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R-U106 (Y-DNA)

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This project is a meeting place for users who share the R-U106 Y-DNA haplogroup, which means they are related along their paternal lines. Users in this group may want to share their family trees with each other to find overlaps and merge duplicate profiles in order to join or expand the World Family Tree and discover new relatives.

Background

R1b-U106 is one of the large Western European subclades of haplogroup R along with R1b-P312 (Proto-Celtic) and R1b-S1194 (potentially Nordwestblock / Belgae). Balkano-Anatolian R1b-Z2103 that earlier split off from R1b-M269 is considered a Proto-Indo-Iranian subclade (Dorians, Hittites, Trojans, Phrygians, and Armenians) like R1a-Z93 (Indo-Aryans, Persians, Medes, Mitanni, and Tatars). R1b-U106 is considered Proto-Germanic.

Time to Most Recent Common Ancestor

Geographical Origins

All in all, we know that Wielbark probably represented the initial migration period of East Germanic tribes, traditionally believed to be from Northern Scandinavia, into territory later inhabited by Slavic tribes (and potentially earlier by a Balto-Slavic community).

In the 2nd century eastern German tribes – the Goths, who left monuments of the Wielbark archeological culture on the territories of modern Poland. They moved from the Southern Baltic to the territories in the northern part of the Black Sea. This caused the formation of a new type of the ancient life style, united by a common name – the Chernyahivska culture, the area of which during its prosperity (3rd–4th centuries) covered Southern East of Poland, the bigger part of modern Ukrainian and Moldavian lands and the bordering with them regions of Romania, and Russia.

On the basis of the Wielbark cultural element present on the Chernyahivska culture’s monuments, it was established that in the upper area of the Southern Bug Goths were divided: one group moved towards the Black Sea and the Azov Sea area, the second one turned south-west. Those who lived west of the Dnieper, were called the Tervingi, or the Visigoths. Those who moved to its left bank were called the Greutungs and the Ostrogoths. At different stages of the development of the Chernyahivska culture the Goths actively interacted with the other German tribes, in particular: the Heruli, the Vandals, the Taifals, the Burgundes, the Peukini or the Boranes.

For some time the Gothes headed big military-political unions, occurring in the Chernyahivska culture, uniting the late Scythes, Sarmatians, Getons and Dacians, and the Slavs. The Goths reached the biggest power during the reign of the Greutung king – Ermanaric (330–375 A.D.). In 375 A.D. the Goths’ state was defeated by the Huns. Afterwards the majority of the Greutungs were forced back to the west bank of the Dniester. Soon afterwards, under pressure of the Hun conquest, the Goths’ left the area of the Chernyahivska culture (in Southern, Central, and Western Europe, and even in Northern Africa, where through the Biskaysky channel the fighting tribe of vandals moved), the rest was dissolved among the local people of different tribes, having left no significant impact on its material and spiritual culture. The Goths stayed in Crimea for the longest period of time. The last records were Crimean Goths – the inhabitants of cave cities Mangup-Kale, Eski-Kermen and others – were mentioned in the written sources of the 17th century.

Y-DNA SNP Reference Trees & Tools

Testing

Y-DNA Next Generation Sequencing (NGS) is recommended over STR or individual SNP testing to confirm haplogroup R-U106 and downstream SNPs (NGS at least 15 Mbp at 30x depth). Whole Genome Sequencing (WGS) will also provide up to 100% Y-DNA coverage, but is not required.

Autosomal DNA (atDNA)

atDNA testing can help locate additional candidates for testing.

Whole Genome Sequencing (WGS) & Health

Contribution

It is recommended you share your BAM files with NGS Y-DNA scientists & researchers.

Sharing & Additional Matches

If you have an Autosomal DNA kit (atDNA), it is recommended you share/link your kit with relevant sites to locate additional matches.

Additional Resources

Ancient DNA

R1b computed Jan 04 2023 using YFull v10.08.00
Total in System: 32620 samples, 57 ancient
R1b on YFull has 620 ancient or scientific study samples not in the FTDNA Big Y Tree

Historical Context

European early modern humans (EEMH) lineages between 40 and 26 ka (Aurignacian) were still part of a large Western Eurasian "meta-population", related to Central and Western Asian populations. Divergence into genetically distinct sub-populations within Western Eurasia is a result of increased selection pressure and founder effects during the Last Glacial Maximum (LGM, Gravettian).

By the end of the LGM, after 20 ka, A Western European lineage, dubbed West European Hunter-Gatherer (WHG) emerged from the Solutrean refugium during the European Mesolithic.

These mesolithic hunter-gatherer cultures are subsequently replaced in the Neolithic Revolution as a result of the arrival of Early European Farmers (EEF) lineages derived from mesolithic populations of West Asia (Anatolia and the Caucasus).

