First taste of Early Medieval DNA from the Ural region (Csaky et al. 2020 preprint)

Over at bioRxiv at this LINK. From the preprint:

The ancient Hungarians originated from the Ural region of Russia, and migrated through the Middle-Volga region and the Eastern European steppe into the Carpathian Basin during the 9th century AD. Their Homeland was probably in the southern Trans-Ural region, where the Kushnarenkovo culture disseminated. In the Cis-Ural region Lomovatovo and Nevolino cultures are archaeologically related to ancient Hungarians. In this study we describe maternal and paternal lineages of 36 individuals from these regions and nine Hungarian Conquest period individuals from today's Hungary, as well as shallow shotgun genome data from the Trans-Uralic Uyelgi cemetery. We point out the genetic continuity between the three chronological horizons of Uyelgi cemetery, which was a burial place of a rather endogamous population. Using phylogenetic and population genetic analyses we demonstrate the genetic connection between Trans-, Cis-Ural and the Carpathian Basin on various levels. The analyses of this new Uralic dataset fill a gap of population genetic research of Eurasia, and reshape the conclusions previously drawn from 10-11th century ancient mitogenomes and Y-chromosomes from Hungary.

...

Majority of Uyelgi males belonged to Y chromosome haplogroup N, and according to combined STR, SNP and Network analyses they belong to the same subclade within N-M46 (also known as N-tat and N1a1-M46 in ISOGG 14.255). N-M46 nowadays is a geographically widely distributed paternal lineage from East of Siberia to Scandinavia 33 . One of its subclades is N-Z1936 (also known as N3a4 and N1a1a1a1a2 in ISOGG 14.255), which is prominent among Uralic speaking populations, probably originated from the Ural region as well and mainly distributed from the West of Ural Mountains to Scandinavia (Finland). Seven samples of Uyelgi site most probably belong to N-Y24365 (also known as N-B545 and N1a1a1a1a2a1c2 in ISOGG 14.255) under N-Z1936, a specific subclade that can be found almost exclusively in todays’ Tatarstan, Bashkortostan and Hungary 17 (ISOGG, Yfull).

Csaky et al., Early Medieval Genetic Data from Ural Region Evaluated in the Light of Archaeological Evidence of Ancient Hungarians, bioRxiv, Posted July 13, 2020, doi: https://doi.org/10.1101/2020.07.13.200154

See also...

Hungarian Conquerors were rich in Y-haplogroup N

On the association between Uralic expansions and Y-haplogroup N

More on the association between Uralic expansions and Y-haplogroup N

Ancient DNA confirms the link between Y-haplogroup N and Uralic expansions

On the exotic origins of the Hungarian Arpad Dynasty (Nagy et al. 2020)

Hungarians speak a Uralic and Finno-Ugric language. However, the founders of the Medieval Hungarian state, the Arpad Dynasty, probably had Irano-Turkic paternal origins. There's a very interesting new paper on this topic at the European Journal of Human Genetics (see here). From the paper, emphasis is mine:

The phylogenetic origins of the Hungarians who occupied the Carpathian basin has been much contested [40]. Based on linguistic arguments it was proposed that they represented a predominantly Finno-Ugric speaking population while the oral and written tradition of the Árpád dynasty suggests a relationship with the Huns. Based on the genetic analysis of two members of the Árpád Dynasty, it appears that they derived from a lineage (R-Z2125) that is currently predominantly present among ethnic groups (Pashtun, Tadjik, Turkmen, Uzbek, and Bashkir) speaking Iranian or Turkic languages. However, their closest kin, the Bashkirs live in close proximity with Finno-Ugric speaking populations with the N-B539 haplogroup. A recent study shows that this haplogroup is also found in modern Hungarians [41]. Intriguingly, the most recent separation of the N-B539 derived lineages found in Hungarians and Bashkirs is estimated to have occurred ~2000 years before present [42]. This would suggest that a group of people consisting of a Turkic (R-SUR51) component and a Finno-Ugric (N-B539) component left the Volga Ural region about 2000 years ago and started a migration that eventually culminated in settlement in the Carpathian Basin.

Citation...

Nagy, P.L., Olasz, J., Neparáczki, E. et al. Determination of the phylogenetic origins of the Árpád Dynasty based on Y chromosome sequencing of Béla the Third. Eur J Hum Genet (2020). https://doi.org/10.1038/s41431-020-0683-z

See also...

