fort in the world about 8,000 years ago in Siberia, a new study finds. Archaeologists have long associated fortresses with permanent agricultural settlements. However, this cluster of fortified structures reveals that prehistoric groups were constructing protective edifices much earlier than originally thought.” ref
“These hunter-gatherers “defy conventional stereotypes that depict such societies as basic and nomadic, unveiling their capacity to construct intricate structures,” study co-author Tanja Schreiber, an archaeologist at Free University of Berlin, told Live Science in an email. Located along the Amnya River in western Siberia, remains of the Amnya fort include roughly 20 pit-house depressions scattered across the site, which is divided into two sections: Amnya I and Amnya II. Radiocarbon dating confirmed that the settlement was first inhabited during the Mesolithic, or Middle Stone Age, according to the study. When constructed, each pit house would have been protected by earthen walls and wooden palisades — two construction elements that suggest “advanced agricultural and defensive capabilities” by the inhabitants, the archaeologists said in a statement.” ref
“One of the Amnya fort’s most astonishing aspects is the discovery that approximately 8,000 years ago, hunter-gatherers in the Siberian Taiga built intricate defense structures,” Schreiber said. “This challenges traditional assumptions that monumental constructions were solely the work of agricultural communities.” It’s unknown what triggered the need for these fortified structures in the first place, but the strategic location overlooking the river would have not only been an ideal lookout point for potential threats but also allowed hunter-gatherers to keep tabs on their fishing and hunting grounds, the researchers noted.” ref
“The Samara culture was an Eneolithic (Copper Age) culture that flourished around the turn of the 5th millennium BCE, at the Samara Bend of the Volga River (modern Russia). The Samara culture is regarded as related to contemporaneous or subsequent prehistoric cultures of the Pontic–Caspian steppe, such as the Khvalynsk, Repin, and Yamna (or Yamnaya) cultures.” ref
“Genetic analyses of a male buried at Lebyazhinka, radiocarbon dated to 5640-5555 BCE, found that he belonged to a population often referred to as “Samara hunter-gatherers”, a group closely associated with Eastern Hunter-Gatherers. The male sample carried Y-haplogroup R1b1a1a and mitochondrial haplogroup U5a1d.” ref
“Pottery consists mainly of egg-shaped beakers with pronounced rims. They were not able to stand on a flat surface, suggesting that some method of supporting or carrying must have been in use, perhaps basketry or slings, for which the rims would have been a useful point of support. The carrier slung the pots over the shoulder or onto an animal. The decoration consists of circumferential motifs: lines, bands, zig-zags, or wavy lines, incised, stabbed, or impressed with a comb. These patterns are best understood when seen from the top. They appear then to be a solar motif, with the mouth of the pot as the sun. Later developments of this theme show that in fact the sun is being represented.” ref
“The culture is characterized by the remains of animal sacrifice, which occur over most of the sites. There is no indisputable evidence of riding, but there were horse burials, the earliest in the Old World. Typically the head and hooves of cattle, sheep, and horses are placed in shallow bowls over the human grave, smothered with ochre. Some have seen the beginning of the horse sacrifice in these remains, but this interpretation has not been more definitely substantiated. We know that the Indo-Europeans sacrificed both animals and people, like many other cultures.” ref
“The graves found are shallow pits for single individuals, but two or three individuals might be placed there. Some of the graves are covered with a stone cairn or a low earthen mound, the very first predecessor of the kurgan. The later, fully developed kurgan was a hill on which the deceased chief might ascend to the sky god, but whether these early mounds had that significance is doubtful.” ref
“Grave offerings included ornaments depicting horses. The graves also had an overburden of horse remains; it cannot yet be determined decisively if these horses were domesticated and ridden or not, but they were certainly used as a meat-animal. Most controversial are bone plaques of horses or double oxen heads, which were pierced. The graves yield well-made daggers of flint and bone, placed at the arm or head of the deceased, one in the grave of a small boy. Weapons in the graves of children are common later. Other weapons are bone spearheads and flint arrowheads. Other carved bone figurines and pendants were found in the graves.” ref
“The Yamnaya culture or the Yamna culture, also known as the Pit Grave culture or Ochre Grave culture, was a late Copper Age to early Bronze Age archaeological culture of the region between the Southern Bug, Dniester, and Ural rivers (the Pontic–Caspian steppe), dating to 3300–2600 BCE or around 5,300 to 4,600 years ago. It was discovered by Vasily Gorodtsov following his archaeological excavations near the Donets River in 1901–1903. Its name derives from its characteristic burial tradition: Я́мная (romanization: yamnaya) is a Russian adjective that means ‘related to pits (yama)’, as these people used to bury their dead in tumuli (kurgans) containing simple pit chambers.” ref
“The Yamnaya economy was based upon animal husbandry, fishing, and foraging, and the manufacture of ceramics, tools, and weapons. The people of the Yamnaya culture lived primarily as nomads, with a chiefdom system and wheeled carts and wagons that allowed them to manage large herds. They are also closely connected to Final Neolithic cultures, which later spread throughout Europe and Central Asia, especially the Corded Ware people and the Bell Beaker culture, as well as the peoples of the Sintashta, Andronovo, and Srubnaya cultures.” ref
“Back migration from Corded Ware also contributed to Sintashta and Andronovo. In these groups, several aspects of the Yamnaya culture are present. Yamnaya material culture was very similar to the Afanasevo culture of South Siberia, and the populations of both cultures are genetically indistinguishable. This suggests that the Afanasevo culture may have originated from the migration of Yamnaya groups to the Altai region or, alternatively, that both cultures developed from an earlier shared cultural source.” ref
“Genetic studies have suggested that the people of the Yamnaya culture can be modelled as a genetic admixture between a population related to Eastern European Hunter-Gatherers (EHG) and people related to hunter-gatherers from the Caucasus (CHG) in roughly equal proportions, an ancestral component which is often named “Steppe ancestry”, with additional admixture from Anatolian, Levantine, or Early European farmers. Genetic studies also indicate that populations associated with the Corded Ware, Bell Beaker, Sintashta, and Andronovo cultures derived large parts of their ancestry from the Yamnaya or a closely related population.” ref
“The origin of the Yamnaya culture continues to be debated, with proposals for its origins pointing to both the Khvalynsk and Sredny Stog cultures. The Khvalynsk culture (4700–3800 BCE) (middle Volga) and the Don-based Repin culture (c. 3950–3300 BCE) in the eastern Pontic-Caspian steppe, and the closely related Sredny Stog culture (c. 4500–3500 BCE) in the western Pontic-Caspian steppe, preceded the Yamnaya culture (3300–2500 BCE). The Yamnaya culture was succeeded in its western range by the Catacomb culture (2800–2200 BCE); in the east, by the Poltavka culture (2700–2100 BCE) at the middle Volga. These two cultures were followed by the Srubnaya culture (18th–12th century BCE).” ref
“Further efforts to pinpoint the location came from Anthony (2007), who suggested that the Yamnaya culture (3300–2600 BCE) originated in the Don–Volga area at c. 3400 BCE, preceded by the middle Volga-based Khvalynsk culture and the Don-based Repin culture (c. 3950–3300 BCE), arguing that late pottery from these two cultures can barely be distinguished from early Yamnaya pottery. Earlier continuity from eneolithic but largely hunter-gatherer Samara culture and influences from the more agricultural Dnieper–Donets II are apparent.” ref
“He argues that the early Yamnaya horizon spread quickly across the Pontic–Caspian steppes between c. 3400 and 3200 BCE:
The spread of the Yamnaya horizon was the material expression of the spread of late Proto-Indo-European across the Pontic–Caspian steppes.
[…] The Yamnaya horizon is the visible archaeological expression of a social adjustment to high mobility – the invention of the political infrastructure to manage larger herds from mobile homes based in the steppes.” ref
“Alternatively, Parpola (2015) relates both the Corded ware culture and the Yamnaya culture to the late Trypillia (Tripolye) culture. He hypothesizes that “the Tripolye culture was taken over by PIE speakers by c. 4000 BCE,” and that in its final phase the Trypillian culture expanded to the steppes, morphing into various regional cultures which fused with the late Sredny Stog (Serednii Stih) pastoralist cultures, which, he suggests, gave rise to the Yamnaya culture. Dmytro Telegin viewed Sredny Stog and Yamna as one cultural continuum and considered Sredny Stog to be the genetic foundation of the Yamna.” ref
“The Yamnaya culture was nomadic or semi-nomadic, with some agriculture practiced near rivers, and a few fortified sites, the largest of which is Mikhaylivka. Characteristic for the culture are the burials in pit graves under kurgans (tumuli), often accompanied by animal offerings. Some graves contain large anthropomorphic stelae, with carved human heads, arms, hands, belts, and weapons. The dead bodies were placed in a supine position with bent knees and covered in ochre. Some kurgans contained “stratified sequences of graves.” ref
“Kurgan burials may have been rare, and were perhaps reserved for special adults, who were predominantly, but not necessarily, male. Status and gender are marked by grave goods and position, and in some areas, elite individuals are buried with complete wooden wagons. Grave goods are more common in eastern Yamnaya burials, which are also characterized by a higher proportion of male burials and more male-centred rituals than western areas.” ref
“The Yamnaya culture had and used two-wheeled carts and four-wheeled wagons, which are thought to have been oxen-drawn at this time, and there is evidence that they rode horses. For instance, several Yamnaya skeletons exhibit specific characteristics in their bone morphology that may have been caused by long-term horseriding. Metallurgists and other craftsmen are given a special status in Yamnaya society, and metal objects are sometimes found in large quantities in elite graves.” ref
“New metalworking technologies and weapon designs are used. Stable isotope ratios of Yamna individuals from the Dnipro Valley suggest the Yamnaya diet was terrestrial protein based with insignificant contribution from freshwater or aquatic resources. Anthony speculates that the Yamnaya ate meat, milk, yogurt, cheese, and soups made from seeds and wild vegetables, and probably consumed mead.” ref
“Mallory and Adams suggest that Yamnaya society may have had a tripartite structure of three differentiated social classes, although the evidence available does not demonstrate the existence of specific classes such as priests, warriors, and farmers.” ref
“According to Jones et al. (2015) and Haak et al. (2015), autosomal tests indicate that the Yamnaya people were the result of a genetic admixture between two different hunter-gatherer populations: distinctive “Eastern Hunter-Gatherers” (EHG), from Eastern Europe, with high affinity to the Mal’ta–Buret’ culture or other, closely related people from Siberia and a population of “Caucasus hunter-gatherers” (CHG) who probably arrived from the Caucasus or Iran. Each of those two populations contributed about half the Yamnaya DNA. This admixture is referred to in archaeogenetics as Western Steppe Herder (WSH) ancestry.” ref
“Admixture between EHGs and CHGs is believed to have occurred on the eastern Pontic-Caspian steppe starting around 5,000 BCE, while admixture with Early European Farmers (EEF) happened in the southern parts of the Pontic-Caspian steppe sometime later. More recent genetic studies have found that the Yamnaya were a mixture of EHGs, CHGs, and to a lesser degree Anatolian farmers and Levantine farmers, but not EEFs from Europe due to lack of WHG DNA in the Yamnaya. This occurred in two distinct admixture events from West Asia into the Pontic-Caspian steppe.” ref
“Haplogroup R1b, specifically the Z2103 subclade of R1b-L23, is the most common Y-DNA haplogroup found among the Yamnaya specimens. This haplogroup is rare in Western Europe and mainly exists in Southeastern Europe today. Additionally, a minority are found to belong to haplogroup I2. They are found to belong to a wider variety of West Eurasian mtDNA haplogroups, including U, T, and haplogroups associated with Caucasus Hunter-Gatherers and Early European Farmers. A small but significant number of Yamnaya kurgan specimens from Northern Ukraine carried the East Asian mtDNA haplogroup C4.” ref
“In 2014, a study discovered a new mtDNA subclade C1f from the remains of 3 people found in north-western Russia and dated to 7,500 years ago. The subclades C1b, C1c, C1d, and C4c are found in the first people of the Americas. C1a is found only in Asia.” ref
“C4 – Upper Palaeolithic (14050 – 13770 years ago) Ust-Kyakhta (Buryatia), Late Neolithic-Bronze Age Irkutsk Oblast, Late Neolithic-Iron Age Yakutia, Tubalar (Ederbes), Todzhin (Toora-Hem, Iiy, Adir-Kezhig), Yukaghir (Andrushkino), Yukaghir/Chuvan (Markovo), Russian, Myanmar
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- C4a’b’c – Irkutsk Oblast (6815 years ago), India (Jenu Kuruba)
- C4a – China (Guangdong, Han from Beijing)
- C4a1 – Mongol from Chifeng and Hulunbuir, Tashkurgan (Kyrgyz, Sarikoli, Wakhi), Czech Republic, Denmark
- C4a1a – Korea, China, Uyghur, Buryat (South Siberia), Denmark, Sweden, France, Scotland, Canada
- C4a1a1
- C-T195C! – Ireland, Scotland, England, USA, Hungary (Szeged region), Poland, Belarus, Russia (Russian, Buryat), Turkey, Pakistan (Hazara), India (Jammu and Kashmir), China (Bargut and Mongol in Inner Mongolia, etc.), Korea
- C4a1a2 – China
- C4a1a2a – China (Han from Ili, Han from Henan, etc.)
- C4a1a2b
- C4a1a2b1 – China
- C4a1a2b2 – Uyghur
- C4a1a3 – Bronze Age Irkutsk Oblast (Ust’-Belaya, Khaptsagai, Silinskij, Chastaja Padi), Russian (Kemerovo Oblast), Koryak, Yukaghir, Yakut, Evenk (Nyukzha, Chumikan, Nelkan/Dzhigda), Even (Sakkyryyr, Sebjan, Tompo, Markovo, Kamchatka), Udinsk Buryat (Kushun), Todzhin (Toora-Hem, Adir-Kezhig), Altai Kizhi, Iran (Qashqai), Sweden
- C4a1a3a – Yakut, Buryat (Buryat Republic, Irkutsk Oblast), Bargut, Nentsi
- C4a1a3a1 – Yakut, Nganasan (Vadei of Taimyr Peninsula)
- C4a1a3a1a – Evenk (Taimyr, Stony Tunguska)
- C4a1a3a1b – Tofalar
- C4a1a3a1 – Yakut, Nganasan (Vadei of Taimyr Peninsula)
- C4a1a3b – Bargut, Uyghur
- C4a1a3b1 – Chelkan, Tubalar
- C4a1a3c – Evenk (Taimyr Peninsula, Stony Tunguska)
- C4a1a3d – Yakut
- C4a1a3a – Yakut, Buryat (Buryat Republic, Irkutsk Oblast), Bargut, Nentsi
- C4a1a4 – Buryat, Kazakhstan
- C4a1a4a – Evenk (Okhotsk region), Shor
- C4a1a2 – China
- C4a1a5 – Teleut, Ladakh
- C4a1a6
- C4a1a6a – Russia (Bashkortostan, Khamnigan), Kyrgyzstan (Kyrgyz), Inner Mongolia (Bargut, Buryat)
- C4a1a6b – Buryat (South Siberia, Inner Mongolia), Uyghur
- C4a1a7 – Denmark
- C4a1b – China, Thailand (Palaung)
- C4a1c – Russia (Bashkortostan, Adygei), Iran (Azerbaijanian), China (Xibo, Mongol from Tianjin)
- C4a1a – Korea, China, Uyghur, Buryat (South Siberia), Denmark, Sweden, France, Scotland, Canada
- C4a2
- C4a2a – Yakut, Evenk (Chumikan)
- C4a2a1 – Bronze Age (2275 – 2040 cal BCE or around 4,275 to 4,040 years ago) Irkutsk Oblast (specimen irk076 from burial 3 at the Shamanka 2 site, South Baikal), Shor, Chelkan, Teleut, Altai Kizhi, Yakut, Kazakh, Ket, Evenk (Stony Tunguska, Taimyr), Buryat (Irkutsk Oblast, Inner Mongolia), China, Korea
- C4a2a1a – Yukaghir, Yakut, Evenk (Nyukzha, Iyengra, Nelkan/Dzhigda), Even (Tompo)
- C4a2a1b – Evenk (Nyukzha), Yakut
- C4a2a1b1 – Evenk (Nyukzha)
- C4a2a1c – China (Zhejiang, Uyghurs), Buryat, Todzhin (Iiy), Karanogay (Dagestan)
- C4a2a1c1 – Tofalar (Alygdzher, Nerkha, V. Gutara), Khamnigan
- C4a2a1c2 – Uyghurs
- C4a2a1d – Uyghurs
- C4a2a1d1 – Udinsk Buryat (Kushun), Tofalar (V. Gutara), Evenk (Central Siberia)
- C4a2a1d2 – Evenk (Nelkan/Dzhigda), Evenk/Nivkh (Val)
- C4a2a1e – Bargut (Inner Mongolia), Buryat (Irkutsk Oblast)
- C4a2a1f – Buryat (South Siberia, Irkutsk Oblast)
- C4a2a1g – Ket
- C4a2a1 – Bronze Age (2275 – 2040 cal BCE or around 4,275 to 4,040 years ago) Irkutsk Oblast (specimen irk076 from burial 3 at the Shamanka 2 site, South Baikal), Shor, Chelkan, Teleut, Altai Kizhi, Yakut, Kazakh, Ket, Evenk (Stony Tunguska, Taimyr), Buryat (Irkutsk Oblast, Inner Mongolia), China, Korea
- C4a2b – Tibet, Korea
- C4a2b1 – Wancho
- C4a2b2 – China (Han from Beijing)
- C4a2b2a – Tibet (Sherpa)
- C4a2c – Bargut (Inner Mongolia)
- C4a2c1 – India (Jenu Kuruba)
- C4a2c2 – Lepcha
- C4a2c2a – Ladakh
- C4a2a – Yakut, Evenk (Chumikan)
- C4a1 – Mongol from Chifeng and Hulunbuir, Tashkurgan (Kyrgyz, Sarikoli, Wakhi), Czech Republic, Denmark
- C4b – Mongol from Jilin and Hulunbuir, Yukaghir, Altai Kizhi, Ukraine, Slovakia
- C4c – Ijka
- C4c1 – Sioux (Carson County of South Dakota), Shuswap, Canada, USA, France, Spain
- C4c1a – Cherokee (Flint District of Oklahoma)
- C4c1b – Chippewa (Trempealeau in Wisconsin), Ottawa or Chippewa (Sault Saint Marie, Chippewa County, Michigan), Canada
- C4c2 – Métis (Red River, Manitoba), USA
- C4c1 – Sioux (Carson County of South Dakota), Shuswap, Canada, USA, France, Spain
- C4a – China (Guangdong, Han from Beijing)
- C4-T152C! – Russia (Bashkortostan), England
- C4-T152C!-A12780G – Uyghur
- C4d – Turkey, Tibet (Chamdo, Nyingchi, Shannan, Lhoba), Thailand (Khon Mueang from Chiang Mai Province), Han from Beijing, Mongol from Tongliao
- C4-T152C!-T4742C – Altai Republic (ancient DNA), Uyghur
- C4-T152C!-T4742C-T16093C – Kyrgyz (Kyrgyzstan), Tibet (Nyingchi)
- C4-T152C!-T4742C-T8602C – Sarikoli (Tashkurgan), Burusho (Pakistan)
- C4-T152C!-T4742C-T8602C-G11176A – Pamiri (Gorno-Badakhshan Autonomous Region of Tajikistan)
- C4e – Teleut, Shor” ref
- C4-T152C!-A12780G – Uyghur
- C4a’b’c – Irkutsk Oblast (6815 years ago), India (Jenu Kuruba)
“People of the Yamnaya culture are believed to have had mostly brown eye colour, light to intermediate skin, and brown hair colour, with some variation.” ref
“Some Yamnaya individuals are believed to have carried a mutation to the KITLG gene associated with blond hair, as several individuals with Steppe ancestry are later found to carry this mutation. The Ancient North Eurasian Afontova Gora group, who contributed significant ancestry to Western Steppe Herders, are believed to be the source of this mutation. A study in 2015 found that Yamnaya had the highest ever calculated genetic selection for height of any of the ancient populations tested. It has been hypothesized that an allele associated with lactase persistence (conferring lactose tolerance into adulthood) was brought to Europe from the steppe by Yamnaya-related migrations.” ref
“A 2022 study by Lazaridis et al. found that the typical phenotype among the Yamnaya population was brown eyes, brown hair, and intermediate skin colour. None of their Yamnaya samples were predicted to have either blue eyes or blond hair, in contrast with later Steppe groups in Russia and Central Asia, as well as the Bell Beaker culture in Europe, who did carry these phenotypes in high proportions.” ref
“The geneticist David Reich has argued that the genetic data supports the likelihood that the people of the Yamnaya culture were a “single, genetically coherent group” who were responsible for spreading many Indo-European languages. Reich’s group recently suggested that the source of Anatolian and Indo-European subfamilies of the Proto-Indo-European (PIE) language may have been in west Asia and the Yamna were responsible for the dissemination of the latter. Reich also argues that the genetic evidence shows that Yamnaya society was an oligarchy dominated by a small number of elite males.” ref
“The genetic evidence for the extent of the role of the Yamnaya culture in the spread of Indo-European languages has been questioned by Russian archaeologist Leo Klejn and Balanovsky et al., who note a lack of male haplogroup continuity between the people of the Yamnaya culture and the contemporary populations of Europe. Klejn has also suggested that the autosomal evidence does not support a Yamnaya migration, arguing that Western Steppe Herder ancestry in both contemporary and Bronze Age samples is lowest around the Danube in Hungary, near the western limits of the Yamnaya culture, and highest in Northern Europe, which Klejn argues is the opposite of what would be expected if the geneticists’ hypothesis is correct.” ref
Yamnaya culture and the Proto-Indo-Europeans (PIE) Language
“Marija Gimbutas identified the Yamnaya culture with the late Proto-Indo-Europeans (PIE) in her Kurgan hypothesis. In the view of David Anthony, the Pontic-Caspian steppe is the strongest candidate for the Urheimat (original homeland) of the Proto-Indo-European language, citing evidence from linguistics and genetics which suggests that the Yamnaya culture may be the homeland of the Indo-European languages, with the possible exception of the Anatolian languages. On the other hand, Colin Renfrew has argued for a Near Eastern origin of the earliest Indo-European speakers.” ref
“According to David W. Anthony, the genetic evidence suggests that the leading clans of the Yamnaya were of EHG (Eastern European hunter-gatherer) and WHG (Western European hunter-gatherer) paternal origin and implies that the Indo-European languages were the result of “a dominant language spoken by EHGs that absorbed Caucasus-like elements in phonology, morphology, and lexicon.” It has also been suggested that the PIE language evolved through trade interactions in the circum-Pontic area in the 4th millennium BCE, mediated by the Yamna predecessors in the North Pontic steppe.” ref
“Guus Kroonen et al. 2022 found that the “basal Indo-European stage”, also known as Indo-Anatolian or Pre-Proto-Indo-European language, largely but not totally, lacked agricultural-related vocabulary, and only the later “core Indo-European languages” saw an increase in agriculture-associated words. According to them, this fits a homeland of early core Indo-European within the westernmost Yamnaya horizon, around and west of the Dnieper, while its basal stage, Indo-Anatolian, may have originated in the Sredny Stog culture, as opposed to the eastern Yamnaya horizon.” ref
“The Corded Ware culture may have acted as major source for the spread of later Indo-European languages, including Indo-Iranian, while Tocharian languages may have been mediated via the Catacomb culture. They also argue that this new data contradicts a possible earlier origin of Pre-Proto-Indo-European among agricultural societies South of the Caucasus, rather “this may support a scenario of linguistic continuity of local non-mobile herders in the Lower Dnieper region and their genetic persistence after their integration into the successive and expansive Yamnaya horizon”. Furthermore the authors mention that this scenario can explain the difference in paternal haplogroup frequency between the Yamnaya and Corded Ware cultures, while both sharing similar autosomal DNA ancestry.” ref
ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref, ref
Early Russian Pottery in Cisbaikal Kitoi culture 7,500 years ago, Samara culture 7,000 years ago, and Yamnaya culture 5,600–4,600 years ago, as well as Proto-Indo-European emergence
“The area east of Lake Baikal in Siberia is one of the few regions in Eurasia where pottery was already used during the Late Pleistocene and Early Holocene. Such early pottery complexes were identified in Ust’-Karenga XII, Studenoye 1, Ust’-Menza 1, and Ust’-Khyakhta 3, dated at about 12-000-11,000 years ago. While around 20,000 years ago East Asian hunter-gatherers were already making ceramic pots. (It seems to Damien) that ceramics spread continually from the earliest centers in China, then Japan, and next the Russian Far East, lastly towards the west, all the way to Europe.” ref
(ANA) Ancient Northeast Asian ancestry today
“Genetically, ANA/Amur ancestry peaks among modern Tungusic, Mongolic, and
“Abstract: Occupied between ~10,300 and 9300 years ago, the Pre-Pottery Neolithic site of Aşıklı Höyük in Central Anatolia went through early phases of sheep domestication. Analysis of 629 mitochondrial genomes from this and numerous sites in Anatolia, southwest Asia, Europe, and Africa produced a phylogenetic tree with excessive coalescences (nodes) around the Neolithic, a potential signature of a domestication bottleneck. This is consistent with archeological evidence of sheep management at Aşıklı Höyük which transitioned from residential stabling to open pasturing over a millennium of site occupation. However, unexpectedly, we detected high genetic diversity throughout Aşıklı Höyük’s occupation rather than a bottleneck. Instead, we detected a tenfold demographic bottleneck later in the Neolithic, which caused the fixation of mitochondrial haplogroup B in southwestern Anatolia. The mitochondrial genetic makeup that emerged was carried from the core region of early Neolithic sheep management into Europe and dominates the matrilineal diversity of both its ancient and the billion-strong modern sheep populations.” ref
“The establishment of Neolithic sedentary societies in southwest Asia was associated with the development of farming practices between 10,000 and 12,000 years ago. Those practices included the cultivation of cereals and legumes and the management of sheep, goats, cattle, and pigs, which ultimately resulted in their domestication. Crop-livestock subsistence strategies started gaining ground around 10,500 years ago in the northern “Fertile Crescent,” and by 9,500 years ago, this mode of subsistence had replaced the foraging lifestyle in parts of southwest Asia and Cyprus. Archaeobotanical and zooarcheological analyses showed that these millennia-long practices of crop cultivation and ungulate management led to phenotypic changes in both plants and animals. To understand these changes, it is often necessary to integrate the evidence of multiple sites and millennia. However, only a few Pre-Pottery Neolithic sites preserved a sufficiently long occupation history and representative faunal assemblages to track morphological, biometric, and demographic changes related to early livestock management at a single location. The list of such exceptional sites in Anatolia includes Çayönü, Cafer Höyük, and Nevalı Çori in Southeastern Anatolia and Aşıklı Höyük in Central Anatolia.” ref
“Aşıklı Höyük is situated on the bank of the Melendiz River. Here, large numbers of caprine bones (i.e., sheep and goats) have been excavated from occupational phases spanning over a thousand years, between ~10,300 and 9,300 years ago. The importance of small livestock management at the site was such that, during the thousand-years occupation, the composition of animal remains identified as sheep and goat increased from ~50% to 87%. Analyses of this extraordinary assemblage provided a unique glimpse into early strategies of sheep management. This includes mortality curves that are indicative of the culling of juvenile males, which in turn reflects exploitation for meat, and spatial patterns of skeletal distribution that imply that slaughtering took place near the living quarters. Further analyses of intra-articular joint pathologies suggested restricted mobility close to the village, including residential stabling, which led to the accumulation of dung and urine salts in the sediments.” ref