In the European Bronze Age, there were again substantial population replacements in parts of Europe by the intrusion of Western Steppe Herders (WSH) lineages from the Pontic–Caspian steppes, being deeply related to Mesolithic European hunter-gatherers. These Bronze Age population replacements are associated with the Bell Beaker and Corded Ware cultures archaeologically and with the Indo-European expansion linguistically.

As a result of the population movements during the Mesolithic to Bronze Age, modern European populations are distinguished by differences in WHG, EEF and Ancient North Eurasian (ANE) ancestry. Admixture rates varied geographically; in the late Neolithic, WHG ancestry in farmers in Hungary was at around 10%, in Germany around 25% and in Iberia as high as 50%. The contribution of EEF is more significant in Mediterranean Europe, and declines towards northern and northeastern Europe, where WHG ancestry is stronger; the Sardinians are considered to be the closest European group to the population of the EEF.

Ethnogenesis of the modern ethnic groups of Europe in the historical period is associated with numerous admixture events, primarily those associated with the Roman during the Migration period and the Germanic, Norse, Slavic and Turkic expansions.

Due to natural selection, the percentage of Neanderthal DNA in ancient Europeans gradually decreased over time. From 45,000 BP to 7,000 BP, the percentage dropped from around 3–6% to 2%. The removal of Neanderthal-derived alleles occurred more frequently around genes than other parts of the genome.

Upper Paleolithic

It is thought that modern humans began to inhabit Europe during the Upper Paleolithic about 40,000 years ago. Some evidence shows the spread of the Aurignacian culture.: 59 

From a purely patrilineal, Y-chromosome perspective, it is possible that the old Haplogroup C1a2, F and/or E may be those with the oldest presence in Europe. They have been found in some very old human remains in Europe. However, other haplogroups are far more common among living European males because of later demographic changes.

Haplogroup I (M170), which is now relatively common and widespread within Europe, may represent a Palaeolithic marker – its age has been estimated at ~ 22,000 BP. While it is now concentrated in Europe, it probably arose in a male from the Middle East or Caucasus, or their near descendants, c. 20–25,000 years BP, when it diverged from its immediate ancestor, haplogroup IJ. At about this time, an Upper Palaeolithic culture also appeared, known as the Gravettian.

Earlier research into Y-DNA had instead focused on haplogroup R1 (M173): the most populous lineage among living European males; R1 was also believed to have emerged ~ 40,000 BP in Central Asia. However, it is now estimated that R1 emerged substantially more recently: a 2008 study dated the most recent common ancestor of haplogroup IJ to 38,500 and haplogroup R1 to 18,000 BP. This suggested that haplogroup IJ colonists formed the first wave and haplogroup R1 arrived much later.

Last Glacial Maximum

The Last Glacial Maximum ("LGM") started c. 30 ka BCE, at the end of MIS 3, leading to a depopulation of Northern Europe. According to the classical model, people took refuge in climatic sanctuaries (or refugia) as follows:

• Northern Iberia and Southwest France, together making up the "Franco-Cantabrian" refugium

• The Balkans

• Ukraine and more generally the northern coast of the Black Sea

• Italy.

This event decreased the overall genetic diversity in Europe, a "result of drift, consistent with an inferred population bottleneck during the Last Glacial Maximum". As the glaciers receded from about 16,000–13,000 years ago, Europe began to be slowly repopulated by people from refugia, leaving genetic signatures.

Some Y haplogroup I clades appear to have diverged from their parental haplogroups sometime during or shortly after the LGM. Haplogroup I2 is prevalent in the western Balkans, as well as the rest of southeastern and central-eastern Europe in more moderate frequencies. Its frequency drops rapidly in central Europe, suggesting that the survivors bearing I2 lineages expanded predominantly through south-eastern and central-eastern Europe.

Cinnioglu sees evidence for the existence of an Anatolian refuge, which also harboured Hg R1b1b2. Today, R1b dominates the y chromosome landscape of western Europe, including the British Isles, suggesting that there could have been large population composition changes based on migrations after the LGM.

Semino, Passarino and Pericic place the origins of haplogroup R1a within the Ukrainian ice-age refuge. Its current distribution in eastern Europe and parts of Scandinavia are in part reflective of a re-peopling of Europe from the southern Russian/Ukrainian steppes after the Late Glacial Maximum.

From a study of 51 individuals, researchers were able to identify five separate genetic clusters of ancient Eurasians during the LGM: the Věstonice Cluster (34,000–26,000 years ago), associated with the Gravettian culture; the Mal'ta Cluster (24,000–17,000), associated with the Mal'ta-Buret' culture, the El Mirón Cluster (19,000–14,000 years ago), associated with the Magdalenian culture; the Villabruna Cluster (14,000–7,000 years ago) and the Satsurblia Cluster (13,000 to 10,000 years ago).

From around 37,000 years ago, all ancient Europeans began to share some ancestry with modern Europeans. This founding population is represented by GoyetQ116-1, a 35,000 year old specimen from Belgium. This lineage disappears from the record and is not found again until 19,000 BP in Spain at El Mirón, which shows strong affinities to GoyetQ116-1.