Hungarian Conquerors were rich in Y-haplogroup N

On the association between Uralic expansions and Y-haplogroup N

More on the association between Uralic expansions and Y-haplogroup N

Ancient DNA confirms the link between Y-haplogroup N and Uralic expansions

Hungarian Conquerors were rich in Y-haplogroup N (Fóthi et al. 2020)

Open access at Archaeological and Anthropological Sciences at this LINK. Below is the paper abstract. Emphasis is mine:

According to historical sources, ancient Hungarians were made up of seven allied tribes and the fragmented tribes that split off from the Khazars, and they arrived from the Eastern European steppes to conquer the Carpathian Basin at the end of the ninth century AD. Differentiating between the tribes is not possible based on archaeology or history, because the Hungarian Conqueror artifacts show uniformity in attire, weaponry, and warcraft. We used Y-STR and SNP analyses on male Hungarian Conqueror remains to determine the genetic source, composition of tribes, and kin of ancient Hungarians. The 19 male individuals paternally belong to 16 independent haplotypes and 7 haplogroups (C2, G2a, I2, J1, N3a, R1a, and R1b). The presence of the N3a haplogroup is interesting because it rarely appears among modern Hungarians (unlike in other Finno-Ugric-speaking peoples) but was found in 37.5% of the Hungarian Conquerors. This suggests that a part of the ancient Hungarians was of Ugric descent and that a significant portion spoke Hungarian. We compared our results with public databases and discovered that the Hungarian Conquerors originated from three distant territories of the Eurasian steppes, where different ethnicities joined them: Lake Baikal-Altai Mountains (Huns/Turkic peoples), Western Siberia-Southern Urals (Finno-Ugric peoples), and the Black Sea-Northern Caucasus (Caucasian and Eastern European peoples). As such, the ancient Hungarians conquered their homeland as an alliance of tribes, and they were the genetic relatives of Asiatic Huns, Finno-Ugric peoples, Caucasian peoples, and Slavs from the Eastern European steppes.

Fóthi, E., Gonzalez, A., Fehér, T. et al., Genetic analysis of male Hungarian Conquerors: European and Asian paternal lineages of the conquering Hungarian tribes, Archaeol Anthropol Sci (2020) 12: 31. https://doi.org/10.1007/s12520-019-00996-0

See also...

On the association between Uralic expansions and Y-haplogroup N

More on the association between Uralic expansions and Y-haplogroup N

Ancient DNA confirms the link between Y-haplogroup N and Uralic expansions

The Balkan connection

The hot topic at the moment is social inequality in Bronze Age Europe, thanks to a new paper by Mittnik et al. at Science. The full article is sitting behind an exceedingly robust paywall here.

However, the genotype dataset from the paper is freely available at the Max Planck Society's Edmond data repository here. Below is my Principal Component Analysis (PCA) of ancient West Eurasian genetic variation featuring 41 of the highest quality ancients from the new dataset. Almost all of them are from the Lech Valley in the Bavarian Alps, covering the period from the Bell Beaker culture (BBC) to the Middle Bronze Age (MBA). Two of the samples are from a mass Corded Ware culture (CWC) burial in the more northerly Tauber Valley.

I've also highlighted other ancients on the plot associated with the BBC and CWC from present-day Netherlands and Germany, respectively. The relevant PCA datasheet can be downloaded here.

Social stratification in ancient Europe is a fascinating topic, and it's an issue that I've started looking at myself (see here). However, I can't see any correlation between the inferred social standing of the individuals from the Lech and Tauber valleys and their positions in my PCA.

Nevertheless, the PCA is interesting in that it highlights considerable genetic heterogeneity within the Lech Valley BBC population. Indeed, how is this heterogeneity even possible, if, as per Mittnik et al., ancient DNA "has shown that the spread of the BBC throughout continental Europe did not involve large-scale migrations"?

Below is another version of my PCA, but this time focusing on three males: Lech Valley Beakers UNTA58_68Sk1 and WEHR_1192SkA, as well as ALT_4 from the aforementioned mass CWC grave in the Tauber Valley. Note that UNTA58_68Sk1 and WEHR_1192SkA represent genetically the most southern and northern, respectively, Lech Valley BBC samples that had enough data to be run in my analysis. I chose to focus on males because they carry the Y-chromosome, which can be informative about male-mediated ancient population expansions.