“By 9,700 years ago, however, sheep management strategies apparently shifted toward extensive herding. Evidence for this includes a decrease in urine salt and dung concentrations in residential areas, an increase in carcass size (table S2), an improvement in joint health implying greater mobility, and shifts in phytolith and stable isotope profiles that imply more extensive grazing. Together, the evidence obtained at Aşıklı Höyük demonstrates that sheep management in early Neolithic communities was a dynamic process of learning by doing. Although management strategies at Aşıklı Höyük likely affected the phenotype of sheep populations, it is not clear whether they initiated evolutionary changes that ultimately led to the strong genetic differentiation between wild and domestic populations that we observe today.” ref
“A common assumption is that capture and spatial isolation of a subset of a wild population induced a “domestication bottleneck,” provoking the general reduction of genetic diversity evident in modern domestic sheep populations. Here, to address whether the initial management of sheep at Aşıklı Höyük caused shifts in their genetic makeup, we analyzed 629 whole mitogenomes sourced from 15 countries, including 62 from Aşıklı Höyük, spanning a period of over 10,000 years. This allowed us to infer the mitochondrial phylogeography and maternal demographic history of Anatolian and European sheep, and the contribution of the Aşıklı Höyük community to the formation of the Neolithic package dispersing across north and southwestern Anatolia between 10,000 and 8,000 years ago, and subsequently into Europe.” ref
“When we tested the temporal changes in haplogroup frequencies, we found, in general, significant differences between, but not within, groups. The western group included Africa, Europe (modern and ancient), and Neolithic southwestern Anatolia, while the eastern group included Neolithic Aşıklı Höyük, Chalcolithic Güvercinkayası, Caucasus, Levant, and eastern Asia. This pattern was true when the tests were performed with effective population sizes (Ne) between 104 and 106 (we refrained from writing P values here because of the numerous combinations of Ne used in these analyses; see table S6 for the exact values and fig. S2 for a graphical summary of these results). The whole pattern of significance makes sense of the patterns of diversity/neutrality indexes and leads to an important observation that the divergence between eastern and western groups seems to have originated between central and southwestern Anatolia, during the Neolithic. In addition, the temporal tests detected no significant changes in haplogroup frequencies across Aşıklı Höyük’s occupation layers (P = 0.28 to 0.97).” ref
Variable kinship patterns in Neolithic Anatolia revealed by ancient genomes
“Genetic kinship estimated from co-buried individuals’ genomes in Neolithic Anatolia. Close relatives are common among co-burials in Aşıklı and Boncuklu. Many unrelated infants were found buried in the same building in Çatalhöyük and Barcın. Neolithic societies in Southwest Asia may have held diverse concepts of kinship. The social organization of the first fully sedentary societies that emerged during the Neolithic period in Southwest Asia remains enigmatic, mainly because material culture studies provide limited insight into this issue.” ref
“However, because Neolithic Anatolian communities often buried their dead beneath domestic buildings, household composition and social structure can be studied through these human remains. Here, we describe genetic relatedness among co-burials associated with domestic buildings in Neolithic Anatolia using 59 ancient genomes, including 22 new genomes from Aşıklı Höyük and Çatalhöyük. We infer pedigree relationships by simultaneously analyzing multiple types of information, including autosomal and X chromosome kinship coefficients, maternal markers, and radiocarbon dating.” ref
“In two early Neolithic villages dating to the 9th and 8th millennia BCE, Aşıklı Höyük and Boncuklu, we discover that siblings and parent-offspring pairings were frequent within domestic structures, which provides the first direct indication of close genetic relationships among co-burials. In contrast, in the 7th millennium BCE sites of Çatalhöyük and Barcın, where we study subadults interred within and around houses, we find close genetic relatives to be rare. Hence, genetic relatedness may not have played a major role in the choice of burial location at these latter two sites, at least for subadults. This supports the hypothesis that in Çatalhöyük and possibly in some other Neolithic communities, domestic structures may have served as burial location for social units incorporating biologically unrelated individuals. Our results underscore the diversity of kin structures in Neolithic communities during this important phase of sociocultural development.” ref
“This study focuses on social organization across two Neolithic periods. The Aceramic period is represented by Aşıklı Höyük (c. 8,350–7,300 cal BCE) and Boncuklu (c. 8,300–7,600 cal BCE) (Figure 1A), which are among the earliest sedentary communities in Central Anatolia. During the 9th millennium these sites were characterized by small curvilinear buildings, and both maintained mainly forager subsistence practices. The subsequent Ceramic Neolithic period communities were increasingly reliant on food production, and they lived in larger settlements characterized by rectilinear, clustered architecture. In our study, this later period is represented by Çatalhöyük (c. 7,100–5,950 cal BCE), Tepecik-Çiftlik (c. 7,500–5,800 cal BCE), and Barcın Höyük (c. 6,600–6,000 cal BCE). For this study, we screened Neolithic period human remains from Aşıklı Höyük (n = 30) and Çatalhöyük (n = 60) by shotgun DNA sequencing. Owing to adverse conditions and the antiquity of the material, only n = 8 (26%) and n = 14 (23%) skeletons (all petrous bones), respectively, contained ≥0.1% human DNA.” ref
Increased genetic diversity from the Aceramic to the Ceramic period
“We first analyzed genetic relationships at the population level. Principal components analysis (Figure 1B), ADMIXTURE analysis, as well as FST, f3– and D-statistics (Figures S2A) showed that Aşıklı Höyük and Çatalhöyük people belonged to the Central and West Anatolian early Holocene gene pool, represented by Boncuklu Höyük, Tepecik-Çiftlik, and Barcın Höyük individuals, as well as an Epipalaeolithic Central Anatolian individual from Pınarbaşı. Within this regional group, we discern genetically distinct communities, such that individuals from these sites (except for Tepecik-Çiftlik) tended to share more recent common ancestry with individuals from the same settlement compared with those of other settlements (among 576–11,780 D-tests per site, 84%–93% were nominally significant in this direction; Figures S3D). FST, f3– and D-statistics also showed that residents of the two Aceramic Neolithic settlements, Aşıklı Höyük and Boncuklu Höyük, were genetically highly similar to each other (Figures 1C, S2A, S2B, and S2D–S2F) relative to Ceramic Neolithic-period populations.” ref
“Aceramic Neolithic-period populations had lower within-group genetic diversity (measured using the f3-statistic) than did Ceramic Neolithic groups (Figures 1D and S2C, and Tables Z8 and Z9) and carried a higher fraction of short runs of homozygosity (ROH) than most Ceramic Neolithic genomes (Figure S3G). This temporal increase in diversity, also noted in earlier studies, could be explained by two non-exclusive scenarios, namely population growth and genetic admixture. By testing D(Outgroup, X; Aceramic Anatolian, Ceramic Anatolian), where X represents an early Holocene Zagros or Levantine population, we found results compatible with southern and eastern gene flow into Central and West Anatolia between roughly 7,500 and 6,500 cal BCE (Figure 1E and Table Z4) as previously suggested. Using qpAdm, we could also model Ceramic Neolithic Anatolian populations as mixtures of c.90% Aceramic Neolithic Anatolian ancestry (estimate ± 1 standard error: 89%–92% ± 2%–4%) and c.10% Levantine ancestry (8%–11% ± 2%–4%) (models that included Zagros or Caucasus populations were not supported) (Table Z10). Notably, the timing of increased population mobility is contemporaneous with a stronger reliance on agriculture and animal husbandry as food sources, a shift to larger buildings, likely population growth, and possible shifts in patterns of social organization, as we describe below.” ref
Estimating pedigree relationships among Neolithic co-burials
“Neolithic Southwest Asian settlements contain structures that are usually interpreted as domestic dwellings that served as focal points for the socialization of household members. These societies frequently interred their dead, including subadults and adults of both sexes, beneath the floors of these buildings while they were inhabited by the living. A common assumption has been that these burials were of household members who were related in some way, possibly genetically or through social kinship.However, it is not yet clear if individuals buried under house floors necessarily lived in those structures as part of a single co-resident group. The extent of dietary similarity among individuals interred within the same building, for instance, is currently ambiguous. Nevertheless, the assemblage of burials within or around domestic structures is expected to carry information about household composition and/or burial practices, and it may shed light on the relative importance of genetic relatedness as an organizing principle within these early Neolithic communities. In previous studies at Çatalhöyük, analyses of dental morphometrics and of mitochondrial DNA have suggested that individuals interred within the same building are often not genetically closely related. The question has remained unresolved, however, due to the inability of either data type to sufficiently identify exact pedigree relationships on any one site.” ref
“Here, we re-address the question of co-burial relationships using genome data from Neolithic Anatolian communities. In order to infer reliable pedigree relationships, we used different sources of information simultaneously. First, we employed three allele frequency-based methods to infer genetic kinship coefficients: NgsRelate, lcMLkin, and READ (Figures 2, S4A, and S4B). Second, to distinguish different pedigree relationships among putative first-degree pairs (e.g., siblings, mother-son, father-daughter), we used the probabilities of sharing 0, 1, or 2 alleles identical-by-descent (Cotterman coefficients; k0, k1, k2), although the low coverage of our genome data constrained the utility of this latter approach. Therefore, for inferring pedigree relationships we combined (a) kinship coefficients (θ) estimated from autosomal and from X chromosomal loci, (b) mitochondrial haplotype sharing, (c) osteological age-at-death estimates, and (d) radiocarbon dates. Finally, we performed pedigree simulations to determine the power of kinship coefficient estimation using low coverage data (Figure S4C). In addition, we studied the performance of the kinship estimation algorithms on negative controls, that is, real data from pairs of individuals who could historically not be close relatives. We hence limited the kinship tests to pairs of individuals sharing a minimum of 5,000 single nucleotide polymorphisms (SNPs) (Figure S1B). This permits reliable estimations of genetic relatedness up to the 3rd degree (e.g., cousins). Pairs related beyond the 3rd degree are here referred to as “unrelated.” ref
“The final dataset included a total of 223 pairs of individuals buried within the same sites, who were broadly contemporaneous, and who had sufficient genomic data for reliably inferring genetic relatedness (Tables S1, S3, and Z11). Of these, co-burials comprised 32 individuals and 50 pairs, including 2–6 burials associated with the same building or building clusters (i.e., co-burials). In Çatalhöyük and Barcın, co-buried individuals who could be genetically sampled only included subadults. Importantly, all these buildings either had evidence of domestic use (e.g., hearths) or lacked evidence of systematic non-domestic use (e.g., use as animal penning), and did not deviate from others of the same layer in terms of structure or elaboration.” ref
Co-buried pairs in Aceramic period sites frequently include relatives
“The data from Aşıklı Höyük included genomes of five individuals from the same stratigraphic layer who produced statistically consistent radiocarbon ages (χ2 = 7.6, χ 2(5%) = 9.5, n = 4; Table Z2) and could have lived at the same time. These individuals, all females, were interred in two buildings in close proximity and that shared a workspace, likely used by a single household (Figure 3A). All three methods identified two pairs of first-degree relatives (Figure 2A, and Tables S3 and Z11). One pair buried in the same building included an adult and child (individuals 136 and 131). The other pair, buried in separate but proximate buildings, included an old adult and child (individual 133 and 128). The genetic and skeletal evidence indicated both pairs to be sisters (Figures 2A–2C and Tables S3 and Z11). However, we cannot exclude parent-offspring relationships. An adult female (individual 129), buried in the same building as individual 128, had no genetically close relatives among the other four individuals. Thus, although only a minority of studied individual pairs (2 of 10 pairs) were closely related, the majority of individuals studied (4 out of 5) had one close relative identified in the same or adjacent building (Figure 2D and Table S1).” ref
“The Boncuklu Höyük data comprised nine genomes of individuals who were buried in three buildings or in external spaces. Five individuals formed a co-burial cluster in two adjacent consecutive buildings (Figure 3B). Among these, two pairs of first-degree relatives were identified (Figures 2A–2C; Tables S3 and Z11) (also reported earlier). The first was a possible mother and her adult son (individuals ZHF and ZHJ), who were buried in the same building (B14). Their radiocarbon results were different at the 1% significance level (χ2 = 8.8, χ2 (5%) = 3.8, n = 1; Table Z2), and suggested that the woman (ZHJ) died first with 90% probability. The second included a possible pair of adult male and female siblings (individuals ZHBJ and ZHAF). These individuals were buried in the proximate consecutive buildings (B12 and B14). Thus, as at Aşıklı Höyük, we could identify close relatives across the majority of individuals (4 out of 5) associated with neighboring building pairs (Figure 2D and Table S1). The only exception was a perinatal infant (individual ZHAG). Intriguingly, this infant was buried in the same grave with an adult female (individual ZHAF). The infant also shared the woman’s mitochondrial haplotype but was closely related to neither the woman nor any other individual studied. Other individuals also lacked close relatives in this dataset.” ref
Relatives are rare among Çatalhöyük and Barcın intramural burials
“The Çatalhöyük data contained genomes of 14 individuals from multiple stratigraphic levels. All except one individual were subadults; 10 and 4 were genetically determined to have been females and males, respectively. Ten subadults, buried in three buildings dating to the mid-7th millennium BCE, constituted three co-burial clusters (Figure 3C). We identified a single pair of female siblings (individuals 2728 and 2842), an infant and a child, buried within the same building (Building 50) (Figures 2A–2C, and Tables S3, Z2, and Z11). The pair produced statistically consistent radiocarbon measurements (χ2 = 0.0, χ2 (5%) = 3.8, n = 1). None of the other pairs of individuals tested were closely related. Hence, among Çatalhöyük individuals co-buried in these three buildings and tested genetically, only 2 out of 10 had genetic kin identified (Figure 2D and Table S1).” ref
“The Barcın Höyük data included genomes of 23 individuals from multiple phases (VIa, VIb and VIc or VId2/3) (Figure 3D). Ten of these individuals were inserted into three or possibly four buildings (Table Z2). We determined two pairs of relatives, including a pair of subadult sisters (associated with Building 5) and a pair of subadult males who were second- or third-degree relatives (associated with Buildings 14/15) (Figures 2A–2C and Tables S3 and Z11). Both pairs were buried in close proximity to each other and produced statistically consistent radiocarbon measurements (L11 213 & 215, χ2 = 0.7; M10 271 & 275, χ2 = 0.2, χ2 (5%) = 3.8, n = 1 for both; Table Z2). None of the other individuals had close relatives identified, including four infants buried in Building 4. Hence, among co-buried individuals we could identify relatives for only 4 out of 10 (Figure 2D and Table S1).” ref
“The Tepecik-Çiftlik data included genomes of a total of five individuals from two strata. We identified a probable pair of a mother and her adult son (individuals 37 and 21) buried in different parts of the same building (Building AY/AK) (Tables S3 and Z11). These individuals produced radiocarbon results that are different at the 1% significance level (χ2 = 8.0, χ2 (5%) = 3.8, n = 1), which suggests that it is 96% probable that the woman (individual 37) died first (Table Z2).” ref
Temporal or age-dependent variability in co-burial kinship patterns
“The identification of multiple instances of close genetic relatedness among co-burials across all Neolithic Anatolian settlements studied suggests that early Neolithic social arrangements and possibly household composition were to some extent linked to genetic ties. Although long assumed, genetic relatedness within Neolithic house-related social groups is documented here directly for the first time. This is particularly salient in the evidence from 9th and early 8th-millennium BCE Aşıklı Höyük and Boncuklu Höyük and could be considered suggestive of elements of close genetic kin relationships among groups buried together within Aceramic Neolithic houses.” ref
“Nevertheless, a notable fraction of our sample also contained individuals (nearly all subadults) buried in buildings together with genetically unrelated individuals (50% of 32 individuals; Figure 3). Genetic relatedness among co-burials was especially low in the 7th-millennium BCE Çatalhöyük and Barcın Höyük, with the majority of co-burials lacking identifiable genetically related kin (the sample size from Tepecik-Çiftlik is too small to reach a general conclusion). Indeed, the combined frequencies of individuals among co-burials with and without identified relatives appeared different between Aşıklı and Boncuklu versus Çatalhöyük and Barcın Höyük (odds ratio = 8.6, Fisher’s exact test p = 0.019; Figure 2D and Table S1). However, the difference becomes non-significant when including the co-buried adult pair from Tepecik-Çiftlik in the temporal comparison between Aceramic and Ceramic period sites (odds ratio = 6.6 and p = 0.054).” ref
“Two points need further mention. First, although all age groups are represented archaeologically among Çatalhöyük and Barcın Höyük burials, among samples with sufficient DNA data we had high proportions of subadults (13/14 and 16/23, respectively). This effect appears to be caused by better DNA preservation in subadult bones, at least at Çatalhöyük (Figure S1C; STAR Methods), possibly as a result of age-based differences in burial treatment. As a consequence, in our study, no adult co-burials could be genetically examined from these two sites. Second, Çatalhöyük and Barcın Höyük buildings were significantly larger and contained more burials than those of the Aceramic Neolithic sites (Figure 3).” ref
“The infrequency of close relatives among subadults buried together in relatively large structures at Çatalhöyük and Barcın Höyük is intriguing. It raises the question of whether these buildings may have been used by extended families, such that the co-buried subadults could be distant cousins who were not identified by the methods employed. We thus tested whether individuals buried in closer proximity shared greater genetic similarity, using genetic distances based on the f3-statistic (STAR Methods). After excluding close relatives, we found no correlation between genetic distance and spatial distance across burial pairs in either Çatalhöyük or Barcın Höyük (Pearson r < 0.02, Mantel test p > 0.3; Table S4).” ref
“Researchers also tested the hypothesis that overall genetic similarity among co-burials might be higher within buildings than between buildings. Again, we found no evidence for this (one-sided permutation test p > 0.8; Table S4). These results corroborate previous analyses that found no significant correlation between burial location and dental similarities in Neolithic Çatalhöyük adults and also a lack of mitochondrial DNA shared among co-burials. We note that we do not expect all individuals associated with these buildings to have been buried within those structures. Also, not all individuals interred in these buildings could be sampled in this study. Still, the presence of individuals without identified relatives implies that the choice of the same structure for the burial of community members may be motivated, among other factors, by additional forms of social connectedness.” ref
“For instance, co-burials, including juveniles, may have included “adoptive, foster or fictive kin held together by memory and history-making.” Accordingly, co-burial and perhaps household composition in these later Neolithic settlements may have included—but also extended beyond—close genetic kin. It is also possible that the practice of co-burying subadults with genetically unrelated individuals was already present in the Aceramic period in Anatolia, but we did not sample these sufficiently in Aşıklı and Boncuklu. Indeed, the Boncuklu adult female-infant pair sharing a grave, found to be unrelated, may reflect such a tradition. It, therefore, remains unclear, yet, whether the difference among sites in co-burial patterns reflects a temporal shift or differential treatment of adults versus subadults in Neolithic Anatolia.” ref
Varying traditions linking sex and space
“Another set of observations involves burial patterns with respect to sex. First, we find co-burial of closely related adults of both sexes at Boncuklu Höyük and possible adult-child sister pairs at Aşıklı Höyük. Although our sample size is limited to reach a definitive conclusion, it is worth noting that the pattern is consistent with adult females retaining close ties to their natal households, symbolically or residentially, over significant periods of their lives. This scenario, at least at Boncuklu Höyük, could equally have applied to the males. Second, the sex patterning observed in Anatolian Neolithic burials appear distinct from those described for Neolithic and Bronze Age cemeteries in Europe, where male burials predominate, and patrilocality is evident.” ref
“For instance, in a study of multiple cemeteries, Mittnik and colleagues identified only 2 first-degree related female pairs out of 21 first-degree relationships. This proportion is different in our data, which reveals 4 first-degree related female pairs out of 7 first-degree relationships (odds ratio = 11.1, Fisher’s exact test p = 0.02). This result, as well as the contrast between co-burial of related adult females in the Aceramic period buildings and the stark patrilocal patterns in 6th-3rd millennium European cemeteries, are consistent with the notion that sex role differences intensified following the initial adoption of agriculture. Meanwhile, both sister pairs we identified at Barcın and Çatalhöyük were subadults. In this regard, patrilocal traditions in Ceramic period Anatolian sites remain a possibility (as suggested earlier based on dental and mtDNA data).” ref
“In summary, in addition to evidence for the existence of close genetic ties among putative households in the Aceramic period, we find that genetic relatedness among subadult co-burials was infrequent at Ceramic period Çatalhöyük and Barcın. Although we cannot yet pinpoint when and where this latter practice emerged, it appears plausible that during the transition from the Aceramic to the Ceramic Neolithic period in Anatolia, in parallel with changes in subsistence and population mobility, genetic relatedness may have become less important in the structuring of intramural burial traditions.” ref
Neolithic buildings and households
“The concept of “house” refers to a social institution through which societies define a particular type of membership group, i.e., the “household.” What defines a household is based upon the cooperating individuals’ criteria for relatedness, task-orientation, and co-residence. These criteria are socio-culturally constructed and, therefore, highly variable across societies. For example, household members can be genetically related, as in genetic kin-based family organizations, but a household can also be composed of individuals who co-reside and share tasks with reference to relatedness criteria other than genetic ties. Nevertheless, these criteria of relatedness, genetic or otherwise, are considered legitimate only if they express continuity through successful invention and manipulation of concepts such as descent, belonging and other social differences based on age, sex and skill, all of which are also actively employed in terminologies of kinship or affinity. Within this context, long-lasting architecture has been the most potent embodiment of inclusion and relatedness, through which a household membership and its history can be represented via a variety of symbolic activities.” ref
“Some of the earliest long-lasting residential architecture, considered to be the primary context for the socialization of household members, is found in early Neolithic SW Asia, c. 10th-7th millennia cal BCE. The criteria that define relatedness among the household members of these societies, however, have long been debated: were the co-residents genetic kin, or did other factors determine household membership? Based on the size and form of the buildings, it has been suggested that the earlier curvilinear structures of the c.10th-9th millennium cal BCE were used by extended families, perhaps related to polygamous social structures, whereas the adoption of larger rectilinear and compartmentalized buildings of 8th-7th millennium cal BCE reflects a shift to close genetic kin-based organization.” ref
“Alternatively, given the relatively small size of most Neolithic residential structures, regardless of shape, it has been postulated that these buildings were mostly used by nuclear families. Other researchers hold that the transition from some form of nuclear family household to increasingly autonomous family households occurred during the Pre-Pottery Neolithic B (PPNB). Yet others argue that the increasingly autonomous households only occurred in the Late Neolithic as an element of multiscalar transformations of Neolithic communities in this period. Research on mortuary practices also underlines the broad regional changes through time, including suggested shifts from community membership to increasingly separate and autonomous household organizations in the PPNB. Meanwhile, the repeated construction of mudbrick buildings at the same location over multiple generations, sometimes even maintaining the position of internal structures such as hearths, implies the presence of distinct household identities in these societies.” ref
Do co-burials represent households?