During this interval, the distinct Věstonice Cluster is predominant in Europe, even at Goyet. The re-expansion of the El Mirón Cluster coincided with warming temperatures following the retreat of the glaciers during the Last Glacial Maximum. From 37,000 to 14,000 years ago, the population of Europe consisted of an isolated population descended from a founding population that didn't interbreed significantly with other population

Mesolithic

Mesolithic (post-LGM) populations had diverged significantly due to their relative isolation over several millennia, to the harsh selection pressures during the LGM, and to the founder effects caused by the rapid expansion from LGM refugia in the beginning Mesolithic. By the end of the LGM, around 19 to 11 ka, the familiar varieties of Eurasian phenotypes had emerged. However, the lineage of Mesolithic hunter-gatherers of Western Europe (WHG) does not survive as a majority contribution in any modern population. They were most likely blue eyed, and retained the dark skin pigmentation of pre-LGM EEMH. The HERC2 and OCA2 variations for blue eyes are derived from the WHG lineage were also found in the Yamnaya people.

Around 14,000 years ago, the Villabruna Cluster shifted away from GoyetQ116-1 affinity and started to show more affinity with the Near East, a shift which coincided with the warming temperatures of the Bølling-Allerød interstadial. This genetic shift shows that Near East populations had probably already begun moving into Europe during the end of the Upper Paleolithic, about 6,000 years earlier than previously thought, before the introduction of farming. A few specimens from the Villabruna Cluster also show genetic affinities for East Asians that are derived from gene flow. The HERC2 variation for blue eyes first appears around 13,000 to 14,000 years ago in Italy and the Caucasus. The light skin pigmentation characteristic of modern Europeans is estimated to have spread across Europe in a "selective sweep" during the Mesolithic (19 to 11 ka). The associated TYRP1 SLC24A5 and SLC45A2 alleles emerge around 19 ka, still during the LGM, most likely in the Caucasus.

In 2009, DNA was extracted from the remains of a male hunter-gatherer from Kostenki-12 who lived circa 30,000 BP and died aged 20–25. His maternal lineage was found to be mtDNA haplogroup U2. He was buried in an oval pit in a crouched position and covered with red ochre. Kostenki 12 was later found to belong to the patrilineal Y-DNA haplogroup C1* (C-F3393).

A male from Kostenki-14 (Markina Gora), who lived approximately 38,700–36,200 year ago, was also found to belong to mtDNA haplogroup U2. His Y-DNA haplogroup was C1b* (C-F1370).

The Kostenki-14 genome represents early evidence for the separation of Europeans and East Asian lineages. It was found to have a close relationship to both "Mal'ta boy" (24 ka) of south-east Siberia (Ancient North Eurasian) and to the later Mesolithic hunter-gatherers of Europe and western Siberia, as well as with a basal population ancestral to Early European Farmers, but not to East Asians. Yang et al. 2020 found that the early hunter-gatherers lineage of Kostenski-14 may have contributed (c. 68%) ancestry to the Ancient North Eurasian Yana and Mal'ta samples, with the remainder ancestry (c. 32%) being contributed from an East-Eurasian Tianyuan-related population. Kostenki-14 had some level of ancient Neanderthal admixture, which has been dated as going back to circa 54,000 BP.

Fu et al. (2016) examined the remains of fourteen Gravettians. The eight male included three samples of Y-chromosomal haplogroup CT, one of I, one IJK, one BT, one C1a2, and one sample of F.

Teschler et al. (2020) examined the remains of one adult male and two twin boys from a Gravettian site in Austria. All belonged to haplogroup Y-Haplogroup I. and all had the same mtDNA, U5.

The remains were covered in red ochre, a compound known to have religious significance, indicating that this woman’s burial was ceremonial in nature.

In the Vestonice 13 sample, the Y chromosomal haplogroup CT (not IJK-L16) (CTS109+, CTS5318+, CTS6327+, CTS8243+, CTS9556+, Z17718+, Y1571+, M5831+) was determined, for the Vestonice 15 sample, the Y chromosome haplogroup BT (PF1178+), in the Vestonice 43 sample, the Y chromosome haplogroup F (not I) (P145+, P158+). In the Vestonice 16 sample, the Y chromosomal haplogroup C1a2 (V20+, V86+).

In 2016, researchers successfully extracted DNA from several ancient human fossils at Goyet (with direct dates): GoyetQ116-1 (35,160-34,430 BP) and GoyetQ376-3 (33,940-33,140 BP) from the Aurignacian; GoyetQ376-19 (27,720-27,310 BP), GoyetQ53-1 (28,230-27,720 BP), GoyetQ55-2 (27,730-27,310 BP), GoyetQ56-16 (26,600-26,040 BP) and Goyet2878-21 (27,060-26,270 BP) from the Gravettian; and GoyetQ-2 (15,230-14,780 BP) from the Magdalenian.