The PCA outcomes for these individuals are generally in line with their results in other types of genetic analyses, including those based on formal statistics. For instance, compared to the other two, ALT_4 harbors excess early steppe herder ancestry, UNTA58_68Sk1 excess early European farmer ancestry, and WEHR_1192SkA excess European hunter-gatherer ancestry. Moreover...

- UNTA58_68Sk1 shows a non-local isotopic signature and belongs to Y-haplogroup G2a, a marker essentially missing from BBC populations north of the Alps, and is best modeled as a two-way mixture between Bronze Age populations from the Balkans and the Pontic-Caspian steppe (see here), which probably means that he was a migrant to the Lech Valley from south of the Alps

- importantly, UNTA58_68Sk1 is not an isolated case, at least in the sense that several other BBC individuals from Bavaria, Bohemia, Hungary and Poland show varying ratios of Balkan-related ancestry, although almost all of these people are women

- WEHR_1192SkA is very similar to Bell Beakers from the northern Netherlands with whom he shares the R1b-P312 Y-haplogroup, suggesting that he was part of a population that moved into the Lech Valley from potentially as far away as the North Sea coast

- although ALT_4 probably shares the R1b-L51 Y-haplogroup with WEHR_1192SkA and many other BBC and Bronze Age individuals from the Bavarian Alps and surrounds, this can't be used as evidence of significant local genetic continuity after the CWC period, especially considering the comparatively eastern genome-wide structure of ALT_4.

Of course, archeological data suggest that the BBC was influenced in some important ways by the Copper and Bronze Age cultures of the Balkans and Carpathian Basin. So much so, in fact, that Marija Gimbutas, author of The Civilization of the Goddess, believed that the BBC originated in the Balkans from a synthesis of the local Vucedol culture and the intrusive Yamnaya culture from the Pontic-Caspian steppe.

Considering the ancient DNA evidence, however, the main demographic center of the early BBC could not have been south of the Alps.

Rather, it appears that early BBC and even CWC groups from north of the Alps moved into the Balkans and Carpathian Basin, where they may have established contacts with the local elites. If so, this might explain the significant southern cultural influences on the BBC, but limited accompanying genetic impact. This scenario also has support from archeological data (for instance, see here).

See also...

Is Yamnaya overrated?

The Boscombe Bowmen

Single Grave > Bell Beakers

More on the association between Uralic expansions and Y-haplogroup N

Genes don't speak languages, people do. Thus, associations between genetic markers and languages may often not be easy to discern, especially with time. This is the case when it comes to Y-chromosome haplogroup N and the Hungarian language.

I briefly discussed this problem not long ago in the context of new ancient DNA samples from medieval Hungary (see here). Today, a detailed paper on the topic by Post et al. was published at Scientific Reports (open access here). It brings together evidence from modern and ancient DNA, linguistics and archeology to argue that Hungarian was introduced into the Carpathian Basin during the Middle Ages by migrants from near the Ural Mountains rich in Y-haplogroup N3a4-B539. Below is the paper abstract, emphasis is mine:

Hungarians who live in Central Europe today are one of the westernmost Uralic speakers. Despite of the proposed Volga-Ural/West Siberian roots of the Hungarian language, the present-day Hungarian gene pool is highly similar to that of the surrounding Indo-European speaking populations. However, a limited portion of specific Y-chromosomal lineages from haplogroup N, sometimes associated with the spread of Uralic languages, link modern Hungarians with populations living close to the Ural Mountain range on the border of Europe and Asia. Here we investigate the paternal genetic connection between these spatially separated populations. We reconstruct the phylogeny of N3a4-Z1936 clade by using 33 high-coverage Y-chromosomal sequences and estimate the coalescent times of its sub-clades. We genotype close to 5000 samples from 46 Eurasian populations to show the presence of N3a4-B539 lineages among Hungarians and in the populations from Ural Mountain region, including Ob-Ugric-speakers from West Siberia who are geographically distant but linguistically closest to Hungarians. This sub-clade splits from its sister-branch N3a4-B535, frequent today among Northeast European Uralic speakers, 4000–5000 ya, which is in the time-frame of the proposed divergence of Ugric languages.

Post et al., Y-chromosomal connection between Hungarians and geographically distant populations of the Ural Mountain region and West Siberia, Scientific Reports 9, Article number: 7786 (2019), DOI: https://doi.org/10.1038/s41598-019-44272-6

See also...