“One potential source of information that could help resolve the nature of Neolithic household composition and social organization comes from burials within buildings during their occupancy. Neolithic SW Asian societies frequently practiced the burial of individuals beneath the floors of domestic buildings, usually during the time these structures were inhabited. A common assumption has been that these burials were of household members and were related in some way, possibly genetically or through kinship based on other factors. This could include households composed of families of closely genetically related individuals, extended families, multi-family households, or social units where genetic relatedness had little role. In reality, however, it remains unclear whether individuals buried under house floors lived in the same building as part of a co-resident group, i.e., whether they represented households.” ref
“If co-burials were indeed household members, we may expect them to share specific attributes more with each other than with other co-burial groups; most notably, elements of their diet. Evidence on dietary similarity among Neolithic Anatolian households is currently equivocal. A 2015 study reported no dietary differentiation among Çatalhöyük co-burials in different buildings. A 2020 study using a wider dataset again from Çatalhöyük reported statistically significant differentiation in carbon and nitrogen isotope values among buildings. This same work further reported significant dietary differences among neonates buried in different buildings. Still, possible confounding factors that could influence stable isotope profiles (age and sex for adults, pathological conditions for neonates) were not explicitly controlled for in these analyses, and we therefore consider these results as preliminary.” ref
“There exist additional arguments against the hypothesis that co-burials represent households. It appears that the average number of burials per residential structure is generally too small to represent full households. For instance, in Aşıklı Höyük, only 90 burials have been discovered from more than 400 rooms excavated. This suggests additional factors influenced the choice of burial locations and type of funerary treatments of individuals. Furthermore, an apparent excess of burials in some residential buildings, in sites such as Çatalhöyük, and occasionally at other sites such as Abu Hureyra and Bestansur (although the relevant buildings here may not be ordinary residential structures), implies a special role of some residential buildings for burial of individuals who probably had originally lived in other residences. Düring and Marciniak’s (2005) analysis of Çatalhöyük houses also indicates that human burials in buildings may have served to advertise the temporal continuity (history) of the buildings, which thus ensured the continuity and success of the household, regardless of their genetic ties.” ref
“If co-burials do not represent household members, their interment in the same buildings could be driven by at least two distinct traditions. First, individuals may be buried together because they died at the same time. This could also include mass burials following disease outbreaks. However, in the case of co-burials in Neolithic Anatolian settlements, the mortuary context and mortality profiles do not indicate mass burials. The evidence overall suggests these were collective burials, where individuals were buried sequentially, as is prevalent at Neolithic Çatalhöyük as well as other sites.” ref
“Second, co-burial patterns may reflect local traditions stipulating specific burial arrangements of individuals who do not belong to the same household. Such traditions could involve burying individuals of specific status or social backgrounds together. The motivation behind these traditions may be to maintain social and economic ties among groups and to “consolidate community membership”. For instance, it has been suggested that the emergence of cemeteries during the Natufian period could have represented “the establishment or strengthening of special interest groups, inheritance of corporate property, and territorial ownerships”. Another example could be traditions such as described for Aboriginal Australian groups where the corpses of deceased young children were retained by the mothers to be interred with an adult male who dies next (Musgrave 1930, cited in). If such arrangements were in place also in Neolithic Anatolian settlements, we might expect no direct social or genetic connection among co-burials.” ref
Relatedness among co-burials
“Studies on genetic relatedness among co-burials in Neolithic SW Asia have yet been limited. Most work to date relies on dental metric and non-metric traits as proxies for genetics, and one recent study used mitochondrial DNA. These studies have reported patterns consistent with endogamy or with matrilocality in Neolithic Levantine sites, and with patrilocality at Çatalhöyük. Meanwhile, the Çatalhöyük studies, based either on dental analysis or mitochondrial DNA, found no evidence for individuals buried in the same building being more closely related to each other than to individuals buried in other buildings.” ref
“Still, owing to the inability to estimate the degree of kinship using dental traits and mitochondrial data, the question of kinship among co-burials in Neolithic SW Asia has remained largely unresolved. Ancient genomics, in turn, can be used as a powerful tool to determine genetic relatedness and kinship among households of the dead, allowing further consideration of how burial locations might have structured relationships between households of the living and the construction of kinship, as well as social memory and social traditions in general. With some temporal depth to our study we are also able to consider if there might be temporal trends in these social practices over the long term.” ref
Description of archaeological sites
Description of Aşıklı Höyük
“Aşıklı Höyük, located in the Volcanic Cappadocia Region in eastern Central Anatolia is one of the earliest sedentary communities of the region, radiocarbon dated to the mid-9th and 8th millennium BCE (8350-7300 cal BCE). Excavations at the site started in 1989 as salvage excavations under the direction of Prof. Ufuk Esin (İstanbul University). Since 2010, the research and excavation project has been led by Prof. Mihriban Özbaşaran (İstanbul University) and Güneş Duru (Mimar Sinan Fine Arts University) in collaboration with an international team from various universities and institutions.” ref
“The first inhabitants of Aşıklı settled near the western bank of the Melendiz River. The river, flowing from the Ihlara Valley, and the volcanic landscape provided a rich habitat for various animal and plant species. A warm climate and park-woodland vegetation was dominant in the region during the beginning of the Holocene. The mid-9th millennium BCE inhabitants of the site lived in semi-subterranean, oval mudbrick buildings that were reconstructed and renewed periodically at the same location. Characteristics of these buildings include hearths, a small platform, grinding stones and burial pits. Daily life was organized outside the buildings, in open activity areas, where many of the daily tasks were conducted.” ref
“Archaeozoological data attest to broad spectrum hunting during the 9th millennium BCE, including a variety of small prey animals, birds and fish, although the main focus was always on sheep/goat. Analyses of micromorphology and soil chemistry, and the presence of primary dung layers attest to the fact that animals were kept on-site, inside wattle and daub enclosures. Archaeozoological data, as well as isotope analysis show that caprines, specifically sheep, were kept in the settlement from the earliest levels; management and the domestication process continued all through the sequence. The community had the knowledge and the experience of growing plants and cultivating wild and domestic cereals. Wild plants, legumes and fruits were among the gathered plants.” ref
“With the start of the 8th millennium BCE, changes took place in architecture and settlement patterns. Rectangular structures replaced the oval and semi-subterranean buildings. These rectangular buildings were mostly single-roomed. Although few in number, buildings with two or three-room buildings are also present. Toward the end of the settlement occupation, buildings started to cluster. Building clusters, generating neighborhoods, were separated by narrow spaces or passages with access to communal middens. Separated by a “gravel street” from the residential area, to the southwest of the present mound, lies a building complex distinguished from domestic buildings in terms of its plan, size, construction material, internal architectural features and floor and wall treatment. The architectural features and the characteristics of the archaeological material (i.e., the dominance of wild cattle) permit interpretation of this area as a “public area” where communal consumption and certain ceremonies took place. Evidence of communal activities in this area indicates the continuity of the collective way of living, while the daily activities in the residential area most probably took place on the flat roofs and inside the dwellings. During this period, hunting and gathering continued, though with less importance, and subsistence was based mainly on sheep and goat, but these animals were no longer kept within the settlement.” ref
“Two concepts, a communal way of life and continuity, characterized the social organization of the Aşıklı community. Interaction with other regions and communities had a certain tempo during the mid-9th millennium BCE, as evidenced by the material culture. However, simultaneous with the increasing focus on the full establishment of sedentism and caprine management, the pace of interaction decreased, only to increase again during the last 200-300 years of site occupation, corresponding roughly to the second half of the 8th millennium BCE. This is well illustrated by the sudden appearance of non-local materials and technologies during this period. In contrast with this pattern of temporal change, continuity of certain elements, such as the location of the buildings and interior architectural features, constant renewal and maintenance of the floors and walls of buildings, and the transferring of objects and know-how was another factor that characterized the social fabric of the community. The inhabitants managed to live in cohesion throughout the occupation sequence and the communal way of life was maintained with new solutions, but also continuity through temporal changes and transformations was the main characteristic of the newly established Neolithic way of life at Aşıklı Höyük.” ref
“The burial customs consist of intramural, single sub-floor inhumations. The deceased were buried in pits under the floors of the buildings in a flexed position. To date, 90 burials have been found in 400 rooms. Although this tradition was not subject to change for hundreds of years, new practices arose during the latest levels of the occupation at the site. The dead were not buried with any items of personal adornment during the mid-9th millennium BCE. However, changes can be observed toward the mid-8th millennium when some individuals are found buried with ornaments. Of the 82 individuals subjected to bioarchaeological analysis, adults constitute 60% while children make up 40%. Of the 46 adults for whom sex can be determined, females constitute 65% while males constitute 35%, a marginally significant difference (binomial test p = 0.054). In terms of the daily activities conducted by the Aşıklı Höyük individuals, task-related pathologies of adults show that the shoulders, hips, ankles, elbows and knees were affected by osteoarthritis, possibly stemming from habitual stress. Males exhibit significant degrees of shoulder osteoarthritis, followed by their elbows and hips; for females the ankles were most affected by this disease, followed by the shoulders and hips. This may suggest that males were routinely engaged in activities such as carrying heavy loads, throwing, walking and kneeling, and females were probably engaged in activities that involved walking and squatting.” ref
“Five of these burials genetically studied here were interred in Building 1 and Building 3 of Aşıklı Höyük Layer 4 (Figure 3A). These are buildings in direct proximity with less than 1 m between them, which showed temporal overlap in their periods of use, and which shared a common open workspace between them. We therefore treated the individuals from both buildings as a cluster of co-burials, who might represent members of the same household.” ref
Description of Boncuklu Höyük
“Boncuklu is situated in the middle of the SW Konya basin (37°45’N 32°52’E) and lies 33.4 km northwest of the site of Pınarbaşı and 9.5 km northeast from Çatalhöyük. The site was discovered during the archaeological survey under the direction of Prof. Douglas Baird from the University of Liverpool, UK. Excavations directed by Baird began in 2006 and continue at the present time. Baird was joined by co-directors, Prof. Andrew Fairbairn of University of Queensland, Australia and Dr. Gökhan Mustafaoğlu of Ankara Haci Bayram Veli University, Turkey, since 2011. Occupation of the site is documented from 8300-7600 cal BCE directly through radiocarbon dating. However, stratigraphic and material evidence suggest a slightly longer span of occupation.” ref
“The exploitation of wild resources seems to have predominated, especially wild cattle and boar, fish and wetland birds, along with nuts and fruits from surrounding hill areas. Small-scale cultivation of wheat, lentils and peas was an additional modest component of subsistence activities. The chipped stone industry was microlithic, in significant contrast to broadly contemporary Levantine PPNB and northern Fertile Crescent assemblages and thus shows significant continuities with the earlier, local Epipalaeolithic and the earlier 10th/early 9th millennium BC community at Pınarbaşı in technology and raw material. Continuities between Epipalaeolithic and early Holocene forager communities and the community at Boncuklu are clear. This evidence is supported by significant genetic continuity. By 8300 cal BCE it appears local foragers adopted domestic plants from areas to the south and east, incorporating them into their traditional wetland exploitation practices. They were probably introduced to the region as a consequence of the far-reaching and continuous interactions with neighboring regions from the Epipalaeolithic through the 10th-early 9th millennia cal BCE, as also documented at earlier and contemporary Pınarbaşı.” ref
“The site possessed a number of sunken-floored sub-oval domestic buildings with mudbrick walls. The households display highly structured use of internal house space, divided into a ‘clean’ area presumably for sleeping, socializing and food consumption and a ‘dirty’ kitchen area. The houses were very regularly refurbished, plastered and modified, especially the hearth areas, showing the intensity of domestic use. The floor area of these houses is small and modeling shows small intimate household units with intensive and repetitive domestic practices. Evidence of ritual and symbolism in the ‘clean’ areas, including burials, is regular and differentiated from house to house suggesting creation and maintenance of distinctive household identities. The Boncuklu houses were also repeatedly and continuously reconstructed over multiple generations in the same location, a practice at some other 10th-7th millennia cal BCE sites in the surrounding regions, for example to the northeast at Aşıklı from 8300 cal BCE, just to the south at Çatalhöyük from 7100 cal BCE, in the Levant at PPNA Jericho and in PPNB Tell Halula. This seems to be a symbolic statement of household continuity. This expression of continuity and identity suggests small tight-knit households in continuous occupation of these domestic structures, whatever the nature and dynamism of their composition, which we can start to grasp through aDNA evidence. Nevertheless, there seems evidence that some broader corporate social practices cross-cut households, including some practices involved in food and resource exploitation in the wider landscape.” ref
“Primary inhumations were placed under the ‘clean’ area of the houses during their occupation. It seems the dead ‘ancestors’, whether biologically related or not, were kept close to the living. In the case of Boncuklu the modest numbers of burials under house floors, maximum 5 and more usually 1-3, per house, suggest many of these could easily be members of the household that lived in these buildings, although we certainly cannot assume that to be so. Nevertheless, reflecting the fact they occurred within the houses while still in use and that these were small-sized buildings with very intimate spaces, presumably means the co-burial of the dead expressed some type of relationship to the households of the living, and thus represented a symbolic statement of connection between the dead and the living. Indeed, evidence attests to ongoing attention to burials and knowledge of their location.” ref
“There were also primary burials and burials of deliberately disarticulated human remains, including human crania, in open areas between buildings in areas of midden accumulation. More than 37 Neolithic burials, plus a minimum of 274 individual bones and 129 isolated finds of human remains have been studied, although more have been excavated. Nine skeletal samples from securely stratified 9th-8th millennia cal BCE burials in Areas H, K, and M provided sufficient aDNA preservation for genetic analysis (Table Z2), and thus genomic data. Boncuklu human remains do not reflect significant disproportionate representation of males or females and there is an even spread across most age categories, including children and young, middle and old adults, with a slight, but not unusual, lesser presence of older children/adolescents.” ref
“Five of these burials (ZHF – Grave 14, ZHJ – Grave 15, ZHAF – Grave 18, ZHAG – Grave 18 and ZHBJ – Grave 30), including 2 pairs of individuals with first-degree genetic relationships were all articulated primary inhumations stratified within a sequence of 2 buildings in Area H, Building 12 and Building 14. Building 12 predates Building 14 and, indeed, the foundation cut for Building 14 removed the northern edge of Building 12 (Figure 3B). Building 14 seemed a direct replacement for Building 12, an example of the continuous reconstruction of the buildings in the same locations, although in this case with some shift of the house to the north. ZHBJ, the likely brother of ZHAF, was buried in the northern part of Building 12 (Figure 3B). ZHAF, his likely sister, was buried in the southern part of Building 14 (Figure 3B). This seems a deliberate attempt to keep these individuals close at death and points to the close connections between the living and dead in these households. Both these burials had similar orientations, approximately west-east/northwest-southeast with heads at the West. It is thus tempting to think this might also reflect their close relationship.” ref
“It may well have done, but these are the most common burial orientations at Boncuklu, among c. 70% of the analyzed burials and so might simply reflect these broader patterns. ZHAG, a female perinatal child that likely died at birth, was placed directly against the pelvis of ZHAF, but was genetically unrelated to that adult female ZHAF and also unrelated to ZHF and ZHJ in the same building. It is, of course, possible her mother lived in Building 14 but was genetically unrelated to the other adults buried there, or that as a result of some form of connection to the child and/or her mother she was buried with ZHAF, albeit from a household who lived elsewhere. ZHF and ZHJ, most likely adult son and mother respectively, were located in the more eastern parts of Building 14 (Figure 3B). The orientation of their bodies was not dissimilar, ZHF had the common northwest-southeast and ZHJ a north-south orientation. However, their heads were at opposite ends of the grave-cut, ZHF to the northwest and ZHJ to the south. It is, therefore, difficult to suggest that orientation at Boncuklu was a direct expression of close family relationships.” ref
“ZHAJ was a primary inhumation burial of an adult male that predated Building 14 and seems to have been located in an open area. ZHB was the burial of an adult female burial post-dating Building 14. Overlying stratigraphy was eroded so it was unclear whether the grave for ZHB was cut through the floor of a building or was placed in an external area. These two burials do not show any close genetic relationships to the other sampled individuals. The other burials analyzed, genetically unrelated to any of these burials in Area H, was one adult male primary inhumation, ZKO, buried in Building 9 in Area K, broadly contemporary but c. 15 m from the cluster in Area H. ZMOJ was a primary inhumation in an external area in the middens of Area M, located c. 25 m from the cluster in Area H. Although well stratified in Neolithic deposits the chronological relationship with the Area H cluster and ZMOJ is not clear.” ref
Description of Çatalhöyük
“Located 9 km to the south of Boncuklu Höyük on the Konya Plain in Central Anatolia, the site of Çatalhöyük was discovered and first excavated between 1961-1965 by James Mellaart (British Institute of Archaeology at Ankara), and later between 1995-2015 by Ian Hodder (Stanford University). Çatalhöyük was designated a UNESCO World Heritage Site in 2012. The site consists of two separate mounds or “tells.” The larger East Mound covers an area of 13 ha and has been dated to c. 7100-5950 cal BCE, corresponding roughly to the Ceramic Neolithic period. The smaller West Mound dates to the Early Chalcolithic and was occupied until the middle of the 6th millennium BCE. The Neolithic East Mound, until c. 6300 cal BCE, is characterized by dense clusters of mudbrick domestic structures interspersed with external spaces used for refuse disposal, animal penning and other daily activities.” ref
“To date, large-scale, clearly identifiable public structures have not been documented at the site. Instead, individual houses at Çatalhöyük appear to have served as the focal point not only for domestic activities such as craft production, food storage and processing, but also ritual behaviors such as burials, wall paintings and other architectural embellishments associated with an elaborate symbolic repertoire and reflecting a complex socio-cultural environment. There is ample evidence for the cultivation of domesticated cereal crops and the keeping of domesticated sheep and goats at the site. Wild animal species, including aurochs, also formed part of the diet, and in the later occupation phases (6500-5950 cal BCE) there is evidence for the herding of domesticated cattle.” ref
“Between 1993 and 2017 the skeletal remains of over 700 individuals had been recovered from stratified Neolithic contexts at Çatalhöyük. Primary inhumations (n = 471 individuals) placed beneath the floors of houses are the dominant burial type at the site. Individuals were typically buried in narrow oval pits under the eastern and northern platforms of the central room, although prenates, neonates and infants were also recovered from within side rooms and near ovens and hearths. Secondary burials of loose or partially articulated skeletal remains, often in association with primary burials, are also observed, although less frequently. Intramural burials became increasingly rare toward the end of the occupation of Çatalhöyük East, while burials are almost completely absent within the settlement on the Chalcolithic West Mound.” ref
“Of the 471 individuals from primary burial contexts, there are 178 adults (20+ years), 29 adolescents (12-20 years), 90 children (3-12 years), 67 infants (2 months-3 years), 85 neonates (0-2 months), and 22 prenates (> 38 weeks in utero). Among the adults and adolescents whose sex could be determined (n = 155), 89 individuals (57%) were assessed as females or possible females, while 66 individuals (43%) were assessed as males or possible males, a marginally significant difference (binomial test p = 0.077).” ref
Description of Barcın Höyük
“Located in the Yenişehir Valley in the province of Bursa in northwestern Turkey, the site of Barcın Höyük yielded an uninterrupted stratigraphic sequence from 6600 to 6000 cal BCE. The settlement was built on a low natural elevation in what would have been a marshland valley. The Neolithic levels at Barcın Höyük, which lie beneath a relatively thin deposit of later levels dating to the Chalcolithic, Bronze Ages and the Byzantine period, are thick and exceed 4.5 m in most places at the site. Called level VI, the Neolithic phase is divided into seven subphases: VIe (earliest level) through VIa. The VIe levels of the site represent the earliest farming community known to date in the Marmara Region. The initial pioneer communities who arrived here around 6600 cal BCE brought with them crops to cultivate and animals to herd. With regards to plants, domesticated varieties of cereals and pulses were plentiful. Sheep and cattle were the preferred herd animals although goats were also present while hunting only contributed a minor part of the diet. Extensive organic residue analyses on pottery demonstrate that the inhabitants of Barcın Höyük relied heavily on dairy products. This observation confirms those made by Evershed and colleagues for later sites in the Marmara Region.” ref
“The initial settlers in the region were accomplished potters even though pottery use was initially limited and indirect methods of heating foods were preferred. Within a century however, thin-walled finely made burnished pots become plentiful. Building on a consistent tradition, recipes of manufacture and temper changed over the ensuing centuries. The residents of Barcın Höyük lived in rectilinear timber frame houses with wood and mud walls. Houses tended to be in rows, surrounded by courtyard areas where a variety of activities were carried out. Burials associated with the settlement were placed within and near structures. Interestingly, many infants were buried within the house proper beneath floors while adults were typically placed in the courtyard areas. Children often tended to be buried outside but closer to the structures, sometimes beneath the floors of the verandas in front of the houses.” ref
“Although intensively analyzed for DNA, the Barcın Höyük skeletons await final anthropological analyses. Based on preliminary data, adults appear to comprise 38% (46 burials) of the 121 burials that come from primary burial contexts. Of the skeletons that can be identified based on sexual characteristics, nearly two thirds of these appear to be females or possibly females. Subadults including adolescents, children, infants and neonates comprise the remaining 62% of the assemblage.” ref
Description of Tepecik-Çiftlik
“Tepecik-Çiftlik is located in the Volcanic Cappadocia region of Central Anatolia in the Melendiz/Çiftlik Plain. The excavators suggest it was occupied from the end of the Aceramic Neolithic Period until the early Chalcolithic Period, between c.7500-5800 cal BCE. The Pottery Neolithic levels show evidence of agriculture and animal breeding, as well as continued hunting and gathering. The site is in close proximity to major obsidian ore beds in the region and is notable for its large amount of obsidian tool remains. Further information about the site may be found at. A 2016 report on Tepecik-Çiftlik indicated that over 170 individuals’ remains dating to the Neolithic levels, buried inside buildings and in open areas had been excavated. A collective burial was also found, and is thought to have been used for successive burials, both primary and secondary. It includes at least 42 individuals of both sexes and various ages.” ref
Description of archaeological material
“This section describes bioarchaeological characteristics of the individuals from Aşıklı Höyük, Çatalhöyük and Boncuklu Höyük. Some of this data are unpublished. Barcın Höyük and Tepecik-Çiftlik individuals included in this study have been described in the supplementary material of Mathieson et al. and Kılınç et al., respectively. Sex was estimated using dimorphic markers, and individual ages-at-death were estimated using standard methods such as human growth and epiphyseal fusion, dental calcification, and bone maturity/size. The sex of subadult individuals listed below have been determined based on genetic data produced in this study.” ref
Description of Aşıklı Höyük individuals
“SK2 (Level 1/2A; Building AB): the burial of a young adult female. Double burial. SK2 was buried in the same burial pit of a male, slightly later. The pit is located in a one-room rectangular building of the mid-8th millennium BCE settlement. Radiocarbon dating places the individual to 7585-7475 cal BCE (Table Z2). SK33 (Level 2C, Building C): the burial of a child, buried under the floor of a rectangular planned kerpiç (mudbrick) building. Radiocarbon dating places the individual to 7945–7890 cal BCE (9%) or 7870–7595 cal BCE (86%). Building C was renewed 10 times at the same location (Figure 3A), where this child’s burial was contemporary with its eighth renewal phase. Excavated in 1991. SK40 (Level 2B, Building BH): the burial of an old adult female. Sub-floor inhumation in a rectangular kerpiç building of the 8th millennium BCE settlement. One of the three individuals buried in the same building: a one-month old infant and a middle adult female. Radiocarbon dating places the individual at 7935–7915 cal BCE (1%) or 7825–7590 cal BCE (94%).” ref
“SK128 (Level 4, Building 3): the burial of a female child. She is one of the two individuals buried in the same building. Radiocarbon dated to 8225–7955 cal BCE (95%). SK129 (Level 4, Building 3): the burial of a young adult female, buried in a semi-subterranean oval building. She is one of the two individuals buried in the same building. Excavated in 2011; primary burial; radiocarbon dated to 8170–8115 cal BCE (6%), 8060–8045 cal BCE (1%), 8010–7985 cal BCE (1%), 7970–7735 cal BCE (86%). SK131 (Level 3E/4, Building 1): the burial of a female child, exposed lying on the pavement of a hearth in a semi-subterranean oval building. This is an exceptional burial, in position and in location. Four more individuals were buried in the same building. The burial was exposed in 2012. She was radiocarbon dated to 8200–8110 cal BCE (16%) or 8095–8035 cal BCE (7%) or 8015–7740 cal BCE (72%).