GoyetQ376-19, Goyet53-1 and Goyet56-16 were found to cluster genetically with several other Gravettian individuals from Europe in the Věstonice Cluster, while GoyetQ-2 was found to cluster genetically with several other Magdalenian individuals from Europe in the El Mirón Cluster.

All later Europeans after GoyetQ116-1 show some genetic affinity for this individual. GoyetQ116-1 also exhibits more genetic affinity for the Tianyuan man than any other ancient individual from West Eurasia. Culturally, the Aurignacian cultural complex is chronologically associated with the human remains of Goyet Q116-1, while the subsequent Gravettian is associated with the Vestonice cluster.

In the single dispersal Out of Africa theory, it is believed that populations related to the Initial Upper Palaeolithic population of Bacho Kiro cave contributed ancestry to later Asian populations, because of genetic similarity and to some early West Europeans such as the c. 35,000 year old individual from the Goyet Caves, Belgium, known as 'GoyetQ116-1'. Populations related to these earlier individuals did not contribute detectable ancestry to later European populations.

Ust'-Ishim man belongs to Y-DNA haplogroup K2. The two subclades of K2 are K2a and K2b In the original paper, he was classified only as Haplogroup K-M9 (KxLT).

Research by Poznik et al. (2016) suggests that Oase 1 Y-DNA belongs to haplogroup K2a*.

He belonged to mitochondrial DNA haplogroup B, and his Y-chromosomal haplogroup was K2b.

Tianyuan man exhibits a unique genetic affinity for GoyetQ116-1 from the Goyet Caves in Namur province, Belgium. GoyetQ116-1 shares more alleles with Tianyuan man than does any other sampled ancient individual from West Eurasia

The individuals at Sungir show closest genetic affinity to the individuals from Kostenki, while showing closer affinity to the individual from Kostenki 12 than to the individual from Kostenki 14. The Sungir individuals descended from a lineage that was related to the individual from Kostenki 14, but were not directly related. The individual from Kostenki 12 was also found to be closer to the Sungir individuals than to the individual from Kostenki 14. The Sungir individuals also show close genetic affinity to various individuals belonging to Vestonice Cluster buried in a Gravettian context, such as those excavated from Dolní Věstonice.

DNA analysis shows that the medieval individual Sungir 6 (730-850 cal BP) belongs to mtDNA Haplogroup W3a1, and Y-DNA Haplogroup I2a1b2 (I-A16681).

In 2022 nuclear and mitochondrial DNA from a female found in the cave was analysed. Like human remains from other Magdalenian sites, her genome shares most drift with the individuals belonging to the ~19,000–14,000-year-old Goyet Q2 genetic cluster.

3D microscopy showed that the flesh had been removed from the bones using the same tools and techniques used on animal bones. The human skulls of the same date found at the cave around 1987, may have been deliberately fashioned into ritual drinking cups or bowls. These de-fleshing marks and secondary treatment of human material at Gough's Cave, also found at other Magdalenian culture sites such as Brillenhöhle and Hohle Fels in Germany and Maszycka Cave in Poland, has been taken as evidence of cannibalism.

In 1903 the remains of a human male, since named Cheddar Man, were found a short distance inside Gough's Cave. He is Britain's oldest complete human skeleton, having been dated to approximately 7150 BC.

In a genetic study published in Nature in March 2023, the authors found that the ancestors of the WHGs were populations associated with the Epigravettian culture, which largely replaced populations associated with the Magdalenian culture about 14,000 years ago (the ancestors of the Magdalenian-associated individuals were the populations associated with the western Gravettian, Solutrean and Aurignacian cultures). In the study, WHG ancestry is renamed 'Oberkassel ancestry', first found north of the Alps in two 14,000 year-old individuals at the eponymous site at Oberkassel, who can be modeled as an admixture of Villabruna ancestry (itself modeled as an admixture between a lineage related to the Věstonice cluster and a lineage ancestral to the Kostenki-14 and Goyet Q116-1 individuals), and Goyet-Q2 ancestry related to individuals found in Europe prior to the Last Glacial Maximum. The study states that all of the individuals of the Oberkassel cluster could be modeled as c. 75% Villabruna and 25% Goyet-Q2 ancestry or, alternatively, as c. 90% Villabruna and 10% Fournol ancestry, a newly identified cluster described as a sister lineage of the Goyet Q116-1 ancestry found in individuals associated with the Gravettian culture of southwestern Europe.

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.

A grave that contained a well-preserved skeleton was discovered at the base of the archaeological layers in 1988. Direct AMS dating of the skeletal remains revealed an age of 14,160 to 13,820 years.

Villabruna 1 is significant in terms of the history of population genetics: the remains were found to carry Y-DNA haplogroup R1b1a-L754* (xL389,V88). This is the oldest documented example of haplogroup R1b found anywhere.