“SK133 (Level 3E/4, Building 1): the burial of an old adult female, the oldest member of the community thus far excavated. She was one of the five individuals buried in the same oval, semi-subterranean building, B.1. She was a primary burial and was radiocarbon dated to 8170–8115 cal BCE (8%), 8060–8040 cal BCE (1%), 8010–7980 cal BCE (2%), 7975–7735 cal BCE (84%). Excavated in 2012. SK136 (Level 3E/4, Building 1): the burial of a young adult female, one of the five individuals from Building 1. She was a primary burial, and was radiocarbon dated to 8175–8110 cal BCE (7%) or 8090–8075 cal BCE (1%) or 8065–8040 cal BCE (1%) or 8015–7705 cal BCE (84%) or 7695–7655 cal BCE (2%). Excavated in 2015.” ref
Description of Çatalhöyük individuals
“Sk.5357 (burial feature 576, Level South K, Early period, Building 17): primary burial of a male infant. He was 9 months ± 3 months at death based on dental development. It was buried in a flexed position along the east wall of B.17 in association with red pigment and traces of reed basketry. The burial was excavated in 1999. Radiocarbon dating places this individual between 7035–6680 cal BCE (93%) or 6670–6650 cal BCE (2%). Sk.21855 (burial feature 8214, Level South K, Early period, Building 17): the primary burial of a female child. She was 4 years ± 1yr at death based on dental development. It was placed in a flexed position in a burial cut along the west wall of B.17. The burial was excavated in 2016.” ref
“Sk.1885 (burial feature 84, Level South M, Middle period, Building 50): the primary flexed burial of a male child. He was 7 years ± 2yrs at death, excavated in 1995. This individual was interred directly above Sk.2033 (see below) in the southwest corner of B.50. Radiocarbon dating places this individual between 6905–6885 cal BCE (1%) or 6825–6635 cal BCE (92%) or 6625–6600 cal BCE (2%). Sk.2033 (burial feature 84, Level South M, Middle period, Building 50): the primary flexed burial of a male child 3 years ± 1yr at death, excavated in 1995. This individual was interred directly below Sk.1885 (see previous) in the southwest corner of B.50. Radiocarbon dating places this individual between 6690-6590 cal BCE (95%).” ref
“Sk.2017 (burial feature 96, Level South M, Middle period, Building 50): the primary burial of a female neonate (0-2 months at death based on measurements of the basi-occipital bone), excavated in 1997. The burial was located near the oven along the southern wall of B.50. The bones of this individual were scorched as a result of the burial’s proximity to the oven. Radiocarbon dating places this individual between 6815–6790 cal BCE (2%) or 6775–6595 cal BCE (93%) (Table Z2). Sk.2728 (burial feature 258, Level South M, Middle period, Building 50): an undisturbed primary burial of a female infant aged 9 months (±3 months) at death based on dental development. It was excavated in 1997 from Building 50, located in the South Area of the site. The body was placed in a small pit near the eastern wall of the main room. Radiocarbon dating of the petrous bone places this individual between 6695-6505 cal BCE (95%).” ref
“Sk.2779.1 (burial feature 265, Level South M, Middle period, Building 50): the primary burial of a male neonate (0-2 months at death based on measurements of the basi-occipital bone), excavated in 1997. The burial was heavily disturbed by Mellaart’s earlier excavations in this building during the 1960s. Sk.2842 (burial feature 274, Level South M, Middle period, Building 50): a disturbed primary burial of a female infant aged 18 months (±6 months) at death based on dental development. It was excavated in 1998 from Building 50, located in the South Area of the site. The body was placed in a small pit near the center of the main room and was partially disturbed by a later burial. Radiocarbon dating of the petrous bone places this individual between 6690-6505 cal BCE (95%).” ref
“Sk.21981 (burial feature 8153, Level South N, Middle period, Building 89): a disturbed primary burial of a female infant/child aged 3 years (±1 year) at death based on dental development. It was excavated in 2015 from Building 89, located in the South Area of the site. The body was placed in a small pit within the north platform of the main room and was subsequently truncated by the digging of a post retrieval pit. Sk.5747 (burial feature 1064, Level South M, Middle period, Building 91): a primary burial of a female infant aged 18 months (±6 months) at death based on dental development. It was excavated in 2002 from Building 91, located in the South Area of the site. The body was placed in a small pit located in the northeast corner of B.91. Radiocarbon dating of the petrous bone places this individual between 6640-6490 cal BCE (95%) (Table Z2).” ref
“Sk.30006 (burial feature 7615, Level North G, Middle period, Building 114): a primary burial of a female infant aged 9 months (±3 months) at death based on dental development. It was excavated in 2015. The body was interred with a middle adult female in an oval pit along the south wall of the main room. Radiocarbon dating of the petrous bone places this individual between 6645–6495 cal BCE (94%) or 6490–6480 cal BCE (1%). Sk.8587 (burial feature 1013), Level North G, Middle period, Building 114): a primary burial of a female neonate (0-2 months at death – based on long bone length) excavated in 2002 and located under the southeast platform. The burial was partially disturbed by subsequent burials in this location, and likely also by rodent burrowing.” ref
“Sk.11739 (burial feature 1912, Level TP Q-R, Final period): a heavily disturbed set of human remains belonging to a middle adult (35-50 years of age-at-death) based on dental occlusal wear. The individual was assessed as a possible male based on cranial morphology, although aDNA suggested the individual was genetically female. These remains, potentially representing a secondary burial, were excavated in 2005 from Space 411, located in the TP Area of the site. Radiocarbon dating of the petrous bone places this individual between 6235-6075 cal BCE (95%).” ref
“Sk.20217 (burial feature 3931, Level TP Q-R, Final period?): a female child aged 6 years (±2 years) at death based on dental development. This individual, excavated in 2012, is one of three individuals recovered from burial feature 3931 in the TPC Area. The burial was badly damaged as it was found directly beneath the surface. Hence, it could not be associated with any Neolithic buildings or spaces. Its stratigraphic position indicates that it post-dates B.122 from the Late period, which implies it most likely comes from the Final period. However, this is not corroborated by radiocarbon dating of the petrous bone that places this individual significantly earlier, between 6415-6240 cal BCE (95%).” ref
“ZHAJ (Area H, Grave 27): this is a primary single inhumation of a middle adult female (as determined by aDNA) buried in a sub-oval cut. The individual was found lying tightly flexed on her left side, positioned east-west with the head toward the west and facing north. ZHAG and ZHAF (Area H, Grave 18): grave 18 contained a double inhumation of a middle adult female (ZHAF) and a perinatal infant (ZHAG) found in an oval cut larger than average. The adult (ZHAF) was found lying tightly flexed on her left side and positioned with a northwest-southeast orientation with the head toward the northwest. The perinate was articulated and found with the head on top of the adult pelvis.” ref
“The female sex of the adult could be confirmed by ancient DNA. The sex of the perinate was determined as a female by aDNA, and it can be ruled out that ZHAF and ZHAG were first or second-degree related. Skeleton ZHAF has been radiocarbon dated to 8285–8175 cal BCE (83%) or 8115–8090 cal BCE (4%) or 8040–8010 cal BCE (8%). ZHB (Area H, Grave 9): a single inhumation of an adolescent-young adult female. The sex of the individual has been confirmed by ancient DNA analysis. The individual was found lying on her right side/partially prone, in a semi-flexed position. The body was orientated east-west with head to the east and facing northeast, and has been radiocarbon dated to between 8280–8165 cal BCE (57%) or 8120–7960 cal BCE (38%).” ref
“ZHF (Area H, Grave 14): single inhumation of an adult male buried in a sub-oval cut. The age-at-death of the individual was difficult to estimate accurately because both the skull and pelvis were highly fragmented. The sex has been confirmed by ancient DNA. The body was found lying on the left side and orientated northwest-southeast with the head orientated toward the northwest and facing northeast. The upper limbs were flexed at the elbow with the palms of the hands together and placed immediately in front of the face. The long bones were highly fragmented and animal burrowing had destroyed much of the skull, most of the axial elements and the feet. The skeleton has been radiocarbon dated to 8225–7940 cal BCE (95%).” ref
“ZHJ (Area H, Grave 15): this is a primary single old adult inhumation found in a sub-oval cut. The individual was found in a flexed position lying on its right side and positioned north-south with the head orientated toward the south. The bones were relatively well preserved compared with other graves, although burrowing animals had destroyed parts of the skull and axial skeleton, including the left foot. Morphological sex determination was difficult because the remains were gracile, probably as a result of the aging process. Ancient DNA analyses demonstrated that this individual was female. She has been radiocarbon dated to 8295–8240 cal BCE (95%).” ref
“ZHBJ (Area H Grave 30): single inhumation of a middle/old adult male in a suboval cut. Sex has been confirmed by ancient DNA. The individual was found lying tightly flexed on his right side, although it should be noted that there was considerable damage from bioturbation that disturbed much of the skeleton and destroyed most of the thorax and skull. The body was positioned east-west with the head toward the west, but the facing direction could not be ascertained due to the aforementioned disturbance. ZKO (Area K, Grave 12): this is a single inhumation of an old adult male in an oval cut. The individual was found lying tightly flexed on his left side and orientated east-west with the head toward the east. The bones were generally well preserved, but rodent burrowing activity caused significant disturbance of the ribs, scapulae and vertebrae. Sex was confirmed through aDNA analysis as male.” ref
“ZMOJ (Area M, Grave 49): a primary but heavily disturbed burial of a young adult male (determined by aDNA) in a sub-circular grave. The individual was orientated east-west with head to the west and facing north. The skull was found at one end of the grave and many of the other bones had been moved by animal action, so their anatomical position was not maintained. Ancient DNA indicates that this individual was male.” ref
Aşıklı Höyük
“Aşıklı Höyük is a settlement mound located nearly 1 kilometre (0.62 mi) south of Kızılkaya village on the bank of the Melendiz brook, and 25 km (16 mi) southeast of Aksaray, Turkey. Aşıklı Höyük is located in an area covered by the volcanic tuff of central Cappadocia, in Aksaray Province. The archaeological site of Aşıklı Höyük was first settled in the Pre-Pottery Neolithic period, around 8,200 BCE. It is situated 1,119.5 metres (3,673 ft) above sea level, a little higher than the region’s average of c. 1,000 metres (3,300 ft). The site itself is about 4 hectares (9.9 acres), considerably smaller than the closely situated site of Çatalhöyük (13 ha (32 acres)). The surrounding landscape is formed by erosion of river valleys into tuff deposits. The Melendiz Valley, where the Aşıklı Höyük is located, constitutes a favourable, fertile, and diverse habitat. The proximity to an obsidian source did become the base of a trade with the material supplying areas as far away as today’s Cyprus and Iraq.” ref
“Aşıklı Höyük was first investigated by Professor Ian A. Todd when he visited the site in the summer of 1964. Todd emphasised the importance of the obsidian in the area, based on over 6,000 obsidian pieces collected from the surface layer alone. The site was classified as a medium sized mound and partly destroyed by the river situated next to it. On the basis of the lithics and animal bones located in the surface layers the site became known as a contemporary to the Palestine PPNB, which later was reinforced by 14C dates (based on five unstratified radiocarbon dates going from 7008 ± 130 to 6661 ± 108). The first comprehensive excavations took place relatively late: first when the government launched a plan that would result in the rise of the waters of the Mamasın Lake located close to Aşıklı Höyük, Professor Ufuk Esin (University of Istanbul) started the salvage excavations in 1989. Nine excavations have been undertaken up to 2003, uncovering approximately 4,200 square metres (45,000 sq ft) on the horizontal plain, making it one of the largest scale excavations in the region.” ref
“The newest dates for Aşıklı Höyük show that the occupational period was from 8200 to 7400 BCE or around 10,200 to 9,400 years ago, extracted from 3 layers with a total of 13 phases; which places it in phase ECA II (correlating with the E/MPPNB in the Levant). It is known as one of the earliest Aceramic Neolithic sites on the Anatolian plateau, and the prior mentioned extraction of the obsidian source was likely to be frequented as far back as the Paleolithic nomadic hunter-gatherers. Due to its date and structural organization Aşıklı Höyük is known to be “a prime example of a first foray into sedentism.” ref
“After more than 400 rooms had been excavated, the total number of individual found to have been buried within the settlement did not surpass 70. All these burials were under building floors. The dead were placed in pits cut through the floor during the occupation of the building. The buried are people of both sexes and all ages. There is a variety of skeletal body postures, from burials in a hocker (fetal) position to extended skeletons facing upwards. Others are lying on one side, occasionally with the legs bent at the knees. The orientation of the burials varies within the buildings, as does the number of individuals buried inside them.” ref
“The male population had individuals up to the age of 55–57 years of age, while the majority of females died between the ages of 20 and 25. The skeletal remains of these women show spinal deformities indicating that they had to carry heavy loads. This does not itself prove that there was a division of labour between the sexes. The fact that the men seem to have outlived the women might be interpreted as sign that the women were subject to more strenuous physical labour than their male counterparts. From Natufian Abu Hureyra there are similar osteological signs, such as pathologies in metatarsals, phalanges, arm, and shoulder joints, being specific to females resulting from habitual kneeling in the use of saddle querns (grinding stones). The Neolithic evidence show indications of increased physical workload in the osteological material on both genders, where the male skeletons show signs of joint disease and trauma arguably caused by cutting timber and tilling.” ref
“Children represent 37.8% of the deceased, with 43.7% mortality within a year of birth. The skeletal remains are complete and with articulations intact, indicating that the burials have been primary. The graves contain either single or double burials. On one occasion two graves were found under the floor of room AB, belonging to an adjacent court (HG) with a large domed mudbrick oven paved with blocks of basalt. In one of the graves were the skeletons of a young woman and an elderly man; in the other a young woman buried together with her baby. The young woman had apparently undergone trepanation and survived only a few days after the operation.” ref
“All skeletons were buried in the hocker position, a fetal-like positioning were the arms are embracing the lower limbs. From a different grave a woman shows signs of being scalped immediately after her death, according to the cut marks on her skull. As many as 55% of the skeletons show signs of being burned. The burial under the floor AB is accommodated by walls with the interior side were painted in a purplish red colour. The oven in HG indicates that this was indeed “special individuals of an elite class”, claiming it can be compared to the “Terrazzo” Building at Çayönü and the “Temple” Building at Nevalı Çori and therefore have been a shrine used for religious ceremonies. Many of the burials contain burial goods consisting of necklaces and bracelets made of beads of various sorts.” ref
“70 burials in over 400 rooms suggest that some form of selection took place of who was buried at the site, implying that AB indeed could be the residence or resting place of people influential in terms of both economy and political power. Rooms containing hearths are more likely to contain burials; as many as 77%. It has been argued that the number of burials could be an underrepresentation inhered at the site, since a large part of the settlement remain unexcavated beneath the baulks. Later excavations which have been published suggest on the other hand that burials were not a general feature at Aşıklı Höyük and therefore the suggestions of burials being a privilege of the elite class do seem plausible. There has not been found a cemetery or any other sign of where the rest of the population might have been disposed of post mortem. This issue is not only limited to Aşıklı Höyük: there is also a lack of cemeteries on the PPNB “mega-sites” in the Levant, such ‘Ain Ghazal in the Jordan Valley.” ref
“It seems that in Aşıklı Höyük, as in the rest of the Anatolian and Levantine area, the burial and any other post mortem treatment was arguably an “upper class” phenomenon. This interpretation has been opposed, referring to the diversity of individuals in both sex and age in the graves. The burials including such a wide range of individuals do not directly coherent with the image of an “upper class” phenomenon. Burials could have been removed or replaced over time, giving a wrong image of the burials as belonging to the elite. An alternative perspective has been suggested: “the Neolithic dead are not under-represented: rather, it is the architecture in settlements that are over-represented”, meaning that in many cases archaeologists have drastically overestimated the extent to which all areas of Neolithic sites were occupied simultaneously. As for Aşıklı Höyük and other sites in the area: low numbers of burials in comparison with occupation span does not directly indicate a cult of the elite.” ref
“At Aşıklı Höyük the hearths are rectangular and usually placed in one of the corners of the rooms, ranging in size from 2.97 to 0.48 square metres (32.0 to 5.2 sq ft). Large stones with a suitable flat shape were used to create an upright edge that stood approximately 20 cm (7.9 in) above the level of the floor. On the short side of the hearth the upright edge is missing to make a fire mouth. It is also here the ash is the most concentrated. Pebbles along the edges and base of the hearth seem to have been covered by a thin layer of plaster. Only in a few cases there is a trace of something recognized as a flue. An estimated 30–40% of all the rooms at Aşıklı Höyük have hearths. This estimate is based on partially damaged and eroded structures possibly giving a number lower than accurate. Based on a limited group of fully exposed buildings 54% of single room units contained a hearth, while only 29% of the multiple room units. The average percentage on base of these building units is 47%, probably a more realistic estimate for the site in total.” ref
“Hearths do not occur in a courtyard context, and are more represented in single room dwellings than multiple room units. Still, multiple room units do have a substantial number of hearths. It has been suggested that the “multiroom dwellings may have functioned as ‘incomplete’ houses for new families still heavily dependent on the larger extended group”. The buildings containing the hearths do not show particular characteristics that distinguish them from structures without hearths; neither do they differ in size or special orientation. Even the hearth itself does not follow an apparent norm in terms of size or location. The position varies considerably, but it always has one side to the wall. The positioning of the hearth does not seem to be determined by general macro-ecological factors, such as prevailing wind directions, nor determined by cultural norms regarding spatial features within buildings.” ref
“The hearth does not seem to be subject to a random placement inside the buildings: it is consistently located at the same spot throughout a very long building sequence. This indicates that their positions were not chosen arbitrarily. When a location for the hearth was chosen it was important that the placement did not change in later rebuilding sequences of the structure (see picture: Deep sounding). There is no evidence for ladderpost scars due to the assumed use of freestanding ladders, making the location of the entrance uncertain. Aşıklı Höyük does not seem to have any evidence for ovens. Aşıklı Höyük had a tradition to reconstruct or rebuild earlier structures. It followed a pattern where the structures were built “exactly on the same spot and with the same alignment as earlier buildings, using older walls as a foundation.” ref
“The structural continuity at Aşıklı Höyük is outstanding, but there is no information to how long the use-life of a building was. If one estimates the same lifespan for a structure at Aşıklı Höyük as it was in Çatalhöyük, one could look at an age of 30 to 60 years before reconstruction occurred. If this assumption is correct, the deep sounding 4H/G from phases 2I up to 2B (eight layers in total) show that the time span of a structure could be from 240 to 480 years. Looking outside of Central Anatolian Neolithic, this type of building continuity is unparalleled both in ethnography and archaeology. This remarkable structural continuity may suggest a social system in which buildings were not privately owned, since one would expect them to be modified on a regular basis. It can be assumed that the rooms were distributed amongst the community members according to the change in both needs and statuses.” ref
“The building practices maintained their characteristics throughout the centuries. It has been claimed that the building continuity is a self-evident feature, since it is deriving from a particular set of foundation practices that can be explained in a functionalistic way. It has otherwise been argued that the extreme degree of continuity is inadequately explained by functionalism alone, since the structures located adjacent to open spaces could have easily been expanded or shrunk according to the specific needs, but instead remained identical. These functionalist parameters can also not explain the continued rebuilding of the hearths, which are always built on the same spot. Individual hearth sequences are often separated with 40 cm (16 in) of soil, and therefore there is no apparent reason (unlike the buildings) why the hearths should consistently be constructed in the same corner as in the successive buildings. In many cases neighbouring buildings do place their respective hearths in different corners. Micro-cosmological special codes or wind direction does not seem to be decisive for the positioning of the hearth. The structural and material remains indicate that the buildings were continuous entities with some form of fixed special identity where the special organization could not be changed by the temporary occupant.” ref
“Structural continuity was of great importance to the inhabitants of Aşıklı Höyük. The reason for this has partly been explained because they (the people) had a rigid adherence to traditions in terms of structural reproduction. The “traditional view” has been disputed because “[In short,] labelling a society as conservative does not answer the question why the people under consideration were conservative”. An alternative approach refers to the historical dimension of the building to be of such great importance that “people came to be bound between walls, metaphorically domesticated”. The interpretation is that the walls are giving historical associations to the people living within them, giving a collective conscience lasting through time. The difference between this interpretation and the “conservative approach” is the potential explanation to why structural reproduction could have been important for the inhabitants of Aşıklı Höyük. The identities of the inhabitants were projected to the structural outcome of the buildings. The generality of this position is not meaningful on its own. It is not certain that the inhabitants of Aşıklı Höyük were aware of the total amount of building there was in the sequence in total. Perhaps the history of a building did not concern them in the same way as archaeologists like to think.” ref
“The lack of change over time suggests that the inhabitants of Aşıklı Höyük had a view of the past as a precedent for the present: a vital part of society that was ‘reborn’ in each reproduction, manifested in its building continuity. The structural reconstruction is a regional feature for Central Anatolia. With the exception of Jericho, most of the evidence from PPNB sites in the Levant indicates that structures were not reconstructed in the same loci, and some location structures differ in dates by several hundred years. The buildings at Aşıklı Höyük are clustered into what has been interpreted as neighbourhoods. As this is a vague perception of the structural outlay of the community it describes them as clustered single and multiroom houses forming compounds, apparently sharing courtyard space for production activities and practising joint cooking and food consumption. Little can be said on the food storage, since there were no remains after storage bins, although storage rooms may be identified due to comparing structures on other sites (e.g. Çatalhöyük).” ref
“The average room size is 12 m2 (130 sq ft) (at this time humans were 1.5 to 1.7 metres (4 ft 11 in to 5 ft 7 in) tall). From two or three up to five or six clustered dwellings formed a ‘neighborhood’ or compound. The interpretation of the borders of these ‘neighbourhoods’ is problematic, since much of the site still lies under the baulks, is in situ or eroded. The distribution of single- and multi-room buildings does not seem to follow a pattern other than that the residential clusters seem to be divided by narrow alleys 0.5–1.0 m (1 ft 8 in – 3 ft 3 in) wide, or open courtyard areas up to a diameter of 4 m (13 ft). The interior of multiroom buildings had openings in the partitioned walls, providing access to the individual rooms. Between the one building and the next there seems to be no communication, since there was no indication of doors in the exterior mudbrick walls. Since the buildings themselves do not have an entrance that can be traced archaeologically on the base of the walls, access had to be provided either through window-like openings high on the walls or from the flat roofs. Roof access is also known from Çatalhöyük, making this entrance more plausible.” ref
“Aşıklı Höyük does also have buildings that are bigger in size but without hearths. These are interpreted as public buildings or ‘building complexes’. These are seen as some of the most enigmatic buildings found at the site, and diverge both in size and spatial organization. One of them (complex HV) being at up to 20 times larger than the largest loam buildings (i.e. 25 m × 20 m (82 ft × 66 ft) = 500 m2 (5,400 sq ft)). They have a multitude of rooms and encompass elaborate and large internal courts; something that is not found in any other buildings. The walls are more robust and massive than other buildings, in some cases being referred to as “monumental walls”, accompanied by parallel outer walls with relatively narrow space in between.” ref
“The interpretation of these buildings is difficult. The fact that they clearly differentiate from the domestic loam buildings indicates that they had special value in the society. They also do not incorporated into the clustered ‘neighbourhoods’, indicating that they served several neighbourhoods or the local community at large. With 500 m2 (5,400 sq ft) the range of activities that could have taken place in this space could easily incorporate several hundred people. Yet, given that the estimated population of Aşıklı Höyük may have run into the thousands, only a selected group in the total population could have used the building at a given occasion. There is a variety of hypotheses regarding the nature of these monumental structures. There are other examples of these restricted monumental spaces on other sites in the Levantine PPNB (such as Nevali Çori, Behida, ‘Ain Ghazal), suggesting that they were used by an elite or for practising different social initiation rites.” ref
“There are no finds of any artefacts carrying religious connotations, symbolic or imagery, at Aşıklı Höyük, in the buildings, courtyards, dumps or open-workshop areas. The only finds include flint tools, which are counted as imports. Other than this there is found one single animal figurine made of clay that can hardly tell us anything of the religious belief of the inhabitants. The limited amount of burials compared to the estimated population makes it very likely that there may have been a cemetery where the deceased were buried, but it has not been found. There is also an absence of storage bins, making the distinction on autonomous households difficult.” ref
Samara, Russia 7,500 years ago carried R1b1a1a
“An EHG buried near Samara, Russia 7,500 years ago carried R1b1a1a. An Eneolithic male buried at Khvalynsk, Russia 7,200-6,000 years ago carried R1b1a.” ref
R1a was not found in Yamnaya remains
“The most common Y-DNA haplogroup found among the Yamnaya specimens was Haplogroup R1b, specifically the Z2103 subclade of R1b-L23. Additionally, a minority are found to belong to haplogroup I2. Autosomal tests also indicate that the Yamnaya are the vector for “Ancient North Eurasian” admixture into Europe. “Ancient North Eurasian” is the name given in literature to a genetic component that represents descent from the people of the Mal’ta–Buret’ culture or a population closely related to them. That genetic component is visible in tests of the Yamnaya people as well as modern-day Europeans.” ref, ref, ref
“Mascarenhas et al. (2015) proposed that the roots of Z93 lie in West Asia, and proposed that “Z93 and L342.2 expanded in a southeasterly direction from Transcaucasia into South Asia“, noting that such an expansion is compatible with “the archeological records of eastward expansion of West Asian populations in the 4th millennium BCE culminating in the so-called Kura-Araxes
Kunda culture
“The earliest traces of human settlement in Estonia are connected with the Kunda culture. The early Mesolithic Pulli settlement is located by the Pärnu River. It has been dated to the beginning of the 9th millennium BCE. The Kunda culture received its name from the Lammasmäe settlement site in northern Estonia, which dates from earlier than 8500 BCE. Bone and stone artifacts similar to those found at Kunda have been discovered elsewhere in Estonia, as well as in Latvia, northern Lithuania, and southern Finland. Among minerals, flint and quartz were used the most for making cutting tools. The beginning of the Neolithic Period is marked by the ceramics of the Narva culture, which appeared in Estonia at the beginning of the 5th millennium BCE. The oldest finds date from around 4900 BCE.” ref
“The Kunda culture, which originated from the Swiderian culture, comprised Mesolithic hunter-gatherer communities of the Baltic forest zone extending eastwards through Latvia into northern Russia, dating to the period 8500–5000 BCE according to calibrated radiocarbon dating. It is named after the Estonian town of Kunda, about 110 kilometres (70 mi) east of Tallinn along the Gulf of Finland, near where the first extensively studied settlement was discovered on Lammasmäe Hill and in the surrounding peat bog. The oldest known settlement of the Kunda culture in Estonia is Pulli. The Kunda culture was succeeded by the Narva culture, who used pottery and showed some traces of food production.” ref
“Most Kunda settlements are located near the edge of the forests beside rivers, lakes, or marshes. Elk were extensively hunted, perhaps helped by trained domestic hunting-dogs. On the coast seal hunting is represented. Pike and other fish were taken from the rivers. There is a rich bone and antler industry, especially in relation to fishing gear. Tools were decorated with simple geometric designs, lacking the complexity of the contemporary Maglemosian Culture communities to the southwest. Jones et al. (2017) determined, based on one sample (6467-6250 BCE) from the Kunda culture and another one from the succeeding Narva culture, closer genetic affinity with Western Hunter-Gatherers (WHGs) than Eastern Hunter-Gatherers (EHGs).” ref
“The Kunda culture appears to have undergone a transition from the Palaeolithic Swiderian culture located previously over much of the same range. One such transition settlement, Pasieniai 1C in Lithuania, features stone tools of both Late Swiderian and early Kunda. One shape manufactured in both cultures is the retouched tanged point. The final Swiderian is dated 7800–7600 BCE by calibrated radiocarbon dating, which is in the Preboreal period, at the end of which time with no gap the early Kunda begins. Evidently, the descendants of the Swiderians were the first to settle Estonia when it became habitable. Other post-Swiderian groups extended as far east as the Ural Mountains. Mittnik et al. (2018) analyzed the remains of a male and female ascribed to the Kunda culture. They found the male to be carrying paternal haplogroup I and maternal haplogroup U5b2c1, while the female carried U4a2.” ref
“They were found to have “a very close affinity” with WHGs, although with “a significant contribution” from Ancient North Eurasians (ANE). Their ANE ancestry was lower than that of Scandinavian Hunter-Gatherers, indicating that ANE ancestry entered Scandinavia without traversing the Baltic. Matthieson et al. (2018) analyzed a large number of individuals buried at the Zvejnieki burial ground, most of whom were affiliated with the Kunda culture and the succeeding Narva culture. The mtDNA belonged to haplotypes U5, U4 and U2, the vast majority of the Y-DNA samples belonged to R1b1a1a and I2a1. The results affirmed that the Kunda and Narva cultures were about 70% WHG and 30% EHG. The nearby contemporary Pit–Comb Ware culture was on the contrary found to be about 65% EHG. Around 3700/3600 BC there was a complete turnover to now y-hg R1a1a1, from 900/800 onward mainly in Estonia with increasing impact of y-hg N-M231 (N1a1).” ref
“The first pottery was made of thick clay mixed with pebbles, shells, or plants. The Narva-type ceramics are found throughout almost the entire Estonian coastal region and on the islands. The stone and bone tools of the era have a notable similarity with the artifacts of the Kunda culture. Around the beginning of the 4th millennium BCE, Comb Ceramic culture arrived in Estonia. Until the early 1980s, the arrival of Finnic peoples, the ancestors of the Estonians, Finns, and Livonians, on the shores of the Baltic Sea was associated with the Comb Ceramic Culture. However, such a linking of archaeologically defined cultural entities with linguistic ones cannot be proven, and it has been suggested that the increase of settlement finds in the period is more likely to have been associated with an economic boom related to the warming of the climate. Some researchers have even argued that a Uralic form of language may have been spoken in Estonia and Finland since the end of the last glaciation.” ref
“The burial customs of the comb pottery people included additions of figures of animals, birds, snakes, and men carved from bone and amber. Antiquities from comb pottery culture are found from Northern Finland to Eastern Prussia. The beginning of the Late Neolithic Period, about 2200 BCE, is characterized by the appearance of the Corded Ware culture, pottery with corded decoration, and well-polished stone axes (s.c. boat-shape axes). Evidence of agriculture is provided by charred grains of wheat on the wall of a corded-ware vessel found in the Iru settlement. Osteological analysis shows an attempt was made to domesticate the wild boar. Specific burial customs were characterized by the dead being laid on their sides with their knees pressed against their breast, one hand under the head. Objects placed into the graves were made of the bones of domesticated animals.” ref
“The beginning of the Bronze Age in Estonia is dated to approximately 1800 BCE. The development of the borders between the Finnic peoples and the Balts was under way. The first fortified settlements, Asva and Ridala on the islands of Saaremaa and Iru in Northern Estonia, began to be built. The development of shipbuilding facilitated the spread of bronze. Changes took place in burial customs, a new type of burial ground spread from Germanic to Estonian areas, and stone cist graves and cremation burials became increasingly common, alongside a small number of boat-shaped stone graves. About the 7th century BCE or around 2,700 years ago, a large meteorite hit Saaremaa island and created the Kaali craters. About 325 BCE, the Greek explorer Pytheas possibly visited Estonia. The Thule island he described has been identified as Saaremaa by Lennart Meri, though this identification is not widely considered probable, as Saaremaa lies far south of the Arctic Circle.” ref
“The Pre-Roman Iron Age began in Estonia about 500 BC and lasted until the middle of the 1st century CE. The oldest iron items were imported, although since the 1st century iron was smelted from local marsh and lake ore. Settlement sites were located mostly in places that offered natural protection. Fortresses were built, although used temporarily. The appearance of square Celtic fields surrounded by enclosures in Estonia date from the Pre-Roman Iron Age. The majority of stones with man-made indents, which presumably were connected with magic designed to increase crop fertility, date from this period. A new type of grave, quadrangular burial mounds, began to develop. Burial traditions show the clear beginning of social stratification.” ref
Hunter-Gatherer Pottery?
“Pottery is one of the oldest human inventions, originating before the Neolithic period, with ceramic objects such as the Gravettian culture Venus of Dolní Věstonice figurine discovered in the Czech Republic dating back to 29,000–25,000 BCE. However, the earliest known pottery vessels were discovered in Jiangxi, China, which date back to 18,000 BC. Other early Neolithic and pre-Neolithic pottery artifacts have been found, in Jōmon Japan (10,500 BCE), the Russian Far East (14,000 BCE), Sub-Saharan Africa (9,400 BCE), South America (9,000s–7,000s BCE), and the Middle East (7,000s–6,000s BCE).” ref
“The earliest forms of pottery were made from clays that were fired at low temperatures, initially in pit-fires or in open bonfires. They were hand-formed and undecorated. Earthenware can be fired as low as 600 °C, and is normally fired below 1200 °C. Because unglazed earthenware is porous, it has limited utility for the storage of liquids or as tableware. However, earthenware has had a continuous history from the Neolithic period to today. It can be made from a wide variety of clays, some of which fire to a buff, brown or black colour, with iron in the constituent minerals resulting in a reddish-brown. Reddish coloured varieties are called terracotta, especially when unglazed or used for sculpture. The development of ceramic glaze made impermeable pottery possible, improving the popularity and practicality of pottery vessels. Decoration has evolved and developed through history.” ref
“Stoneware is pottery that has been fired in a kiln at a relatively high temperature, from about 1,100 °C to 1,200 °C, and is stronger and non-porous to liquids. The Chinese, who developed stoneware very early on, classify this together with porcelain as high-fired wares. In contrast, stoneware could only be produced in Europe from the late Middle Ages, as European kilns were less efficient, and the right type of clay less common. It remained a speciality of Germany until the Renaissance. Stoneware is very tough and practical, and much of it has always been utilitarian, for the kitchen or storage rather than the table. But “fine” stoneware has been important in China, Japan and the West, and continues to be made. Many utilitarian types have also come to be appreciated as art.” ref
“The study of pottery can help to provide an insight into past cultures. Fabric analysis (see section below), used to analyze the fabric of pottery, is an important part of archaeology for understanding the archaeological culture of the excavated site by studying the fabric of artifacts, such as their usage, source material composition, decorative pattern, color of patterns, etc. This helps to understand the characteristics, sophistication, habits, technology, tools, trade, etc., of the people who made and used the pottery. Carbon dating reveals the age. Sites with similar pottery characteristics have the same culture; those sites that have distinct cultural characteristics but with some overlap are indicative of cultural exchange, such as trade or living in the vicinity or continuity of habitation, etc. Examples are black and red ware, redware, Sothi-Siswal culture, and Painted Grey Ware culture. The six fabrics of Kalibangan is a good example of use of fabric analysis in identifying a differentiated culture which was earlier thought to be typical Indus Valley civilisation (IVC) culture.” ref
“Pottery is durable, and fragments, at least, often survive long after artifacts made from less-durable materials have decayed past recognition. Combined with other evidence, the study of pottery artifacts is helpful in the development of theories on the organization, economic condition, and cultural development of the societies that produced or acquired pottery. The study of pottery may also allow inferences to be drawn about a culture’s daily life, religion, social relationships, attitudes towards neighbors, attitudes to their own world, and even the way the culture understood the universe. It is valuable to look into pottery as an archaeological record of potential interaction between peoples. When pottery is placed within the context of linguistic and migratory patterns, it becomes an even more prevalent category of social artifact.” ref
“ Sherds have been found in China and Japan from a period between 12,000 and perhaps as long as 18,000 years ago. As of 2012, the earliest pottery vessels found anywhere in the world, dating to 20,000 to 19,000 years before the present, was found at Xianren Cave in the Jiangxi province of China. Other early pottery vessels include those excavated from the Yuchanyan Cave in southern China, dated from 16,000 BCE, and those found in the Amur River basin in the Russian Far East, dated from 14,000 BCE. The Odai Yamamoto I site, belonging to the Jōmon period, currently has the oldest pottery in Japan. Excavations in 1998 uncovered earthenware fragments which have been dated as early as 14,500 BCE. The term “Jōmon” means “cord-marked” in Japanese. This refers to the markings made on the vessels and figures using sticks with cords during their production. Recent research has elucidated how Jōmon pottery was used by its creators.” ref
“It appears that pottery was independently developed in Sub-Saharan Africa during the 10th millennium BCE, with findings dating to at least 9,400 BCE from central Mali, and in South America during the 9,000s–7,000s BCE. The Malian finds date to the same period as similar finds from East Asia – the triangle between Siberia, China and Japan – and are associated in both regions to the same climatic changes (at the end of the ice age new grassland develops, enabling hunter-gatherers to expand their habitat), met independently by both cultures with similar developments: the creation of pottery for the storage of wild cereals (pearl millet), and that of small arrowheads for hunting small game typical of grassland. Alternatively, the creation of pottery in the case of the Incipient Jōmon civilisation could be due to the intensive exploitation of freshwater and marine organisms by late glacial foragers, who started developing ceramic containers for their catch.” ref
“Around 8000 BCE during the Pre-pottery Neolithic period, and before the invention of pottery, several early settlements became experts in crafting beautiful and highly sophisticated containers from stone, using materials such as alabaster or granite, and employing sand to shape and polish. Artisans used the veins in the material to maximum visual effect. Such objects have been found in abundance on the upper Euphrates river, in what is today eastern Syria, especially at the site of Bouqras. The earliest history of pottery production in the Fertile Crescent starts the Pottery Neolithic and can be divided into four periods, namely: the Hassuna period (7000–6500 BCE), the Halaf period (6500–5500 BCE), the Ubaid period (5500–4000 BCE), and the Uruk period (4000–3100 BCE). By about 5000 BCE pottery-making was becoming widespread across the region, and spreading out from it to neighbouring areas. Pottery making began in the 7th millennium BC. The earliest forms, which were found at the Hassuna site, were hand formed from slabs, undecorated, unglazed low-fired pots made from reddish-brown clays.” ref
“Within the next millennium, wares were decorated with elaborate painted designs and natural forms, incising and burnished. The invention of the potter’s wheel in Mesopotamia sometime between 6,000 and 4,000 BC (Ubaid period) revolutionised pottery production. Newer kiln designs could fire wares to 1,050 °C (1,920 °F) to 1,200 °C (2,190 °F) which enabled increased possibilities. Production was now carried out by small groups of potters for small cities, rather than individuals making wares for a family. The shapes and range of uses for ceramics and pottery expanded beyond simple vessels to store and carry to specialized cooking utensils, pot stands and rat traps. As the region developed new organizations and political forms, pottery became more elaborate and varied. In the Chalcolithic period in Mesopotamia, Halafian pottery achieved a level of technical competence and sophistication, not seen until the later developments of Greek pottery with Corinthian and Attic ware.” ref
“Europe’s oldest pottery, dating from circa 6700 BCE, was found on the banks of the Samara River in the middle Volga region of Russia. These sites are known as the Yelshanka culture. The early inhabitants of Europe developed pottery in the Linear Pottery culture slightly later than the Near East, circa 5500–4500 BCE. In the ancient Western Mediterranean elaborately painted earthenware reached very high levels of artistic achievement in the Greek world; there are large numbers of survivals from tombs. Minoan pottery was characterized by complex painted decoration with natural themes. The classical Greek culture began to emerge around 1000 BC featuring a variety of well crafted pottery which now included the human form as a decorating motif.” ref
“The pottery wheel was now in regular use. Although glazing was known to these potters, it was not widely used. Instead, a more porous clay slip was used for decoration. A wide range of shapes for different uses developed early and remained essentially unchanged during Greek history. Fine Etruscan pottery was heavily influenced by Greek pottery and often imported Greek potters and painters. Ancient Roman pottery made much less use of painting, but used moulded decoration, allowing industrialized production on a huge scale. Much of the so-called red Samian ware of the Early Roman Empire was produced in modern Germany and France, where entrepreneurs established large potteries.” ref
Narva culture (First European Pottery?)
“Preceded by the Kunda culture, and followed by the Pit–Comb Ware culture, Corded Ware culture, and Brushed Pottery culture. The Narva culture or eastern Baltic was a European Neolithic archaeological culture in present-day Estonia, Latvia, Lithuania, Kaliningrad Oblast (former East Prussia), and adjacent portions of Poland, Belarus, and Russia. A successor of the Mesolithic Kunda culture, the Narva culture continued up to the start of the Bronze Age. The culture spanned from c. 5300 to 1750 BCE. The technology was that of hunter-gatherers. The culture was named after the Narva River in Estonia.” ref
“The people of the Narva culture had little access to flint; therefore, they were forced to trade and conserve their flint resources. For example, there were very few flint arrowheads and flint was often reused. The Narva culture relied on local materials (bone, horn, schist). As evidence of trade, researchers found pieces of pink flint from Valdai Hills and plenty of typical Narva pottery in the territory of the Neman culture while no objects from the Neman culture were found in Narva. Heavy use of bones and horns is one of the main characteristics of the Narva culture. The bone tools, continued from the predecessor Kunda culture, provide the best evidence of continuity of the Narva culture throughout the Neolithic period. The people were buried on their backs with few grave goods. The Narva culture also used and traded amber; a few hundred items were found in Juodkrantė. One of the most famous artifacts is a ceremonial cane carved of horn as a head of female elk found in Šventoji.” ref
“The people were primarily fishers, hunters, and gatherers. They slowly began adopting husbandry in the middle Neolithic. They were not nomadic and lived in the same settlements for long periods as evidenced by abundant pottery, middens, and structures built in lakes and rivers to help fishing. The pottery shared similarities with the Comb Ceramic culture, but had specific characteristics. One of the most persistent features was mixing clay with other organic matter, most often crushed snail shells. The pottery was made of 6-to-9 cm (2.4-to-3.5 in) wide clay strips with minimal decorations around the rim. The vessels were wide and large; the height and the width were often the same. The bottoms were pointed or rounded, and only the latest examples have narrow flat bottoms. From mid-Neolithic, Narva pottery was influenced and eventually disappeared into the Corded Ware culture.” ref
“The prehistoric shell middens of Atlantic Europe consist of marine molluscs, but the eastern Baltic did not have exploitable marine species. Here the sole recorded shell midden, at Riņņukalns in Latvia, is on an inland lake and is formed of massive dumps of freshwater shells. Recent excavations indicate that they are the product of a small number of seasonal events during the later fourth millennium BCE. The thickness of the shell deposits suggests that this was a special multi-purpose residential site visited for seasonal aggregations by pottery-using hunter-gatherer communities on the northern margin of Neolithic Europe.” ref
“For a long time, archaeologists believed that the first inhabitants of the region were Finno-Ugric, who were pushed north by people of the Corded Ware culture. In 1931, Latvian archaeologist Eduards Šturms was the first to note that artifacts found near the Zebrus Lake in Latvia were different and possibly belonged to a separate archaeological culture. In early 1950s settlements on the Narva River were excavated. Lembit Jaanits and Nina Gurina grouped the findings with similar artifacts from eastern Baltic region and described the Narva culture. At first, it was believed that Narva culture ended with the appearance of the Corded Ware culture.” ref
“However, newer research extended it up to the Bronze Age. As Narva culture spanned several millennia and encompassed a large territory, archaeologists attempted to subdivide the culture into regions or periods. For example, in Lithuania two regions are distinguished: southern (under influence of the Neman culture) and western (with major settlements found in Šventoji). There is an academic debate what ethnicity the Narva culture represented: Finno-Ugrians or other Europids, preceding the arrival of the Indo-Europeans. It is also unclear how the Narva culture fits with the arrival of the Indo-Europeans (Corded Ware and Globular Amphora cultures) and the formation of the Baltic tribes.” ref
“Mathieson (2015) analyzed a large number of individuals buried at the Zvejnieki burial ground, most of whom were affiliated with the Kunda culture and the succeeding Narva culture. The mtDNA extracted belonged exclusively to haplotypes of U5, U4, and U2. With regards to Y-DNA, the vast majority of samples belonged to R1b1a1a haplotypes and I2a1 haplotypes. The results affirmed that the Kunda and Narva cultures were about 70% WHG and 30% EHG. The nearby contemporary Pit–Comb Ware culture was, on the contrary, found to be about 65% EHG. An individual from the Corded Ware culture, which would eventually succeed the Narva culture, was found to have genetic relations with the Yamnaya culture.” ref
“Jones et al. (2017) examined the remains of a male of the Narva culture buried c. 5780-5690 BCE. He was found to be a carrier of the paternal haplogroup R1b1b and the maternal haplogroup U2e1. People of the Narva culture and preceding Kunda culture were determined to have closer genetic affinity with Western Hunter-Gatherers (WHGs) than Eastern Hunter-Gatherers (EHGs). Saag et al. (2017) determined haplogroup U5a2d in a Narva male. Mittnik et al. (2018) analyzed 24 Narva individuals. Of the four samples of Y-DNA extracted, one belonged to I2a1a2a1a, one belonged to I2a1b, one belonged to I, and one belonged to R1. Of the ten samples of mtDNA extracted, eight belonged to U5 haplotypes, one belonged to U4a1, and one belonged to H11. U5 haplotypes were common among Western Hunter-Gatherers (WHGs) and Scandinavian Hunter-Gatherers (SHGs). Genetic influence from Eastern Hunter-Gatherers (EHGs) was also detected.” ref
Pitted Ware culture
“The Pitted Ware culture (c. 3500 BC–c. 2300 BCE) was a hunter-gatherer culture in southern Scandinavia, mainly along the coasts of Svealand, Götaland, Åland, north-eastern Denmark and southern Norway. Despite its Mesolithic economy, it is by convention classed as Neolithic, since it falls within the period in which farming reached Scandinavia. The Pitted Ware people were largely maritime hunters, and were engaged in lively trade with both the agricultural communities of the Scandinavian interior and other hunter-gatherers of the Baltic Sea. The people of the Pitted Ware culture were a genetically homogeneous and distinct population descended from earlier Scandinavian Hunter-Gatherers (SHGs). The culture emerged in east-central Sweden around 3,500 BC, gradually replacing the Funnelbeaker culture throughout the coastal areas of southern Scandinavia. It subsequently co-existed with the Funnelbeaker culture for several centuries.” ref
“From about 2,800 BCE, the Pitted Ware culture co-existed with the Battle Axe culture, which was the successor of the Funnelbeaker culture in southern Scandinavia. By 2,300 BCE, the Pitted Ware culture had been absorbed by the Battle Axe culture. The subsequent Nordic Bronze Age represents a fusion of elements from the Pitted Ware culture and the Battle Axe culture. Modern Scandinavians, unlike the Sami, display partial genetic origins from the Pitted Ware people. Genetic studies suggest that the Pitted Ware peoples, unlike their Neolithic neighbors, were descended from earlier Scandinavian Hunter-Gatherers (SHGs). At the time of the emergence of the Pitted Ware culture, these hunter-gatherers persisted to the north of the agricultural Funnelbeaker culture. Their ceramic traditions are related to those of the Comb Ceramic culture. Because of this, Pitted Ware Culture is often combined with Comb Ceramic Culture to form Pit Comb Ware Culture.” ref
“The Pitted Ware culture arose around 3,500 BCE. Its earliest sites are found in east-central Sweden, where it appears to have replaced the Funnelbeaker culture. Its subsequent expansion is accompanied by the disappearance of settlements of the Funnelbeaker culture throughout large parts of southern Scandinavia. It came to occupy the coasts of Denmark, southern Sweden, southern Norway and various islands of the Baltic Sea, such as Öland, Gotland, and Åland. There were lively contacts with hunter-gatherer communities of Finland and the eastern Baltic. During its initial years, the Pitted Ware culture co-existed with the Funnelbeaker culture. Although the two cultures exchanged goods with each other, its peoples appear to have had widely different identities, and they did not mix with each other to any notable extent. During the period of Pitted Ware expansion, the Funnelbeakers constructed a number of defensive palisades, which may mean that the two peoples were in conflict with each other. There is archaeological evidence of high levels of violence among the people of the Pitted Ware culture. Throughout its existence of more than 1,000 years, the Pitted Ware culture remained virtually unchanged.” ref