About 85% of his ancestry can be modelled as coming from the c. 14,000–7,000-year-old Villabruna genetic cluster, and only c. 15% from the Goyet Q2 cave cluster whose genes are found in association with the Late Upper Palaeolithic Magdalenian culture. He is not closely related to the earlier Magdalenian individuals found in the same cave, whose ancestry is entirely from the Goyet cluster. The genomes of all British Mesolithic individuals sequenced to date other than Cheddar Man can be modelled as only Villabruna-related (WHG) ancestry, without additional Goyet-related admixture.

Analysis of the 476,347 single nucleotide polymorphisms recovered from Kendricks_074 shows that he shares most drift with the individuals belonging to the ~14,000–7,000-year-old Villabruna genetic cluster. Eleven Mesolithic individuals from elsewhere in the British Isles, the Western Hunter-Gatherer population, can also be modeled as having entirely Villabruna ancestry, except for Cheddar Man with some 85% Villabruna ancestry.

However, a Palaeolithic individual from Gough’s Cave in SW England, who possibly lived at approximately the same time as Kendricks_074, shares most drift with the individuals belonging to the ~19,000–14,000-year-old Goyet Q2 genetic cluster. De-fleshing marks and secondary treatment of human material at Gough’s Cave (also found at other Magdalenian culture sites such as Brillenhöhle and Hohle Fels in Germany and Maszycka Cave in Poland) has been taken as evidence of cannibalism. This suggests that at least two different human groups, with different genetic affinities and different dietary and cultural behaviours, were present in Britain during the Late Glacial.

Loschbour man lived over 8,000 years ago, making the skeleton the oldest human remains found in the country. The remains contained Y-DNA of the Haplogroup I2a-M423*.

Prehistoric Europe

Genetic studies have confirmed that Early European Farmers can be modelled as Anatolian Neolithic Farmers with 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, and WHGs were displaced to the margins.

During the Chalcolithic and early Bronze Age, the Early European Farmer cultures were overwhelmed by new migrations from the Pontic–Caspian steppe by a group related to people of the Yamnaya culture who carried Western Steppe Herder ancestry and probably spoke Indo-European languages. Once again the populations mixed, and EEF ancestry is common in modern European populations, with EEF ancestry highest in Southern Europeans, especially Sardinians and Basque people.

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) around 43,000 BC, and to have split from Caucasian Hunter-Gatherers (CHGs) around 23,000 BC.

Genetic studies demonstrate that the introduction of farming to Europe in the 7th millennium BC 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. The Y-DNA of EEFs was typically types of haplogroup G2a, and to a lesser extent H, T, J, C1a2 and E1b1, while their mtDNA was diverse.

During the Chalcolithic and early Bronze Age, the EEF-derived cultures of Europe were overwhelmed by successive invasions 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. 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. EEF mtDNA however remained frequent, suggesting admixture between WSH males and EEF females.

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, Greeks and Iberians, with the lowest EEF ancestry found in modern Europeans ranging from 35% to 40% in modern Finns, Lithuanians and Latvians.

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.

Prehistoric British Isles & Iberia

Sample: I11149 (Male)
Location: England, Cambridgeshire, Teversham (Marshall’s) Evaluation
Age: 733-397 calBCE
Y-DNA: R-Z156
mtDNA: V

Yamnaya

The genes of the Yuzhny Oleni were transmitted to the people of the Yamnaya Culture, and to Scandinavia through a western route.

The ancestry of the Yamnaya people can be mostly modelled as an admixture of Eastern Hunter-Gatherers (EHGs) and a Near Eastern component related to Caucasus hunter-gatherers, Iranian Chalcolithic people, or a genetically similar population. Each of those two populations contributed about half the Yamnaya DNA.

Lithic artefacts, bone artefacts, charcoal, flax fibers, and pottery were discovered at the cave. The lithic artefacts show similarities to eastern Epigravettian sites. Perforated pendants made out of stalagmite and polished bovid bone were also discovered. The remains of yellow, red and brown ochre were also found at the site.

In 2013, archaeologists found a temporal bone fragment of an ancient human in the cave. Direct AMS dating of the bone yielded an estimated date of 13,300 BP for the age of the bone. Researchers successfully extracted DNA from the petrous part of the temporal bone and managed to recover low coverage genomes.

The ancient individual from Satsurblia was male with black hair and brown eyes; however, the individual is one of the earliest found to carry the derived HERC2 allele for blue eyes. The Satsurblia individual also likely had light skin, as he was found to carry the derived SLC24A5 allele for light skin. The Satsurblia individual was also lactose intolerant and did not carry the derived EDAR allele commonly found in East Asians and Native Americans.

The Satsurblia individual belongs to mtDNA Haplogroup K3 and Y-DNA Haplogroup J1-Y6313*. About 1.7-2.4% of the Satsurblia individual's DNA was Neanderthal in origin.

The Satsurblia individual is genetically closest to an ancient individual, dating to around 9,700 BP, found at the Kotias Klde rock shelter in Georgia. Together, they form a genetically distinct cluster referred to as Caucasus Hunter-Gatherer (CHG).

In comparison to modern human populations, the Satsurblia individual is closest to the modern population in Georgia.

The Caucasus hunter-gatherers contributed significantly to modern European populations by way of the Yamna people. Around half of the Yamna people's DNA come from the Caucasus hunter-gatherers. The Caucasus hunter-gatherers also contributed genetically to modern Central Asians and South Asians.

The expansion of WSHs resulted in the virtual disappearance of the Y-DNA of Early European Farmers (EEFs) from the European gene pool, significantly altering the cultural and genetic landscape of Europe.

Several genetic studies performed since 2015 have given support to the Kurgan theory of Marija Gimbutas regarding the Indo-European Urheimat – that Indo-European languages spread throughout Europe from the Eurasian steppes and that the Yamnaya culture were Proto-Indo-Europeans. According to those studies, haplogroups R1b and R1a, now the most common in Europe (with R1a also being common in South Asia), would have expanded from the Pontic–Caspian 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 in the Bronze Age.

The Sintashta culture is thought to represent an eastward migration of peoples from the Corded Ware culture. It is widely regarded as the origin of the Indo-Iranian languages (or Indo-Iranic languages) 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.

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".

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.

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.

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.

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.

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.

Farming began in Anatolia in what is present-day Turkey. But the DNA shows that the people who experimented with planting wheat and domesticating sheep and goats starting about 10,000 years ago weren’t simply descendants of earlier hunter-gatherers living in the area. Dozens of newly sequenced genomes suggest Anatolia absorbed at least two separate migrations from about 10,000 to 6500 years ago. One came from today’s Iraq and Syria and the other from the Eastern Mediterranean coast. In Anatolia they mixed with each other and with the descendants of earlier hunter-gatherers. By about 6500 years ago, the populations had coalesced into a distinct genetic signature.

Another genetic contribution came from the east about 6500 years ago, as hunter-gatherers from the Caucasus entered the region. Then about 5000 years ago, a fourth group—nomads from the steppes north of the Black Sea, known as the Yamnaya—arrived, adding to the genetic picture but not fundamentally redrawing it. “The people of the Southern Arc are mostly coming from Levantine, Anatolian, and Caucasus components,” Lazaridis says. “The Yamnaya are like a layer of sauce, added after 3000 B.C.E.”

Plague, Violence, Lactose Tolerance & Steppe Cowboy Migration

Genetic studies suggest that Funnelbeaker women were incorporated into the Corded Ware culture through intermixing with incoming Corded Ware males, and that people of the Corded Ware culture continued to use Funnelbeaker megaliths as burial grounds. Subsequent cultures of Late Neolithic, Bronze Age, and Iron Age Central Europe display strong maternal genetic affinity with the Funnelbeaker culture.

The evidence suggested that the Battle Axe culture entered Scandinavia through a migration from Eastern Europe, after which Battle Axe males mixed with Funnelbeaker females.

A genetic study conducted by Haak et al. (2015) found that a large proportion of the ancestry of the Corded Ware culture's population is similar to the Yamna culture, tracing the Corded Ware culture's origins to migrations of the Yamna from the steppes 4,500 years ago. About 75% of the DNA of late Neolithic Corded Ware skeletons found in Germany was a precise match to DNA from individuals of the Yamna culture. The same study estimated a 40–54% ancestral contribution of the Yamna in the DNA of modern Central & Northern Europeans, and a 20–32% contribution in modern Southern Europeans, excluding Sardinians (7.1% or less), and to a lesser extent Sicilians (11.6% or less). Haak et al. also note that their results "suggest" that haplogroups R1b and R1a "spread into Europe from the East after 3,000 BCE.

In terms of phenotypes, Wilde et al. (2014) and Haak et al. (2015) found that the intrusive Yamna population, generally inferred to be the first speakers of an Indo-European language in the Corded Ware culture zone, were overwhelmingly dark-eyed (brown), dark-haired and had a skin colour that was moderately light, though somewhat darker than that of the average modern European. These studies also showed that light pigmentation traits had already existed in pre-Indo-European Neolithic Europeans (in both farmers and hunter-gatherers), so long-standing philological attempts to correlate them with the arrival of Indo-Europeans from the steppes were misguided.

Autosomal DNA tests also indicate that the Yamna migration from the steppes introduced a component of ancestry referred to as "Ancient North Eurasian" admixture into Europe. "Ancient North Eurasian" is the name given in genetic literature to a component that represents descent from the people of the Mal'ta-Buret' culture or a population closely related to them. The "Ancient North Eurasian" genetic component is visible in tests of the Yamna people as well as modern-day Europeans, but not of Western or Central Europeans predating the Corded Ware culture.