“From around 2,800 BCE, the Pitted Ware culture co-existed for some time with the Battle Axe culture and the Single Grave culture, which succeeded the Funnelbeaker culture in southern Scandinavia. Both were variants of the Corded Ware culture. Like the Funnelbeakers, the Corded Ware constructed a series of defensive palisades during this period, which may be a sign of violent conflict between them and the Pitted Ware. Though cultural influences of the Battle Axe culture are detectable in Pitted Ware burials, its peoples do not appear to have mixed with each other. By ca. 2,300 BCE, the Pitted Ware culture had merged with the Battle Axe culture. The subsequent Nordic Bronze Age represents a fusion of elements from the Pitted Ware culture and the Battle Axe culture.” ref
“The economy of the Pitted Ware culture was based on fishing, hunting and gathering of plants. Pitted Ware sites contain bones from elk, deer, beaver, seal, porpoise, and pig. Pig bones found in large quantities on some Pitted Ware sites emanate from wild boar rather than domestic pigs. The hunting of seal was particularly important. For this reason, the Pitted Ware people have been called “hard-core sealers” or the “Inuit of the Baltic”. Seasonal migration was a feature of life, as with many other hunter-gatherer communities. Pitted Ware communities in Eastern Sweden probably spent most of the year at their main village on the coast, making seasonal forays inland to hunt for pigs and fur-bearing animals and to engage in exchange with farming communities in the interior.” ref
“This type of seasonal interaction may explain the unique Alvastra Pile Dwelling in south-western Östergötland, which belongs to the Pitted Ware culture as far as the pottery is concerned, but to the Funnelbeaker culture in tools and weapons. The Pitted Ware peoples appear to have been specialized hunters who engaged in the trade of animal goods with peoples throughout the Baltic. The repertoire of Pitted Ware tools varied from region to region. In part this variety reflected regional sources of raw materials. However, the use of fish hooks, harpoons, nets, and sinkers was fairly widespread. Tanged arrow heads made from blades of flintstone are abundant on Scandinavia’s west coast, and were probably used in the hunting of marine mammals.” ref
“One notable feature of the Pitted Ware Culture is the sheer quantity of shards of pottery on its sites. The culture has been named after the typical ornamentation of its pottery: horizontal rows of pits pressed into the body of the pot before firing. Though some vessels are flat-bottomed, others are round-based or pointed-based, which would facilitate stable positioning in the soil or on the hearth. In shape and decoration, these ceramics reflect influences from the Comb Ceramic culture (also known as Pit-Comb Ware) of Finland and other parts of north-eastern Europe, established in the sixth and fifth millennia BCE. Small animal figurines were modelled from clay, as well as bone. These are also similar to the art of the Comb Ware culture. A large number of ceramic figurines have been found at Jettböle on the island of Jomala in Åland, including some which combine seal and human features.” ref
“The Pitted Ware people buried their dead in cemeteries. Most excavated Pitted Ware burials are located at Gotland, where around 180 graves have been found at numerous sites with several layers. One such site is at Västerbjers. Pitted Ware people were typically buried in flat inhumation graves, although cremation does occur. Unlike the Funnelbeakers, they did not have megalithic graves. Pitted Ware burials are also distinguished from Funnelbeaker burials through their use of red ochre. Grave goods include ceramics, boar tusks, pig jaws, pendants of fox, dog and seal teeth, harpoons, spears, fishhooks of bone, stone and flint axes, and other artifacts. The presence of slate artifacts and battle axes attest wide-ranging contacts between the Pitted Ware people and other cultures of Northern Europe and the Baltic. People of all ages and genders were buried in the same cemetery. There are no indications of difference in social status. Their mortuary houses and secondary burials are nevertheless evidence of complex burial customs.” ref
“The Pitted Ware people had an animistic cosmography similar to that of the people of the Comb Ceramic culture and other Mesolothic hunter-gatherers of the Baltic. Genetic studies of the Pitted Ware peoples have found them to have been strikingly genetically homogenous, suggesting that they originated from a small founder group. Already Malmström (2009) extracted mtDNA from seventeen Pitted Ware people of Gotland. Eight individuals belonged to U4 haplotypes, seven belonged to U5 haplotypes, one to K1a1, one to T2b, and one to HV0. The results debunked previous theories suggesting that the Pitted Ware were related to the Sami people. On the contrary, Pitted Ware people showed closer genetic kinship to modern Balts, Estonians, and modern Scandinavians.” ref
“In a genetic study published in BMC Evolutionary Biology in March 2010, it was discovered that the Pitted Ware possessed a very low level (5%) of an allele (−13910*T) strongly associated with the ability to consume unprocessed milk. This frequency is dramatically different from modern Swedes (74%). Whether the increase of this allele among the Swedes was a result of admixture or natural selection was uncertain. In a genetic study published in Science in April 2012, an individual from the Pitted Ware culture was examined. The individual was found to have “a genetic profile that is not fully represented by any sampled contemporary population”. In another genetic study published in Science in May 2014, the mtDNA of six individuals ascribed to the Pitted Ware culture was extracted. Four samples belonged to U4d, one belonged to U, and one belonged to V.” ref
“A genetic study published in August 2014 found that Pitted Ware peoples were closely genetically similar to people of the Catacomb culture, who like the Pitted Ware people carried high frequencies of the maternal haplogroups U5 and U4. These lineages are associated with Western Hunter-Gatherers and Eastern Hunter-Gatherers. In a genetic study published in Nature in September 2014, members of the Pitted Ware culture were determined to largely belong to the Scandinavian Hunter-Gatherer (SHG) cluster. In a genetic study published in Proceedings of the Royal Society B in January 2015, the mtDNA of thirteen PCW individuals from Öland and Gotland was extracted. The four individuals from Öland carried H1f, T2b, K1a1 and U4a1. Of the ten individuals from Gotland, four carried U4, two carried U5 haplotypes, two carried K1a1, and one carried HV0. The results indicated that the Pitted Ware culture was genetically distinct from the Funnelbeaker culture, and closely genetically related to earlier Mesolithic hunter-gatherers of Scandinavia and Western Europe. It was found that the Pitted Ware culture left a genetic imprint on Scandinavians, although this number is certainly not more than 60%.” ref
“A genetic study published in Nature Communications in January 2018 indicated genetic continuity between SHGs and the Pitted Ware culture, and found that the Pitted Ware people were genetically distinct from the Funnelbeaker culture. A 2018 study described the skin, hair and eye pigmentation of two Pitted Ware individuals from the Ajvide Settlement on the western coast of Gotland. The authors described the Pitted Ware specimens as sharing part of their phenotypic variation with Scandinavian Hunter Gatherers. Both specimens had a high probability of being dark-haired. One exhibited alleles associated with dark skin pigmentation on the SLC24A5 and SLC45A2 genes, while the other specimen carried a mix of alleles at these positions. The two specimens had a high probability of being blue eyed. A 2019 study published in Proceedings of the Royal Society B the remains of a Pitted Ware male were analyzed. He was found to the carrying the maternal haplgroup U5b1d2, and probably a subclade of the paternal haplogroup I2. He was estimated to be 25–35 years old and 165–175 cm tall. It was found that the Pitted Ware people only slightly contributed to the gene pool of the Battle Axe culture, who were almost wholly of Western Steppe Herder descent.” ref
“A genetic study published in Proceedings of the Royal Society B in June 2020 examined the remains of 19 Pitted Ware individuals buried on the island of Gotland. The study included a number of individuals who had been buried in a way typical of the Battle Axe culture. The 6 samples of Y-DNA extracted belonged to the paternal haplogroup I2a-L460 (2 samples), I2-M438 (2 samples), I2a1a-CTS595 and I2a1b1-L161. The 17 samples of mtDNA extracted belonged overwhelmingly to the maternal haplogroups U4 and U5. The study found no evidence of Battle Axe admixture among the Pitted Ware. They were genetically very different from earlier Funnelbeaker inhabitants of Gotland, although they carried a tiny amount of Early European Farmers (EEF) admixture. The evidence suggested that while the Pitted Ware culture was culturally influenced by the Battle Axe culture, it was not genetically influenced by it.” ref
Comb Ceramic culture
“The Comb Ceramic culture or Pit-Comb Ware culture, often abbreviated as CCC or PCW, was a northeast European culture characterised by its Pit–Comb Ware. It existed from around 4200 BCE to around 2000 BCE. The bearers of the Comb Ceramic culture are thought to have still mostly followed the Mesolithic hunter-gatherer (Eastern Hunter-Gatherer) lifestyle, with traces of early agriculture. The distribution of the artifacts found includes Finnmark (Norway) in the north, the Kalix River (Sweden) and the Gulf of Bothnia (Finland) in the west and the Vistula River (Poland) in the south. It would include the Narva culture of Estonia and the Sperrings culture in Finland, among others. They are thought to have been essentially hunter-gatherers, though e.g. the Narva culture in Estonia shows some evidence of agriculture. Some of this region was absorbed by the later Corded Ware horizon. The Pit–Comb Ware culture is one of the few exceptions to the rule that pottery and farming coexist in Europe. In the Near East farming appeared before pottery, then when farming spread into Europe from the Near East, pottery-making came with it. However, in Asia, where the oldest pottery has been found, pottery was made long before farming. It appears that the Comb Ceramic Culture reflects influences from Siberia and distant China.” ref
“By dating according to the elevation of land, the ceramics have traditionally (Äyräpää 1930) been divided into the following periods: early (Ka I, c. 4200 BC – 3300 BC), typical (Ka II, c. 3300 BC – 2700 BC) and late Comb Ceramic (Ka III, c. 2800 BC – 2000 BC). However, calibrated radiocarbon dates for the comb-ware fragments found (e.g., in the Karelian isthmus), give a total interval of 5600 BC – 2300 BC (Geochronometria Vol. 23, pp 93–99, 2004). The settlements were located at sea shores or beside lakes and the economy was based on hunting, fishing, and the gathering of plants. In Finland, it was a maritime culture that became more and more specialized in hunting seals. The dominant dwelling was probably a teepee of about 30 square meters where some 15 people could live. Also, rectangular houses made of timber became popular in Finland from 4000 BC cal. Graves were dug at the settlements and the dead were covered with red ochre. The typical Comb Ceramic age shows an extensive use of objects made of flint and amber as grave offerings.” ref
“The stone tools changed very little over time. They were made of local materials such as slate and quartz. Finds suggest a fairly extensive exchange network: red slate originating from northern Scandinavia, asbestos from Lake Saimaa, green slate from Lake Onega, amber from the southern shores of the Baltic Sea, and flint from the Valdai area in northwestern Russia. The culture was characterized by small figurines of burnt clay and animal heads made of stone. The animal heads usually depict moose and bears and were derived from the art of the Mesolithic. There were also many rock paintings. There are sources noting that the typical comb ceramic pottery had a sense of luxury and that its makers knew how to wear precious amber pendants. The great westward dispersal of the Uralic languages is suggested to have happened long after the demise of the Comb Ceramic culture, perhaps in the 1st millennium BC.” ref
“Saag et al. (2017) analyzed three CCC individuals buried at Kudruküla as belonging to Y-hg R1a5-YP1272 (R1a1b~ after ISOGG 2020), along with three mtDNA samples of mt-hg U5b1d1, U4a and U2e1. Mittnik (2018) analyzed two CCC individuals. The male carried R1 (2021: R1b-M343) and U4d2, while the female carried U5a1d2b. Generally, the CCC individuals were mostly of Eastern Hunter-Gatherer (EHG) descent, with even more EHG than people of the Narva culture. Lamnidis et al. (2018) found 15% Western Hunter-Gatherer (WHG) ancestry, 65% Eastern Hunter-Gatherer (EHG) – higher than among earlier cultures of the eastern Baltic, and 20% Western Steppe Herder (WSH).” ref
Postglacial genomes from foragers across Northern Eurasia reveal prehistoric
mobility associated with the spread of the Uralic and Yeniseian languages
Abstract
“The North Eurasian forest and forest-steppe zones have sustained millennia of sociocultural connections among northern peoples. We present genome-wide ancient DNA data for 181 individuals from this region spanning the Mesolithic, Neolithic, and Bronze Age. We find that Early to Mid-Holocene hunter-gatherer populations from across the southern forest and forest-steppes of Northern Eurasia can be characterized by a continuous gradient of ancestry that remained stable for millennia, ranging from fully West Eurasian in the Baltic region to fully East Asian in the Transbaikal region. In contrast, cotemporaneous groups in far Northeast Siberia were genetically distinct, retaining high levels of continuity from a population that was the primary source of ancestry for Native Americans. By the mid-Holocene, admixture between this early Northeastern Siberian population and groups from Inland East Asia and the Amur River Basin produced two distinctive populations in eastern Siberia that played an important role in the genetic formation of later people. Ancestry from the first population, Cis-Baikal Late Neolithic-Bronze Age (Cisbaikal_LNBA), is found substantially only among Yeniseian-speaking groups and those known to have admixed with them. Ancestry from the second, Yakutian Late Neolithic-Bronze Age (Yakutia_LNBA), is strongly associated with present-day Uralic speakers. We show how Yakutia_LNBA ancestry spread from an east Siberian origin ~4.5kya, along with subclades of Y-chromosome haplogroup N occurring at high frequencies among present-day Uralic speakers, into Western and Central Siberia in communities associated with Seima-Turbino metallurgy: a suite of advanced bronze casting techniques that spread explosively across an enormous region of Northern Eurasia ~4.0kya. However, the ancestry of the 16 Seima-Turbino-period individuals–the first reported from sites with this metallurgy–was otherwise extraordinarily diverse, with partial descent from Indo-Iranian-speaking pastoralists and multiple hunter-gatherer populations from widely separated regions of Eurasia. Our results provide support for theories suggesting that early Uralic speakers at the beginning of their westward dispersal where involved in the expansion of Seima-Turbino metallurgical traditions, and suggests that both cultural transmission and migration were important in the spread of Seima-Turbino material culture.” ref
Sidelkino Ancestry?
“The Sidelkino people in the east had genes associated with dark eyes and light skin.” ref
“The marked shift from Western Hunter-Gatherer-like towards Eastern Hunter-Gatherer-related ancestry from Baltic Mesolithic (ca. 30%) to Combed Ware cultures (ca. 65%-100%) supports this continuous westward expansion, that is possibly best represented in the currently available sampling by the ‘south-eastern’ shift (CHG:ANE-related) of the hunter-gatherer from Lebyazhinka IV (5600 BCE) relative to the older one from Sidelkino (9300 BCE), both from the Samara region in the Middle Volga.” ref
Dzudzuana, Sidelkino, and the Caucasus contribution to the Pontic-Caspian steppe
“It has been known for a long time that the Caucasus must have hosted many (at least partially) isolated populations, probably helped by geographical boundaries, setting it apart from open Eurasian areas. David Reich writes in his book the following about India: The genetic data told a clear story. Around a third of Indian groups experienced population bottlenecks as strong or stronger than the ones that occurred among Finns or Ashkenazi Jews. We later confirmed this finding in an even larger dataset that we collected working with Thangaraj: genetic data from more than 250 jati groups spread throughout India (…)” ref
Genetic ancestry changes in Stone to Bronze Age transition in the East European plain
“The transition from Stone to Bronze Age in Central and Western Europe was a period of major population movements originating from the Ponto-Caspian Steppe. Here, we report new genome-wide sequence data from 30 individuals north of this area, from the understudied western part of present-day Russia, including 3 Stone Age hunter-gatherers (10,800 to 4250 cal BCE) and 26 Bronze Age farmers from the Corded Ware complex Fatyanovo Culture (2900 to 2050 cal BCE). We show that Eastern hunter-gatherer ancestry was present in northwestern Russia already from around 10,000 BCE. Furthermore, we see a change in ancestry with the arrival of farming—Fatyanovo Culture individuals were genetically similar to other Corded Ware cultures, carrying a mixture of Steppe and European early farmer ancestry. Thus, they likely originate from a fast migration toward the northeast from somewhere near modern-day Ukraine—the closest area where these ancestries coexisted from around 3000 BCE.” ref
“The researchers studied the genetic affinities of the higher-coverage (5×) HG PES001 further using outgroup f3 statistics by comparing the affinities of three higher-coverage Russian Mesolithic HGs—PES001 (Peschanitsa; 10,785 to 10,626 cal BCE), I0061 (Yuzhnyy Oleni Ostrov; 6773 to 5886 BCE), and Sidelkino (Sidelkino; 9386 to 9231 cal BCE)—to Mesolithic and Paleolithic HGs from different areas of Europe and Siberia (Fig. 2C and table S9). All three are most similar to individuals from the European part of Russia or from Siberia who lived within 10,000 years of each other—to each other, the West Siberia Neolithic population, and the Afontova Gora 3 individual. These are followed by individuals from Central Europe from the same time window. Geographically close Paleolithic Sunghir and Kostenki individuals from >30,000 cal BCE share less than temporally close HGs from Central Europe with the Russian Mesolithic individuals. We also tried modeling PES001 as a mixture of WHG and either CHG, Mal’ta, or Afontova Gora 3 using qpAdm, but all three models were rejected (P value of 1.96 × 10−85 to 7.93 × 10−13).” ref
EF ancestry in Fatyanovo Culture individuals
“Then, the researchers turned to the Bronze Age Fatyanovo Culture individuals and determined that they carry maternal (subclades of mtDNA hg U5, U4, U2e, H, T, W, J, K, I, and N1a) and paternal (chrY hg R1a-M417) lineages that have also been found in CWC individuals elsewhere in Europe. In all individuals for which the chrY hg could be determined with sufficient depth (n = 6), it is R1a2-Z93, a lineage now spread in Central and South Asia, rather than the R1a1-Z283 lineage that is common in Europe. R1a2-Z93 is also not rejected for the individuals that were determined with less depth due to missing data on more apical markers.” ref
“On the PCA, the Fatyanovo individuals (and the Estonian CWC individual) group together with many European Late Neolithic/Bronze Age (LNBA) and Steppe Middle/Late Bronze Age individuals on top of modern Northern and Eastern Europeans (Fig. 2A). This ancient cluster is shifted toward Anatolian and European EF compared to Steppe Early/Middle Bronze Age populations, including the Yamnaya. The same could be seen in ADMIXTURE analysis where the Fatyanovo individuals are most similar to LNBA Steppe ancestry populations from Central Europe, Scandinavia, and the Eastern Baltic (Fig. 2B). These populations are composed of the “WHG” (blue) and “ancient Caucasus/Iran” (olive) component and small amounts of the “Russian Far East” (orange) component, similarly to Yamnaya populations. However, the European LNBA populations (including Fatyanovo) also display a component most frequent in Anatolian and European EF populations (light green), which is not present in the Yamnaya from Russia.” ref
“The researchers studied the affinities of the Fatyanovo individuals by comparing FST, outgroup f3, and D statistics’ results of different populations and found that Fatyanovo shares more with European EF populations and modern Near Easterners than Yamnaya Samara does (Fig. 3C). This signal can also be seen when using either autosomal or X chromosome (chrX) positions from the 1240K dataset instead of the autosomal positions of the HO dataset with less single-nucleotide polymorphisms (SNPs). We also compared the affinities of Yamnaya Samara and Fatyanovo directly with D statistics and saw that Fatyanovo is significantly more similar (Z > 3) to most EF populations than to Yamnaya Samara, and the latter, in turn, is significantly more similar to most Steppe populations than to Fatyanovo. We studied the apparent EF input in Fatyanovo further using admixture f3 statistics and got significant results (Z < −3) for admixture between different Yamnaya groups and a wide variety of EF populations. Furthermore, when comparing outgroup f3 and D statistics’ results for Fatyanovo and Central CWC, there are no clear differences in their affinities to different ancient or modern population groups (Fig. 3D).” ref
“Because the previous analyses suggested that the genetic makeup of the Fatyanovo Culture individuals was a result of admixture between migrating Yamnaya individuals and contemporary European populations, we used two complementary methods (qpAdm and ChromoPainter/NNLS) to determine suitable proxies for the admixing populations and the mixing proportions (Fig. 4). We tested qpAdm models including Yamnaya from Samara or Kalmykia and a variety of EF populations one at a time and found that the two EF populations, with the highest P values with both Yamnaya populations, are Globular Amphora and Trypillia (P values of 0.02/0.16 and 0.06/0.26, respectively) (table S15). The admixture proportions are 65.5%/66.9% Yamnaya Samara/Kalmykia + 34.5%/33.1% Globular Amphora and 65.5%/69.6% Yamnaya Samara/Kalmykia + 34.5%/30.4% Trypillia, respectively (Fig. 4). The proportions are similar (69 to 75% Yamnaya + 25 to 31% EF) for Central and Baltic CWC (Fig. 4).” ref
“We also tested these four models with the preceding Volosovo Culture HG BER001 added to “right” populations to see whether there is shared drift between Volosovo and Fatyanovo that would cause the models with admixing populations without this drift to be rejected. All four models are still not rejected (P values of 0.97/0.57 and 0.98/0.59, respectively), suggesting that no Volosovo contribution is needed to model Fatyanovo. Ancestry proportions were also calculated using the ChromoPainter/NNLS pipeline with the results 37%/38% Yamnaya Samara + 63%/62% Globular Amphora/Trypillia for Fatyanovo and 51 to 60% Yamnaya + 40 to 49% EF for Central/Baltic CWC.” ref
“Although the estimated proportion of EF ancestry is higher for Fatyanovo compared to the other groups in both cases, the difference is significant (P value of 0.005 to 0.03) only with Trypillia in the model. Note that qpAdm calculates admixture between populations, while ChromoPainter/NNLS uses single individuals as sources, which might influence the results. Although two-way admixture between Yamnaya and an EF population is enough to explain the genetic variation in Fatyanovo, qpAdm models with an HG population added are also not rejected with EHG, WeRuHG, and Volosovo (P values of 0.14 to 0.98). Fatyanovo can be modeled as 60 to 63% Yamnaya Samara + 33 to 34% Globular Amphora + 3 to 6% HG. The results are similar for Central and Baltic CWC with 2 to 11% of HG ancestry (except with Volosovo as a source for Central CWC, which gets a negative mixture coefficient).” ref
“From around 8,00 years ago, we begin to observe admixture events with Sidelkino ancestry in central Europe. This is first detected in an individual from Gross Fredenwalde in northeastern Germany and reaches around 10% in most European hunter-gatherer individuals thereafter. Soon after 8 ka, Sidelkino ancestry was absent in eastern Spain but it had already reached northern Iberia alongside an increase in Oberkassel ancestry (Fig. 5). Conversely, additional Oberkassel ancestry is identified in eastern Europe by at least 7,500 years ago in newly generated genomes from Minino I and Yazykovo from the upper Volga region, whereas a 1,000-years-older individual from Minino I did not have this genetic component. Considering a freshwater reservoir signal in the upper Volga region making radiocarbon dates on human remains appear up to about 500 years older than their true age, there could be an interval of more than 1,000 years between the first evidence of admixture in central European hunter-gatherers with Sidelkino ancestry and eastern European hunter-gatherers with Oberkassel ancestry. However, additional genomes intermediate in time and space are needed to assess whether those two admixture events were independent or part of a common demographic process.” ref
“After 7,500 years ago, as ANF ancestry had reached regions north of the Alps, individuals carrying a hunter-gatherer genetic profile were primarily restricted to the northern fringes of Europe (Fig. 5). In this period, the Oberkassel-ancestry admixture spread further east, reaching Samara by around 6,500 years ago, and an increase in Sidelkino ancestry was detected in hunter-gatherers from the Baltic region, which was previously associated with the transition from the Narva culture to the Comb Ceramic culture (Fig. 8). In central Europe, admixture with ANF ancestry became highly common but not ubiquitous, indicating the co-existence of hunter-gatherer and farmer societies without admixing for several hundred years. The youngest individual carrying large portions of hunter-gatherer ancestry in the analyzed dataset is from Ostorf in northern Germany, dated to around 5,200 years ago (>90% Oberkassel cluster plus Sidelkino-cluster components). Individuals at this site might mark one of the last occurrences of such high levels of hunter-gatherer-related ancestries, just centuries before the emerging European Bronze Age.” ref
“On the basis of PCA and outgroup f3-statistics, the Neolithic Tutkaul 1 individual from Tajikistan is closely related to Upper Palaeolithic individuals from south-central Siberia (Afontova Gora 3 (AG3) and Mal’ta 1), and roughly contemporaneous West Siberian hunter-gatherers (Tyumen and Sosnoviy), both carrying high proportions of ANE ancestry (Fig. 1c and Fig. 6). We tested the affinity of Tutkaul 1 to worldwide ancient and modern populations relative to AG3. Contrary to West Siberian hunter-gatherers, Tutkaul 1 does not carry an extra eastern Eurasian ancestry, but shows affinity to Iranian Neolithic farmers and some younger populations from Iran and the Turan region. Conversely, individuals in the Sidelkino cluster are genetically closer to AG3 than Tutkaul 1. This suggests that the newly reported Neolithic individual from central Asia carries an ancestry that might be a good proxy for the ANE-related contribution to Iran and the Turan region from around 5,500 years ago but not to roughly contemporaneous hunter-gatherers from eastern Europe.” ref