Early papers publishing results on European-wide Y-DNA marker frequencies, such as those of Semino (2000) and Rosser (2000), correlated haplogroup R1b-M269 with the earliest episodes of European colonization by anatomically modern humans (AMH). The peak frequencies of M269 in Iberia (especially the Basque region) and the Atlantic façade were postulated to represent signatures of re-colonization of the European West following the Last Glacial Maximum. However, even prior to recent criticisms and refinements, the idea that Iberian R1b carrying males repopulated most of western Europe was not consistent with findings which revealed that Italian M269 lineages are not derivative of Iberian ones.

More recently, data and calculations from Myres et al. (2011), Cruciani et al. (2011) Arredi et al. (2007), and Balaresque et al. (2010) suggest a Late Neolithic entry of M269 into Europe.

These hypotheses appear to be corroborated by more direct evidence from ancient DNA. R1b was detected in two male skeletons from a German Bell Beaker site dated to 2600–2500 BCE at Kromsdorf, one of which tested positive for M269 but negative for its U106 subclade (note that the P312 subclade was not tested for), while for the other skeleton the M269 test was unclear. A later Bell Beaker male skeleton from Quedlinburg, Germany dated to 2296–2206 BCE tested positive for R1b M269 P312 subclade. Ancient Y-DNA results for the remains of Beaker people from Iberia have yet to be obtained.

By around 4000 BC, the island was populated by people with a Neolithic culture. This neolithic population had significant ancestry from the earliest farming communities in Anatolia, indicating that a major migration accompanied farming. The beginning of the Bronze Age and the Bell Beaker culture was marked by an even greater population turnover, this time displacing more than 90% of Britain's neolithic ancestry in the process. This is documented by recent ancient DNA studies which demonstrate that the immigrants had large amounts of Bronze-Age Eurasian Steppe ancestry, associated with the spread of Indo-European languages and the Yamnaya culture

Modern genetic studies show that its emergence was accompanied by large-scale migrations and genetic displacement. The Battle Axe culture initially absorbed the agricultural Funnelbeaker culture.

People of the Late Neolithic and Bronze Age cultures of Scandinavia were found to be very closely related people of the Corded Ware culture, Bell Beaker culture and Unetice culture, all of whom shared genetic affinity with the Yamnaya culture. The Sintashta culture and Andronovo culture of Central Asia also displayed close genetic relations to the Corded Ware culture.

Celtic Tribes

Later, when the West was better explored, it was found that tin actually came from two regions: Galicia, in the northwest of Iberia, and Devon and Cornwall in southwest Britain.

Belgic Tribes

Germanic Tribes

More than 99% of living men with I1 belong to the DF29 branch which is estimated to have emerged in 2400 BCE. All DF29 men share a common ancestor born between 2500 and 2400 BCE. The oldest ancient individual with I1-DF29 found is Oll009, a man from early Bronze Age Sweden.

Haplogroup I1, as well as subclades of R1b such as R1b-U106 and subclades of R1a such as R1a-Z284, are strongly associated with Germanic peoples and are linked to the proto-Germanic speakers of the Nordic Bronze Age. Current DNA research indicates that I1 was close to non-existent in most of Europe outside of Scandinavia and northern Germany before the Migration Period. The expansion of I1 is directly tied to that of the Germanic tribes. Starting around 900 BC, Germanic tribes started moving out of southern Scandinavia and northern Germany into the nearby lands between the Elbe and the Oder. Between 600 and 300 BC another wave of Germanics migrated across the Baltic Sea and settled alongside the Vistula. Germanic migration to that area resulted in the formation of the Wielbark culture, which is associated with the Goths.

Yet, the frequencies of three main subhaplogroups, namely R1b1b2a1 (R-U106), R1b1b2a2* (R-P312*) and R1b1b2a2g (R-U152), could be reconstructed for two northern French regions Nord-Pas-de-Calais and Île-de-France from data already available (Ramos-Luis et al., 2009; Busby et al., 2012).

In the French surname group, however, frequencies of R-U106 and R-U152 correspond with those observed in Northern France and not with those in the Flemish surname group (R-U106 with 26.78% in AFS and 12.00% in FRS, R-U152 with 10.56% in AFS and 16.00% in FRS; Tables 1 and 2).

A Germanic tribe known as the Eburones had originally inhabited the present-day Cologne Lowland. But they were wiped out in a war of reprisal carried out by Julius Caesar. In 38 BC, the Germanic tribe known as the Ubii, who inhabited the right bank of the Rhine, were resettled by the Roman General Marcus Vipsanius Agrippa in the lands in the Cologne Lowland vacated by the Eburones. This brought the Ubii within Roman-occupied territory.