“In sum, we describe the formation and interaction between the Oberkassel and Sidelkino clusters, the two main hunter-gatherer ancestries present in Europe from 14 ka onwards. The genomic similarity of the Oberkassel cluster to Arene Candide 16 in northwestern Italy might imply that Epigravettian-associated ancestry spread from the south to central Europe passing through the western side of the Alpine region. The Sidelkino ancestry also emerged around 14,000 years ago with its first direct evidence in eastern Europe dated to 13,000 years ago. The increasing level of admixture between distinct hunter-gatherer populations from around 8,000 years ago onwards indicates an intensified mobility of those forager groups. This might have been in part triggered by the concomitant expansion of Neolithic farmers across Europe and/or by environmental factors, such as the climatic event around 8,200 years ago, the largest abrupt cooling in the northern hemisphere during the Holocene epoch.” ref
“Leveraging the substantially increased sample size, we investigated genetically distinct hunter-gatherer groups for allele frequencies at selected loci that are known to be associated with specific phenotypic traits in present-day Europeans (Fig. 5b, Figs. 27 and 28). Consistent with previous findings, none of the analyzed groups show the derived allele at SNP rs4988235 on the LCT gene, which is responsible for lactase persistence. As previously hinted, we find a large frequency variation in alleles related to skin and eye pigmentation among post-LGM hunter-gatherer groups. For the SNP associated with light eye color (HERC2/OCA2 (rs12913832)), individuals from the Villabruna cluster, Oberkassel cluster, Baltic HG, and SHG groups show high frequencies of the derived allele (>90%), which is responsible for the green or blue eye phenotype, whereas Sidelkino cluster, Ukraine HG and Iron Gates HG groups show the low occurrence of this allele (10–25%).” ref
“Instead, for the two SNPs associated with skin color (SLC24A5 (rs1426654) and SLC45A2 (rs16891982)), Sidelkino cluster and Ukraine HG groups show a higher frequency (>90% for SLC24A5 and 29–61% for SLC45A2) of the derived alleles related to light skin color, compared with Oberkassel and Villabruna clusters, where those alleles are almost completely absent (<1%). On the basis of the genetic variation of present-day Europeans, this could imply phenotypic differences between post-14 ka hunter-gatherer populations across Europe, with individuals in the Oberkassel cluster possibly exhibiting darker skin and lighter eyes, and individuals in the Sidelkino cluster possibly lighter skin and darker eye color.” ref
The data generated in this study enabled us to investigate genomic transformations of and interactions between Eurasian hunter-gatherers at high resolution (Fig. 9). We provide five novel insights into the genomic history of hunter-gatherer populations over a time span of 30,000 years from the Upper Palaeolithic to the Neolithic. First, we show that individuals associated with the Gravettian culture across Europe were not a biologically homogeneous population. Culturally, however, we see both widespread general tendencies, such as weaponry and some portable art, and other aspects that have a more regional character, such as mortuary practices, various originalities in lithic and hard organic materials tool kits and adornments.” ref
“The ancestry found in individuals associated with the preceding Aurignacian culture from central Europe (GoyetQ116-1 ancestry) gave rise to Gravettian-associated individuals from western and southwestern Europe. This derived ancestry—the Fournol cluster—survived during the LGM in Solutrean-associated individuals, possibly within the Franco-Cantabrian climatic refugium, leading to later populations associated with the Magdalenian culture (GoyetQ2 cluster and El Mirón). Conversely, the ancestry found in pre-30 ka eastern European individuals (Kostenki cluster and Sunghir group) contributed to Gravettian-associated individuals from central and southern Europe (Věstonice cluster), the latter without descendants retrieved in post-LGM populations from those regions.” ref
“Second, the ancestry of individuals associated with the Epigravettian culture (Villabruna cluster), which was found to genetically connect European and Near Eastern hunter-gatherers, reached southern Europe well before the transition between the Early and Late Epigravettian and possibly as early as the Gravettian–Epigravettian transition. A phylogeographic reconstruction of different lineages carrying this ancestry further suggests its entry point into northeastern Italy from the Balkans followed by a north-to-south expansion into the Italian peninsula alongside a population decline through sequential bottlenecks.” ref
“Third, Magdalenian-associated individuals not only from Iberia but also from the rest of Europe carry Epigravettian-associated ancestry (Villabruna cluster). Genetic analyses of western European individuals associated with the preceding Badegoulian culture might provide clues on the processes that led to the formation of the GoyetQ2 cluster. As inferred from the archaeological record, the spread of the Magdalenian across Europe is linked to southwestern to northern and northeastern post-LGM population expansions and not to movements from southeastern refugia.” ref
“Fourth, we extend the finding of a large-scale genetic turnover as early as 14 ka in central and western European hunter-gatherers associated with multiple techno-complexes—Federmesser, Azilian, and other Final Palaeolithic groups—despite considerable technological continuity with the preceding late Magdalenian. This broadly distributed ancestry (the Oberkassel cluster (also known as WHG)) is most closely related to an Epigravettian-associated individual from northwestern Italy, suggesting that its expansion into continental Europe might have started from the west—and not the east—side of the Alps. Moreover, the almost complete genetic replacement of the Magdalenian-associated gene pool raises the hypothesis that parts of Europe were differentially populated during the abrupt climatic variation starting around 14,700 years ago with the Bølling–Allerød warming period, creating areas where southern European populations could expand. This might also explain the genetic uniformity of the Oberkassel cluster across large parts of western Eurasia but genomic data from between 15,000 and 14,000 years ago is needed to understand the exact dynamics of this turnover.” ref
“Fifth, the Oberkassel ancestry in western and central Europe and the Sidelkino ancestry in eastern Europe remained largely isolated for almost 6,000 years until genetic interactions were first observed—around 8,000 years ago in northeastern Germany, possibly associated with cultural exchanges along the Baltics and around 7,5000 years ago in the upper Volga region, possibly linked to the spread of pottery in the region. This study reveals that western and southwestern Europe served as climatic refugia for the persistence of human groups during the coldest phase of the last Ice Age whereas populations in the Italian peninsula and the eastern European plain were genetically overturned, challenging the role of these regions as glacial refugia for humans.” ref
“The incoming Villabruna ancestry later became the most widespread hunter-gatherer ancestry across Europe. Further palaeogenomic studies on Upper Palaeolithic individuals from the Balkans will be essential for understanding whether southeastern Europe represents the source of the Villabruna ancestry and a climatic refugium for human populations during the LGM. Note added in proof: A companion paper describes genome-wide data of a 23,000-year-old Solutrean-associated individual from southern Iberia that extend the evidence of genetic continuity across the LGM in southwestern Europe.” ref
“The Yamnaya archaeological complex appeared around 3300BCE across the steppes north of the Black and Caspian Seas, and by 3000BCE reached its maximal extent from Hungary in the west to Kazakhstan in the east. To localize the ancestral and geographical origins of the Yamnaya among the diverse Eneolithic people that preceded them, we studied ancient DNA data from 428 individuals of which 299 are reported for the first time, demonstrating three previously unknown Eneolithic genetic clines. First, a “Caucasus-Lower Volga” (CLV) Cline suffused with Caucasus hunter-gatherer (CHG) ancestry extended between a Caucasus Neolithic southern end in Neolithic Armenia, and a steppe northern end in Berezhnovka in the Lower Volga. Bidirectional gene flow across the CLV cline created admixed intermediate populations in both the north Caucasus, such as the Maikop people, and on the steppe, such as those at the site of Remontnoye north of the Manych depression. CLV people also helped form two major riverine clines by admixing with distinct groups of European hunter-gatherers.” ref
“A “Volga Cline” was formed as Lower Volga people mixed with upriver populations that had more Eastern hunter-gatherer (EHG) ancestry, creating genetically hyper-variable populations as at Khvalynsk in the Middle Volga. A “Dnipro Cline” was formed as CLV people bearing both Caucasus Neolithic and Lower Volga ancestry moved west and acquired Ukraine Neolithic hunter-gatherer (UNHG) ancestry to establish the population of the Serednii Stih culture from which the direct ancestors of the Yamnaya themselves were formed around 4000BCE. This population grew rapidly after 3750–3350BCE, precipitating the expansion of people of the Yamnaya culture who totally displaced previous groups on the Volga and further east, while admixing with more sedentary groups in the west. CLV cline people with Lower Volga ancestry contributed four fifths of the ancestry of the Yamnaya, but also, entering Anatolia from the east, contributed at least a tenth of the ancestry of Bronze Age Central Anatolians, where the Hittite language, related to the Indo-European languages spread by the Yamnaya, was spoken. We thus propose that the final unity of the speakers of the “Proto-Indo-Anatolian” ancestral language of both Anatolian and Indo-European languages can be traced to CLV cline people sometime between 4400–4000 BCE.” ref
“The origin of Indo-Anatolian and Indo-European languages. Genetic reconstruction of the ancestry of Pontic-Caspian steppe and West Asian populations points to the North Caucasus-Lower Volga area as the homeland of Indo-Anatolian languages and to the Serednii Stih archaeological culture of the Dnipro-Don area as the homeland of Indo-European languages. The Caucasus-Lower Volga people had diverse distal roots, estimated using the qpAdm software on the left barplot, as Caucasus hunter-gatherer (purple), Central Asian (red), Eastern hunter-gatherer (pink), and West Asian Neolithic (green). Caucasus-Lower Volga expansions, estimated using qpAdm on the right barplot as disseminated Caucasus Neolithic (blue)-Lower Volga Eneolithic (orange) proximal ancestries, mixing with the inhabitants of the North Pontic region (yellow), Volga region (yellow), and West Asia (green).” ref
“Between 3300–1500 BCE, people of the Yamnaya archaeological complex and their descendants, in subsequent waves of migration, spread over large parts of Eurasia, contributing to the ancestry of people of Europe, Central and South Asia, Siberia, and the Caucasus. The spread of Indo-European language and culture transformed all these regions. Despite the centrality of the Yamnaya expansion to the human story of Bronze Age Eurasia, their ancestral origins are poorly understood. A first challenge has been the sparse sampling of the Yamnaya themselves across their enormous geographic distribution. The remarkable long-range mobility of the Yamnaya, quickly spreading over a vast region, adds further difficulties to tracing, from radiocarbon dating, the origins of their material culture and associated genetic profile. Nor can these origins be traced to the numerous earlier Eneolithic cultures that preceded the Yamnaya, and among whom their ancestors must be sought, as these have been sampled even more poorly and systematically.” ref
“The first formal study of the origins of the Yamnaya identified two disparate sources of ancestry: a northern, “Eastern Hunter-Gatherer” (EHG) source from far eastern Europe, and a southern, West Asian source related to present-day Armenians.The latter source was revealed, by ancient DNA, to be related to some of the region’s earliest inhabitants: Paleolithic-Mesolithic “Caucasus Hunter-Gatherers” (CHG) of Georgia, and Neolithic people of the Zagros and South Caucasus. Additional discoveries further complicated the stories of both the northern and southern ancestors of the Yamnaya. First, it was noted that both CHG and EHG were part of an interaction sphere across the boundary between West Asia and eastern Europe, suggesting the existence of intermediate populations and raising the question of when and where these came together to form the Eneolithic antecessors of the Yamnaya. Second, it was recognized that the steppe itself was an admixture zone of EHG with “Western Hunter-Gatherers” (WHG). Mesolithic hunter-gatherers from Ukraine were succeeded by more WHG-admixed Neolithic hunter-gatherers in the Dnipro valley, representing a local reshuffling within the European portion of a ~7,000 km-long trans-Eurasian cline of boreal hunter-gatherers.” ref
“What was the relative contribution of the EHG (who were present in the Volga River at Lebyazhinka ca. 5660–5535 BCE) and these more western Ukraine Neolithic hunter-gatherers (UNHG) of the Dnipro to later populations? Third, it was discovered that the Yamnaya had not only CHG-related, but also Anatolian Neolithic ancestry, absent in the early known steppe inhabitants, and derived from European farmer neighbors west of the steppe. This ancestry was later shown to be of rather Anatolian-Levantine-Mesopotamian origin, and to be mediated not from Europe but from the Caucasus neighbors south of the steppe. Such ancestry must have been added following the expansion of Neolithic farmers into the Caucasus, introduced thence into the steppe as a later exogenous element, distinct from the earlier CHG-related one. Finally, it was recognized that European steppe populations were formed not only by northern-southern admixture, but included, in at least some Eneolithic and Bronze Age people of the North Caucasus, contributions related to Siberians from further east. What was the extent of the spread of this eastern ancestry, and did the Yamnaya themselves possess it?” ref
“Here we present a unified population genetic analysis of 372 newly reported individuals dating from 6400–2000 BCE, as well as increased quality data for 61 individuals. The present study serves as the formal technical report for 299 of the newly reported individuals and 55 of the individuals with increased quality data; more than 80% of the individuals are from Russia, but the dataset is also significant in including dozens of individuals from westward expansion of Steppe cultures along the Danube.” ref
“A parallel study presents a combined archaeological and genetic analysis of population transformations in the North Pontic Region (Ukraine and Moldova) and serves as the formal report for the data from the other 73 of the newly analyzed individuals and the other 5 individuals with increased quality data, with both studies co-analyzing the full dataset. We grouped individuals into analysis labels based on geographical and temporal information, archaeological context, and genetic clustering. The potential of the combined dataset for shedding light on this period can be appreciated from the fact that it adds 79 analyzed Eneolithic people from the steppe and its environs (from Russia or Ukraine, west of 60E longitude and south of 60N latitude, between 5000–3500BCE) to 82 published and a total of 286 Yamnaya/Afanasievo individuals compared to 75 in the literature.” ref
J-M172 (Zagros mountains origin)
“J-M304 (Transcaucasian origin) is defined by the M304 genetic marker, or the equivalent 12f2.1 marker. The main current subgroups J-M267 (Armenian highlands origin) and J-M172 (Zagros mountains origin), which now comprise between them almost all of the haplogroup’s descendant lineages, are both believed to have arisen very early, at least 10,000 years ago. Nonetheless, Y-chromosomes F-M89* and IJ-M429* were reported to have been observed in the Iranian plateau (Grugni et al. 2012). On the other hand, it would seem to be that different episodes of populace movement had impacted southeast Europe, as well as the role of the Balkans as a long-standing corridor to Europe from the Near East is shown by the phylogenetic unification of Hgs I and J by the basal M429 mutation. This proof of common ancestry suggests that ancestral Hgs IJ-M429* probably would have entered Europe through the Balkan track sometime before the LGM. They then subsequently split into Hg J and Hg I in Middle East and Europe in a typical disjunctive phylogeographic pattern.” ref
“Haplogroup J-M172 (J2-M172) is common in modern populations in Western Asia, Central Asia, South Asia, Southern Europe, Northwestern Iran and North Africa. It is thought that J-M172 may have originated in the Caucasus, Anatolia and/or Western Iran. The date of origin for haplogroup J-M172 was estimated by Batini et al in 2015 as between 19,000 and 24,000 years ago. Samino et al in 2004 dated the origin of the parent haplogroup, J-P209, to between 18,900 and 44,500 years ago. Ancient J-M410, specifically subclade J-Y12379*, has been found, in a mesolithic context, in a tooth from the Kotias Klde Cave in western Georgia dating 9.529-9.895 years ago. This sample has been assigned to the Caucasus hunter-gatherers (CHG) autosomal component. J-M410, more specifically its subclade J-PF5008, has also been found in a mesolithic sample from the Hotu and Kamarband Caves located in Mazandaran Province of Iran, dating back to 9,100-8,600 BCE (approximately 11,000 yrats ago). Both samples belong to the Trialetian Culture. It is likely that J2 men had settled over most of Anatolia, the South Caucasus and the Zagros mountains by the end of the Last Glaciation 12,000 years ago.” ref
“Zalloua & Wells (2004) and Al-Zahery et al. (2003) claimed to have uncovered the earliest known migration of J2, expanded possibly from Anatolia and the Caucasus. Nebel et al. (2001) found that, “According to Underhill et al. (2000), Eu 9 (H58) evolved from Eu 10 (H71) through a T→G transversion at M172 (emphasis added),” and that in today’s populations, Eu 9 (the post-mutation form of M172) is strongest in the Caucasus, Asia Minor and the Levant, whilst Eu 10 becomes stronger and replaces the frequency of Eu 9 as one moves south into the Arabian Peninsula, so that people from the Caucasus met with Arabs near and between Mesopotamia (Sumer/Assyria) and the Negev Desert, as “Arabisation” spread from Arabia to the Fertile Crescent and Turkey.” ref
“Di Giacomo et al. (2004) postulated that J-M172 haplogroup spread into Southern Europe from either the Levant or Anatolia, likely parallel to the development of agriculture. As to the timing of its spread into Europe, Di Giacomo et al. points to events which post-date the Neolithic, in particular the demographic floruit associated with the rise of the Ancient Greek world. Semino et al. (2004) derived older age estimates for overall J2 (having used the Zhivotovsky method c.f. Di Giacomo), postulating its initial spread with Neolithic farmers from the Near East. However, its subclade distribution, showing localized peaks in the Southern Balkans, southern Italy, north/central Italy and the Caucasus, does not conform to a single ‘wave-of-advance’ scenario, betraying a number of still poorly understood post-Neolithic processes which created its current pattern. Like Di Giacomo et al., the Bronze Age southern Balkans was suggested by Semino et al. to have been an important vector of spread.” ref
“Haplogroup J-M172 is found mainly in the Fertile Crescent, the Caucasus, Anatolia, Italy, the Mediterranean littoral, and the Iranian plateau. Y-DNA: J2 (J-M172): Syrid/Nahrainid Arabid(s). J-M172 is found at very high frequencies in certain peoples of the Caucasus: among the Ingush 87.4%, Chechens 55.2%, Georgians 21%-72%, Azeris 24%-48%, Abkhaz 25%, Balkars 24%, Ossetians 24%, Armenians 21%-24%, Adyghe 21.8%, and other groups. J-M172 is typical of populations of the Near East, Southern Europe, Southwest Asia and the Caucasus, with a moderate distribution through much of Central Asia, South Asia, and North Africa. The highest reported frequency of J-M172 ever was 87.4%, among Ingush in Malgobek.” ref
“More specifically it is found in Iraq, Kuwait, Syria, Lebanon, Turkey, Georgia, Azerbaijan, North Caucasus, Armenia, Iran, Israel, Palestine, Cyprus, Greece, Albania, Italy, Spain, and more frequently in Iraqis 24%, Chechens 51.0%-58.0%, Georgians 21%-72%, Lebanese 30%, Ossetians 24%, Balkars 24%, Syrians 23%, Turks 13%-40%, Cypriots 12.9%-37%, Armenians 21%-24%, Circassians 21.8%, Iranians 10%-25%, Albanians 16%, Italians 9%-36%, Sephardi Jews 15%-29%, Maltese 21%, Palestinians 17%, Saudis 14%, Jordanians 14%, Omanis 10%-15%, and North Indian Shia Muslim 18%.” ref
“It has been proposed that haplogroup subclade J-M410 was linked to populations on ancient Crete by examining the relationship between Anatolian, Cretan, and Greek populations from around early Neolithic sites in Crete. Haplogroup J-M172 was associated with Neolithic Greece (ca. 8500 – 4300 BCE) and was reported to be found in modern Crete (3.1%) and mainland Greece (Macedonia 7.0%, Thessaly 8.8%, Argolis 1.8%). Sephardi Jews have about 15%-29%, of haplogroup J-M172, and Ashkenazi Jews have 15%-23%. It was reported in an early study which tested only four STR markers that a small sample of Italian Cohens belonged to Network 1.2, an early designation for the overall clade now known as J-L26, defined by the deletion at DYS413. However, a large number of all Jewish Cohens in the world belong to haplogroup J-M267 (see Cohen modal haplotype).” ref
“Haplogroup J-M172 has been shown to have a more northern distribution in the Middle East, although it exists in significant amounts in the southern middle-east regions, a lesser amount of it was found when compared to its brother haplogroup, J-M267, which has a high frequency southerly distribution. It was believed that the source population of J-M172 originated from the Levant/Syria (Syrid-J-M172), and that its occurrence among modern populations of Europe, Central Asia, and South Asia was a sign of the neolithic agriculturalists. However, as stated it is now believed more likely to have been spread in waves, as a result of post-Neolithic processes.” ref
“Haplogroup J-M172 is found in the highest concentrations in the Caucasus and the Fertile Crescent/Iraq and is found throughout the Mediterranean (including the Italian, Balkan, Anatolian and Iberian peninsulas and North Africa). The highest ever reported concentration of J-M172 was 72% in Northeastern Georgia. Other high reports include Ingush 32% (Nasidze 2004), Cypriots 30-37% (Capelli 2005), Lebanese 30% (Wells et al. 2001), Assyrian, Mandean and Arab Iraqis 29.7%, Syrians and Syriacs 22.5%, Kurds 24%-28%, Pashtuns 20-30%, Iranians 23% (Aburto 2006), Ashkenazi Jews 24%, Palestinian Arabs 16.8%-25%, Sephardic Jews 29% and North Indian Shia Muslim 18%, Chechens 26%, Balkars 24%, Yaghnobis 32%, Armenians 21-24%, and Azerbaijanis 24%-48%.” ref
“In South Asia, J2-M172 was found to be significantly higher among Dravidian castes at 19% than among Indo-European castes at 11%. J2-M172 and J-M410 is found 21% among Dravidian middle castes, followed by upper castes, 18.6%, and lower castes 14%. Subclades of M172 such as M67 and M92 were not found in either Indian or Pakistani samples which also might hint at a partial common origin.” ref
“According to a genetic study in China by Shou et al., J2-M172 is found with high frequency among Uygurs (17/50 = 34%) and Uzbeks (7/23 = 30.4%), moderate frequency among Pamiris (5/31 = 16.1%), and also found J-M172 in Han Chinese (10%) and low frequency among Yugurs (2/32 = 6.3%) and Monguors (1/50 = 2.0%). The authors also found J-M304(xJ2-M172) with low frequency among the Russians (1/19 = 5.3%), Uzbeks (1/23 = 4.3%), Sibe people (1/32 = 3.1%), Dongxiangs (1/35 = 2.9%), and Kazakhs (1/41 = 2.4%) in Northwest China. Only far northwestern ethnic minorities had haplogroup J in Xinjiang, China. Uzbeks in the sample had 30.4% J2-M172 and Tajiks of Xinjiang and Uyghurs also had it.” ref
“The oldest evidence of agriculture comes from the Fertile Crescent (~11,000 years ago), the centre for demic diffusion. However, evidences of first agriculture from South Asia indicate a timeline (~9,000-10,000 years ago) closer to the emergence of agriculture in the Fertile Crescent. Previous studies have established that J2a-M410 and J2b-M102 represent the Y-chromosomal component associated with demic diffusion of Neolithic farmers in North Africa and Eurasia from Mesopotamia (Iraq and Syria). The J2-M172 has been associated with different cultures and populations in history, such as Mediterranean/Aegean, Greco-Anatolian, Mesopotamian, and Caucasian. Presence of J2a-M410 and J2b-M102 in India has been considered a result of gene influx from Western Asia. Worldwide spatial distribution of haplogroup (HG) J2a-M410 coincides with presence of archaeological records of painted pottery and ceramic figurine culture. Similar material culture dating ~9,000 years ago has been recovered from the Neolithic sites of Mehrgarh located West of Indus Valley (now in Pakistan).” ref
“The population of the sub-continent shows an outstanding biological and cultural variation, which has been shaped by geographical, ecological, social, and linguistic factors. Many recent studies, particularly from the field of evolutionary genetics, have shown the extent to which each of these factors has contributed to the generation and maintenance of this diversity. It is pertinent that paternal gene pool of India comprises mainly HGs of autochthonous origin of Late Pleistocene ancestry and received very little gene flow from outside. Many Y chromosomal studies of India addressing major demographic questions have included and broadly discussed HG J2-M172.” ref
“However, the arrival and distribution of HG J2-M172 subclades in India have not been studied comprehensively yet. Hence, the aim of this study is to address the following: (1) whether or not, the demic diffusion model stands true for the current distribution of J2-M172 in India in the backdrop of independent origin of agriculture in South Asia; (2) which contemporary populations have the closest affinities with J2-M172 in India; (3) whether distribution of J2a-M410 in India is more dominant in castes than in tribes; and (4) what social, linguistic or geographical factors might have influenced the distribution of J2-M172 subclades. To address the above, we investigated the presence of different subclades of HG J2-M172 in diverse set of Indian populations and their affinities with rest of the world.” ref
Origin, Diffusion, and Differentiation of Y-Chromosome Haplogroups E and J: Inferences on the Neolithization of Europe and Later Migratory Events in the Mediterranean Area
“Abstract: The phylogeography of Y-chromosome haplogroups E (Hg E) and J (Hg J) was investigated in >2,400 subjects from 29 populations, mainly from Europe and the Mediterranean area but also from Africa and Asia. The observed 501 Hg E and 445 Hg J samples were subtyped using 36 binary markers and eight microsatellite loci. Spatial patterns reveal that (1) the two sister clades, J-M267 and J-M172, are distributed differentially within the Near East, North Africa, and Europe; (2) J-M267 was spread by two temporally distinct migratory episodes, the most recent one probably associated with the diffusion of Arab people; (3) E-M81 is typical of Berbers, and its presence in Iberia and Sicily is due to recent gene flow from North Africa; (4) J-M172(xM12) distribution is consistent with a Levantine/Anatolian dispersal route to southeastern Europe and may reflect the spread of Anatolian farmers; and (5) E-M78 (for which microsatellite data suggest an eastern African origin) and, to a lesser extent, J-M12(M102) lineages would trace the subsequent diffusion of people from the southern Balkans to the west. A 7%–22% contribution of Y chromosomes from Greece to southern Italy was estimated by admixture analysis.” ref
“The arrival of haplogroup R1a-M417 in Eastern Europe, and the east-west diffusion of pottery through North Eurasia.” ref
R-M417 (R1a1a1)
“R1a1a1 (R-M417) is the most widely found subclade, in two variations which are found respectively in Europe (R1a1a1b1 (R-Z282) ([R1a1a1a*] (R-Z282) and Central and South Asia (R1a1a1b2 (R-Z93) ([R1a1a2*] (R-Z93).” ref
R-Z282 (R1a1a1b1a) (Eastern Europe)
“This large subclade appears to encompass most of the R1a1a found in Europe.