Sample: SZ2
Location: Szólád, Somogy County, Hungary
Age: Longobard 6th Century
Y-DNA: R-Z338
FTDNA Comment: Shares 5 SNPs with a man from the UK. Forms a new branch down of R-Z338 (U106). New branch = R-BY176786
mtDNA: T1a1

Sample: SZ4
Location: Szólád, Somogy County, Hungary
Age: Longobard 6th Century
Y-DNA: R-ZP200
FTDNA Comment: Splits R-ZP200 (U106). Derived (positive) for 2 SNPs and ancestral (negative) for 19 SNPs. New path = R-Y98441>R-ZP200
mtDNA: H1c9

Sample: SZ11
Location: Szólád, Somogy County, Hungary
Age: Longobard 6th Century
Y-DNA: R-FGC13492
FTDNA Comment: Shares 1 SNP with a man from Italy. Forms a new branch down of R-FGC13492 (U106). New branch = R-BY138397
mtDNA: K2a3a

Sample: SZ16
Location: Szólád, Somogy County, Hungary
Age: Longobard 6th Century
Y-DNA: R-U106
mtDNA: U4b1b

Slavic Tribes

Baltic Tribes

Celtic Galatia Tribes

Iranian Tribes

Turkic Tribes

The Oghuric tribes are also connected with the Hungarians, whose exo-ethnonym is usually believed to be derived from On-Oğur (> (H)Ungari). Hungarians -> Hun Oghur -> (ten oghur tribes): On ogur -> up.chv. Won ogur -> dow.chv. Wun ogur -> belor. Wugorac -> rus. Wenger -> slove. Vogr, Vogrin -> cheh. pol. Węgier, Węgrzyn, -> lit. Veñgras. The Hungarians are culturally of mixed Ugrian / Turkic heritage, with Oghuric-Bulgar and Khazar influences, even though much of the modern-day Hungarian genepool also has strong Slavic, Germanic, and Iranic influences.

Sample HU55 is derived for R-BY41605 (R1b1a1b1a1a1c2). The R-U106 haplogroup upstream of R-BY41605 is most common in western Europe, and quite frequent among Hungarian conquerors.

Hun Conquest, Roman Empire Collapse & Germanic Tribe Migration

So far 6 Y-chromosome Hg-s have been published from the Conquerors; which revealed the presence of N1a1- M46 (previously called Tat or N1c), in two out of 4 men, while detected two R1b-U106 and two I2a-M170 Hg-s.

Three out of 4 samples in the small Karos3 cemetery belonged to Hg R1b1a1b1a1a1-U106 setting apart this cemetery from all other groups, except for the Hun/2 sample which is the only other one with this Hg. Hg U106 is considered a “Germanic” branch as it is most significant today in Germany, Scandinavia, and Britain, and rare in Eastern Europe (Supplementary Table S4). Its ancestral branch Hg R1b1a1b-M262 is assumed to have emerged in the Pontic-Caspian Steppe and arrived to Europe with Bronze Age migrations25. Its presence in Hun and Conqueror samples may derive from Goths, Gepids or other German allies of the Huns.

According to Härke the more modern view is of co-existence between the British and the Anglo-Saxons. He suggests that several modern archaeologists have now re-assessed the traditional model, and have developed a co-existence model largely based on the Laws of Ine. The laws include several clauses that provide six different wergild levels for the Britons, of which four are below that of freeman. Although it was possible for the Britons to be rich freemen in Anglo-Saxon society, generally it seems that they had a lower status than that of the Anglo-Saxons.

By contrast, the early medieval population of England exhibits a substantial fraction of continental-derived haplotypes belonging to haplogroups R1b-U106, R1a-M420, I2a1-L460 and I1-M253, which are commonly found in northern and central Europe (and are also common among ancient continental individuals including the ones that we report). In particular, Y chromosomal haplogroups I1-M253 and R1a-M420 were absent from our Bronze, Iron and Roman Age British and Irish individuals, but were identified in more than one-third of our individuals from early medieval England.

Reconquista, Viking Age, Saxon Wars, Norman Conquest & Crusades

Sample: VK289 / Denmark_Bodkergarden Grav H, sk 1
Location: Bødkergarden, Langeland, Denmark
Age: Viking 9th century CE
Y-DNA: R-U106
mtDNA: J2b1a

Sample: VK290 / Denmark_Kumle Hoje Grav O
Location: Kumle_høje, Langeland, Denmark
Age: Viking 10th century CE
Y-DNA: R-FT264183
FTDNA Comment: Shares at least 4 SNPs with a man from Sweden, forming a new branch downstream R-FT263905 (U106). New branch = R-FT264183. > HG02545 remains at R-FT263905
mtDNA: I1a1

Sample: VK389 / Norway_Telemark 3697
Location: Telemark, Nor_South, Norway
Age: Viking 10th century CE
Y-DNA: R-Z27210
FTDNA Comment: Splits R-Z27210 (U106). Derived for 1 ancestral for 2. New path = R-Y32857>R-Z27210
mtDNA: T2b

Age of Discovery, Conquest, Colonization, Migration & Revolution

Y-chromosomal sub-haplogroup R-Z381* is a subgroup of R-U106, which has been found in Western Europe with the highest frequency of around 35% in the north of the Netherlands and in Denmark but with a steep frequency fall to the south as the frequency of R-U106 is only 7% in France.

Americas & Caribbean

Africa, Egypt & Ancient Near East

Asia, Australia & Pacific Islanders
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