- R1a1a1b1a [R1a1a1a*] (R-Z282*) occurs in northern Ukraine, Belarus, and Russia at a frequency of c. 20%.
- R1a1a1b1a3 [R1a1a1a1] (R-Z284) occurs in Northwest Europe and peaks at c. 20% in Norway.
- R1a1a1c (M64.2, M87, M204) is apparently rare: it was found in 1 of 117 males typed in southern Iran.” ref
R1a1a1b2 (R-Z93) (Asia)
“This large subclade appears to encompass most of the R1a1a found in Asia, being related to Indo-European migrations (including Scythians, Indo-Aryan migrations, and so on).
- R-Z93* or R1a1a1b2* (R1a1a2* in Underhill (2014)) is most common (>30%) in the South Siberian Altai region of Russia, cropping up in Kyrgyzstan (6%) and in all Iranian populations (1-8%).
- R-Z2125 occurs at highest frequencies in Kyrgyzstan and in Afghan Pashtuns (>40%). At a frequency of >10%, it is also observed in other Afghan ethnic groups and in some populations in the Caucasus and Iran.
- R-M434 is a subclade of Z2125. It was detected in 14 people (out of 3667 people tested), all in a restricted geographical range from Pakistan to Oman. This likely reflects a recent mutation event in Pakistan.
- R-M560 is very rare and was only observed in four samples: two Burushaski speakers (north Pakistan), one Hazara (Afghanistan), and one Iranian Azerbaijani.
- R-M780 occurs at high frequency in South Asia: India, Pakistan, Afghanistan, and the Himalayas. The group also occurs at >3% in some Iranian populations and is present at >30% in Roma from Croatia and Hungary.” ref
R-M458 (R1a1a1b1a1)
“R-M458 is a mainly Slavic SNP, characterized by its own mutation, and was first called cluster N. Underhill et al. (2009) found it to be present in modern European populations roughly between the Rhine catchment and the Ural Mountains and traced it to “a founder effect that … falls into the early Holocene period, 7.9±2.6 KYA.” M458 was found in one skeleton from a 14th-century grave field in Usedom, Mecklenburg-Vorpommern, Germany. The paper by Underhill et al. (2009) also reports a surprisingly high frequency of M458 in some Northern Caucasian populations (for example 27.5% among Karachays and 23.5% among Balkars, 7.8% among Karanogays and 3.4% among Abazas).” ref
Ceramic Mesolithic
“Pottery was found in Xianrendong cave, China, dating to 20,000–10,000 years ago. In North-Eastern Europe, Siberia, and certain southern European and North African sites, a “ceramic Mesolithic” can be distinguished between c. 9,000 to 5,850 years ago. Russian archaeologists prefer to describe such pottery-making cultures as Neolithic, even though farming is absent. This pottery-making Mesolithic culture can be found peripheral to the sedentary Neolithic cultures. It created a distinctive type of pottery, with point or knob base and flared rims, manufactured by methods not used by the Neolithic farmers. Though each area of Mesolithic ceramic developed an individual style, common features suggest a single point of origin. The earliest manifestation of this type of pottery may be in the region around Lake Baikal in Siberia. It appears in the Yelshanka culture on the Volga in Russia 9,000 years ago, and from there spread via the Dnieper-Donets culture to the Narva culture of the Eastern Baltic.” ref
“Spreading westward along the coastline it is found in the Ertebølle culture of Denmark and Ellerbek of Northern Germany, and the related Swifterbant culture of the Low Countries. A 2012 publication in the Science journal, announced that the earliest pottery yet known anywhere in the world was found in Xianrendong cave in China, dating by radiocarbon to between 20,000 and 19,000 years before present, at the end of the Last Glacial Period. The carbon 14 datation was established by carefully dating surrounding sediments. Many of the pottery fragments had scorch marks, suggesting that the pottery was used for cooking. These early pottery containers were made well before the invention of agriculture (dated to 10,000 to 8,000 BCE), by mobile foragers who hunted and gathered their food during the Late Glacial Maximum.” ref
Neman culture (7,100 to 5,000 years ago)
“The archaeological Neman culture (German: Memel-Kultur) existed from about 5100 to the 3rd millennium BCE, starting in the Mesolithic and continued into the middle Neolithic. It was located in the upper basin of the Neman River (present-day northern Poland, southern Lithuania, western Belarus and Kaliningrad Oblast). In the north, the Neman culture bordered the Kunda culture during the Mesolithic and the Narva culture during the Neolithic. The Neolithic began with the appearance of pottery in mid 5th millennium BCE. The Semi-Neolithic Neman culture was a successor of the Mesolithic Neman culture. Most of flint tools are very similar between both cultures. A new widespread development was knives with sharpened and flared point.” ref
“Pottery of the Neman culture had pointed bottoms and was made of clay mixed with organic matter or crushed quartzite. Some latter examples have flat bottoms. The vessels were a bit narrower and curvier than of the Narva culture. They were decorated with a thin layer of white clay and rows of small imprints around the rim. The rest of the vessel had diagonal stripes forming a pattern of a fishnet or more rows of small imprints. Some pottery found in settlements of the Neman culture was made by Narva culture. Such phenomenon is explained by trade of flint, which Narva culture in the north did not have. Towards the end of the Neman culture, the pottery became more varied and exhibits influence of the Rzucewo culture: imprints made by a cord or resembling a fir. Eventually, the culture was overtaken by the Globular Amphora culture and the Corded Ware culture. ” ref
Sintashta culture
“The Sintashta culture is a Middle Bronze Age archaeological culture of the Southern Urals, dated to the period c. 2200–1900 BCE. It is the first phase of the Sintashta–Petrovka complex, c. 2200–1750 BCE or around 4,200 to 3,750 years ago. The culture is named after the Sintashta archaeological site, in Chelyabinsk Oblast, Russia, and spreads through Orenburg Oblast, Bashkortostan, and Northern Kazakhstan. The Sintashta culture is thought to represent an eastward migration of peoples from the Corded Ware culture.” ref
“Sintashta settlements are estimated to have a population of between 200 and 700 individuals with economies that “heavily exploited domesticated cattle, sheep, and goats alongside horses with occasional hunting of wild fauna”. Anthony (2007) assumes that probably the people of the Sintashta culture spoke “Common-Indo-Iranian”. This identification is based primarily on similarities between sections of the Rig Veda, a religious text which includes ancient Indo-Iranian hymns recorded in Vedic Sanskrit, and the funerary rituals of the Sintashta culture as revealed by archaeology.” ref
“Some cultural similarities with Sintashta have also been found to be common with the Nordic Bronze Age of Scandinavia. There is linguistic evidence of interaction between Finno-Ugric and Indo-Iranian languages, showing influences from the Indo-Iranians into the Finno-Ugric culture. From the Sintashta culture the Indo-Iranian followed the migrations of the Indo-Iranians to Anatolia, the Iranian plateau and the Indian subcontintinent. From the 9th century BCE onward, Iranian languages also migrated westward with the Scythians back to the Pontic steppe where the proto-Indo-Europeans came from.” ref
“It is widely regarded as the origin of the Indo-Iranian languages (Indo-Iranic languages), whose speakers originally referred to themselves as the Arya. The earliest known chariots have been found in Sintashta burials, and the culture is considered a strong candidate for the origin of the technology, which spread throughout the Old World and played an important role in ancient warfare. Sintashta settlements are also remarkable for the intensity of copper mining and bronze metallurgy carried out there, which is unusual for a steppe culture. Among the main features of the Sintashta culture are high levels of militarism and extensive fortified settlements, of which 23 are known.” ref
“Because of the difficulty of identifying the remains of Sintashta sites beneath those of later settlements, the culture was only distinguished in the 1990s from the Andronovo culture. It was then recognised as a distinct entity, forming part of the “Andronovo horizon”. Koryakova (1998) concluded from their archaeological findings that the Sintashta culture originated from the interaction of the two precursors Poltavka culture and Abashevo culture. Allentoft et al. (2015) concluded from their genetic results that the Sintashta culture should have emerged from an eastward migration of peoples from the Corded Ware culture. In addition, Narasimshan et al. (2019) cautiously cite that “morphological data has been interpreted as suggesting that both Fedorovka and Alakul’ skeletons are similar to Sintashta groups, which in turn may reflect admixture of Neolithic forest HGs and steppe pastoralists, descendants of the Catacomb and Poltavka cultures.” ref
“Sintashta emerged during a period of climatic change that saw the already arid Kazakh steppe region become even colder and drier. The marshy lowlands around the Ural and upper Tobol rivers, previously favored as winter refuges, became increasingly important for survival. Under these pressures both Poltavka and Abashevo herders settled permanently in river valley strongholds, eschewing more defensible hill-top locations. Its immediate predecessor in the Ural-Tobol steppe was the Poltavka culture, an offshoot of the cattle-herding Yamnaya horizon that moved east into the region between 2800 and 2600 BCE. Several Sintashta towns were built over older Poltavka settlements or close to Poltavka cemeteries, and Poltavka motifs are common on Sintashta pottery.” ref
“Sintashta material culture also shows the influence of the late Abashevo culture, derived from the Fatyanovo-Balanovo culture, a collection of Corded Ware settlements in the forest steppe zone north of the Sintashta region that were also predominantly pastoralist. Radiocarbon dating indicates that the Sintashta culture dates to between c. 2200 and 1750 BCE, roughly contemporary with the associated Abashevo and Petrovka cultures. Some authors date the Petrovka culture slightly later, from c. 1900 BCE.” ref
“In Cis-Urals, burial sites Berezovaya and Tanabergen II showed Sintashta culture established there c. 2290–1750 BCE (68.2% probability), and the earliest values of this culture, in Trans-Urals, at the burial sites Sintashta II and Kamenny Ambar-5 (Kurgan 2) are c. 2200–2000 BCE. Chariots appear in southern Trans-Urals region in middle and late phases of the culture, c. 2050-1750 BC. Blöcher et al. (2023) consider Sintashta-Petrovka period came to an end in Trans-Urals c. 1900–1800 BCE.” ref
Genetics
“Allentoft et al. 2015 analyzed the remains of four individuals ascribed to the Sintastha culture. One male carried Y-haplogroup R1a and mt-J1c1b1a, while the other carried Y-R1a1a1b and mt-J2b1a2a. The two females carried U2e1e and U2e1h respectively. The study found a close autosomal genetic relationship between peoples of Corded Ware culture and Sintashta culture, which “suggests similar genetic sources of the two,” and may imply that “the Sintashta derives directly from an eastward migration of Corded Ware peoples.” ref
“Sintashta individuals and Corded Ware individuals both had a relatively higher ancestry proportion derived from the Central Europe, and both differed markedly in such ancestry from the population of the Yamnaya Culture and most individuals of the Poltavka Culture that preceded Sintashta in the same geographic region. Individuals from the Bell Beaker culture, the Unetice culture, and contemporary Scandinavian cultures were also found to be closely genetically related to Corded Ware. A particularly high lactose tolerance was found among Corded Ware and the closely related Nordic Bronze Age. In addition, the study found samples from the Sintashta culture to be closely genetically related to the succeeding Andronovo culture.” ref
“Narasimhan et al. 2019 analyzed the remains of several individuals associated with the Sintashta culture. mtDNA was extracted from two females buried at the Petrovka settlement. They were found to be carrying subclades of U2 and U5. The remains of fifty individuals from the fortified Sintastha settlement of Kamennyi Ambar was analyzed. This was the largest sample of ancient DNA ever sampled from a single site. The Y-DNA from thirty males was extracted. Eighteen carried R1a and various subclades of it (particularly subclades of R1a1a1), five carried subclades of R1b (particularly subclades of R1b1a1a), two carried Q1a and a subclade of it, one carried I2a1a1a, and four carried unspecified R1 clades. The majority of mtDNA samples belonged to various subclades of U, while W, J, T, H and K also occurred. A Sintashta male buried at Samara was found to be carrying R1b1a1a2 and J1c1b1a.” ref
“The authors of the study found the majority of Sintashta people (ca. 80%) to be closely genetically related to the people of the Corded Ware culture, the Srubnaya culture, the Potapovka culture, and the Andronovo culture. These were found to harbor mixed ancestry from the Yamnaya culture and peoples of the Central European Middle Neolithic, like the Globular Amphora culture. The remaining sampled Sintashta individuals belonged to various ancestral types different from the majority population, with affinities to earlier populations such as Eneolithic samples collected at Khvalynsk and hunter-gatherers from Tyumen Oblast in western Siberia. This indicates that the Sintashta settlement of Kamennyi Ambar was a cosmopolitan site that united a genetically heterogenous population in a single social group. Estimates based on DATES (Distribution of Ancestry Tracts of Evolutionary Signals) suggest that genetic characteristics typical of the Sintashta culture formed by c. 3200 BCE.” ref
Warfare
“The preceding Abashevo culture was already marked by endemic intertribal warfare; intensified by ecological stress and competition for resources in the Sintashta period. This drove the construction of fortifications on an unprecedented scale and innovations in military technique such as the invention of the war chariot. Increased competition between tribal groups may also explain the extravagant sacrifices seen in Sintashta burials, as rivals sought to outdo one another in acts of conspicuous consumption analogous to the North American potlatch tradition.” ref
“Sintashta artefact types such as spearheads, trilobed arrowheads, chisels, and large shaft-hole axes were taken east. Many Sintashta graves are furnished with weapons, although the composite bow associated later with chariotry does not appear. Higher-status grave goods include chariots, as well as axes, mace-heads, spearheads, and cheek-pieces. Sintashta sites have produced finds of horn and bone, interpreted as furniture (grips, arrow rests, bow ends, string loops) of bows; there is no indication that the bending parts of these bows included anything other than wood. Arrowheads are also found, made of stone or bone rather than metal. These arrows are short, 50–70 cm long, and the bows themselves may have been correspondingly short.” ref
“Sintashta culture, and the chariot, are also strongly associated with the ancestors of modern domestic horses, the DOM2 population. DOM2 horses originated from the Western Eurasia steppes, especially the lower Volga-Don, but not in Anatolia, during the late fourth and early third millennia BCE. Their genes may show selection for easier domestication and stronger backs.” ref
“The Sintashta economy came to revolve around copper metallurgy. Copper ores from nearby mines (such as Vorovskaya Yama) were taken to Sintashta settlements to be processed into copper and arsenical bronze. This occurred on an industrial scale: all the excavated buildings at the Sintashta sites of Sintashta, Arkaim and Ust’e contained the remains of smelting ovens and slag. Around 10% of graves, mostly adult male, contained artifacts related to bronze metallurgy (molds, ceramic nozzles, ore and slag remains, metal bars and drops). However, these metal-production related grave goods rarely co-occur with higher-status grave goods. This likely means that those who engaged in metal production were not at the top of the social-hierarchy, even though being buried at a cemetery evidences some sort of higher status.” ref
“Much of Sintashta metal was destined for export to the cities of the Bactria–Margiana Archaeological Complex (BMAC) in Central Asia. The metal trade between Sintashta and the BMAC for the first time connected the steppe region to the ancient urban civilisations of the Near East: the empires and city-states of modern Iran and Mesopotamia provided a large market for metals. These trade routes later became the vehicle through which horses, chariots and ultimately Indo-Iranian-speaking people entered the Near East from the steppe.” ref
Genetic history of Europe, Genetics and archaeogenetics of South Asia, and Genetic history of the Middle East
PIE Speakers and Haplogroups R1b as well as R1a
“According to three autosomal DNA studies, haplogroups R1b and R1a, now the most common in Europe (R1a is also very common in South Asia) would have expanded from the Pontic steppes, along with the Indo-European languages; they also detected an autosomal component present in modern Europeans which was not present in Neolithic Europeans, which would have been introduced with paternal lineages R1b and R1a, as well as Indo-European languages. Studies that analyzed ancient human remains in Ireland and Portugal suggest that R1b was introduced in these places along with autosomal DNA from the Pontic steppes.” ref
“The subclade R1a1a (R-M17 or R-M198) is most commonly associated with Indo-European speakers. Data so far collected indicate that there are two widely separated areas of high frequency, one in Eastern Europe, around Poland and the Russian core, and the other in South Asia, around Indo-Gangetic Plain. The historical and prehistoric possible reasons for this are the subject of on-going discussion and attention amongst population geneticists and genetic genealogists, and are considered to be of potential interest to linguists and archaeologists also. Ornella Semino et al. propose a postglacial (Holocene) spread of the R1a1 haplogroup from north of the Black Sea during the time of the Late Glacial Maximum, which was subsequently magnified by the expansion of the Kurgan culture into Europe and eastward.” ref
“A large, 2014 study by Underhill et al., using 16,244 individuals from over 126 populations from across Eurasia, concluded there was compelling evidence, that R1a-M420 originated in the vicinity of Iran. The mutations that characterize haplogroup R1a occurred ~10,000 years ago. Its defining mutation (M17) occurred about 10,000 to 14,000 years ago. Pamjav et al. (2012) believe that R1a originated and initially diversified either within the Eurasian Steppes or the Middle East and Caucasus region.” ref
Yamnaya culture
“All Yamnaya individuals sampled by Haak et al. (2015) belonged to the Y-haplogroup R1b. According to Jones et al. (2015) and Haak et al. (2015), autosomal tests indicate that the Yamnaya-people were the result of admixture between “Eastern Hunter-Gatherers” from eastern Europe (EHG) and “Caucasus hunter-gatherers” (CHG). Each of those two populations contributed about half the Yamnaya DNA. According to co-author Dr. Andrea Manica of the University of Cambridge:
The question of where the Yamnaya come from has been something of a mystery up to now […] we can now answer that, as we’ve found that their genetic make-up is a mix of Eastern European hunter-gatherers and a population from this pocket of Caucasus hunter-gatherers who weathered much of the last Ice Age in apparent isolation.” ref
“Based on these findings and by equating the people of the Yamnaya culture with the Proto-Indo-Europeans, David W. Anthony (2019) suggests that the Proto-Indo-European language formed mainly from a base of languages spoken by Eastern European hunter-gathers with influences from languages of northern Caucasus hunter-gatherers, in addition to a possible later influence from the language of the Maikop culture to the south (which is hypothesized to have belonged to the North Caucasian family) in the later neolithic or Bronze Age involving little genetic impact.” ref
Eastern European hunter-gatherers
“According to Haak et al. (2015), “Eastern European hunter-gatherers” who inhabited Russia were a distinctive population of hunter-gatherers with high affinity to a ~24,000-year-old Siberian from the Mal’ta-Buret’ culture, or other, closely related Ancient North Eurasian (ANE) people from Siberia and to the Western Hunter-Gatherers (WHG). Remains of the “Eastern European hunter-gatherers” have been found in Mesolithic or early Neolithic sites in Karelia and Samara Oblast, Russia, and put under analysis. Three such hunter-gathering individuals of the male sex have had their DNA results published. Each was found to belong to a different Y-DNA haplogroup: R1a, R1b, and J. R1b is also the most common Y-DNA haplogroup found among both the Yamnaya and modern-day Western Europeans. R1a is more common in Eastern Europeans and in the northern parts of the Indian subcontinent.” ref
Near East population
“The Near East population were most likely hunter-gatherers from the Caucasus (CHG) c.q. Iran Chalcolithic related people with a major CHG-component. Jones et al. (2015) analyzed genomes from males from western Georgia, in the Caucasus, from the Late Upper Palaeolithic (13,300 years old) and the Mesolithic (9,700 years old). These two males carried Y-DNA haplogroup: J* and J2a. The researchers found that these Caucasus hunters were probably the source of the farmer-like DNA in the Yamnaya, as the Caucasians were distantly related to the Middle Eastern people who introduced farming in Europe.” ref
“Their genomes showed that a continued mixture of the Caucasians with Middle Eastern took place up to 25,000 years ago, when the coldest period in the last Ice Age started. According to Lazaridis et al. (2016), “a population related to the people of the Iran Chalcolithic contributed ~43% of the ancestry of early Bronze Age populations of the steppe.” According to Lazaridis et al. (2016), these Iranian Chalcolithic people were a mixture of “the Neolithic people of western Iran, the Levant, and Caucasus Hunter-Gatherers.” Lazaridis et al. (2016) also note that farming spread at two places in the Near East, namely the Levant and Iran, from where it spread, Iranian people spreading to the steppe and south Asia.” ref
Northern and Central Europe
“Haak et al. (2015) studied DNA from 94 skeletons from Europe and Russia aged between 3,000 and 8,000 years old. They concluded that about 4,500 years ago there was a major influx into Europe of Yamnaya culture people originating from the Pontic–Caspian steppe north of the Black Sea and that the DNA of copper-age Europeans matched that of the Yamnaya. The four Corded Ware people could trace an astonishing three-quarters of their ancestry to the Yamnaya, according to the paper. That suggests a massive migration of Yamnaya people from their steppe homeland into Eastern Europe about 4500 years ago when the Corded Ware culture began, perhaps carrying an early form of Indo-European language.” ref
Bronze age Greece
“A 2017 archaeogenetics study of Mycenaean and Minoan remains published in the journal Nature concluded that the Mycenaean Greeks were genetically closely related with the Minoans but unlike the Minoans also had a 13-18% genetic contribution from Bronze Age steppe populations.” ref
Haplogroup R1a
“Haplogroup R1a, or haplogroup R-M420, is a human Y-chromosome DNA haplogroup that is distributed in a large region in Eurasia, extending from Scandinavia and Central Europe to southern Siberia and South Asia. While R1a originated ca. 22,000 to 25,000 years ago, its subclade M417 (R1a1a1) diversified ca. 5,800 years ago. The place of origin of the subclade plays a role in the debate about the origins of Proto-Indo-Europeans.” ref
“The split of R1a (M420) is computed to ca. 22,000 or 25,000 years ago, which is the time of the last glacial maximum. A 2014 study by Peter A. Underhill et al., using 16,244 individuals from over 126 populations from across Eurasia, concluded that there was “a compelling case for the Middle East, possibly near present-day Iran, as the geographic origin of hg R1a.” The ancient DNA record has shown the first R1a during the Mesolithic in Eastern Hunter-Gatherers (from Eastern Europe), and the earliest case of R* among Upper Paleolithic Ancient North Eurasians, from which the Eastern Hunter-Gatherers predominantly derive their ancestry. No early samples of R1a have so far been found in Iran.” ref
“According to Underhill et al. (2014), the downstream R1a-M417 subclade diversified into Z282 and Z93 circa 5,800 years ago “in the vicinity of Iran and Eastern Turkey.” Even though R1a occurs as a Y-chromosome haplogroup among various languages such as Slavic and Indo-Iranian, the question of the origins of R1a1a is relevant to the ongoing debate concerning the urheimat of the Proto-Indo-European people, and may also be relevant to the origins of the Indus Valley Civilization. R1a shows a strong correlation with Indo-European languages of Southern and Western Asia, Central and Eastern Europe and to some extent Scandinavia being most prevalent in Eastern Europe, West Asia, and South Asia. In Europe, Z282 is prevalent, particularly while in Asia Z93 dominates. The connection b