Hungarian Conquerors were rich in Y-haplogroup N (Fóthi et al. 2020)

On the association between Uralic expansions and Y-haplogroup N

Ancient DNA confirms the link between Y-haplogroup N and Uralic expansions

On the association between Uralic expansions and Y-haplogroup N

Almost all present-day populations speaking Uralic languages show moderate to high frequencies of Y-chromosome haplogroup N. I reckon there are two likely explanations for this:

- the speakers of Proto-Uralic were rich in N because they lived in an area, probably somewhere around the Ural Mountains, where it was common, and they spread it with them as they expanded from their homeland

- Uralic languages often came to be spoken in areas of North Eurasia where N was already found at moderate to high frequencies

The major exception to this rule are Hungarians, whose language belongs to the Ugric branch of Uralic. Their frequency of N is close to zero and they don't differ much in terms of overall genetic structure from their Indo-European-speaking neighbors in East Central Europe.

This is an issue that has generated much debate over the years about the nature of Uralic expansions, who the Hungarians really were, and how the Hungarian language came to be spoken in the heart of Europe.

But I never understood what the fuss was about, because based on historical sources alone it seemed rather obvious that Hungarian was introduced into the Carpathian Basin during the Middle Ages by a relatively small number of invaders from the east, probably from somewhere around the Ural Mountains, who imposed it on local Indo-European-speaking populations.

As far as I can remember, this has always been the academic consensus, and the results from one of the first ancient DNA studies of human remains soundly corroborated it. Back in 2008, Csányi et al. reported that two out of four skeletons from elite Hungarian conqueror graves dating to the 10th century carried the Tat C allele, which meant that they belonged to Y-haplogroup N (see here).

We've since had to wait over a decade to get a more comprehensive look at the Y-chromosome haplogroups of medieval Hungarians. The most useful effort to date, a manuscript courtesy of Neparáczki et al., was posted this week at bioRxiv (see here).

The results in the preprint suggest a much more complex picture than simply a migration of an obviously Uralic-speaking population rich in Y-haplogroup N into the medieval Carpathian Basin. But they do confirm the presence of N in Hungarian conqueror elites, and, in fact, of very specific subclades of N that link them to the present-day speakers of Uralic languages from around the Ural Mountains. Here are some pertinent quotes from the prepint:

Three Conqueror samples belonged to Hg N1a1a1a1a2-Z1936, the Finno-Permic N1a branch, being most frequent among northeastern European Saami, Finns, Karelians, as well as Komis, Volga Tatars and Bashkirs of the Volga-Ural region. Nevertheless this Hg is also present with lower frequency among Karanogays, Siberian Nenets, Khantys, Mansis, Dolgans, Nganasans, and Siberian Tatars 23.

...

It is generally accepted that the Hungarian language was brought to the Carpathian Basin by the Conquerors. Uralic speaking populations are characterized by a high frequency of Y-Hg N, which have often been interpreted as a genetic signal of shared ancestry. Indeed, recently a distinct shared ancestry component of likely Siberian origin was identified at the genomic level in these populations, modern Hungarians being a puzzling exception 36. The Conqueror elite had a significant proportion of N Hgs, 7% of them carrying N1a1a1a1a4-M2118 and 10% N1a1a1a1a2-Z1936, both of which are present in Ugric speaking Khantys and Mansis 23.

...

Population genetic data rather position the Conqueror elite among Turkic groups, Bashkirs and Volga Tatars, in agreement with contemporary historical accounts which denominated the Conquerors as “Turks” 38. This does not exclude the possibility that the Hungarian language could also have been present in the obviously very heterogeneous, probably multiethnic Conqueror tribal alliance.

Indeed, a large proportion of the 44 males from elite Hun, Avar and Hungarian Conqueror burials analyzed in the study belonged to Y-haplogroups that can't be plausibly associated with the earliest Uralic speakers, but rather with those of various Indo-European languages, such as I1 and R1b-U106 (these are Germanic-specific markers), I2a-L621 and R1a-CTS1211 (obviously Slavic) and R1a-Z2124 (largely Eastern Iranian).

If most of these results aren't due to contamination, then it's likely that both the early Hungarian commoners and elites were, by and large, derived from Indo-European-speaking populations. No wonder then, that present-day Hungarians are basically indistinguishable genetically from their Indo-European-speaking neighbors and, like them, show hardly any Y-haplogroup N.

See also...

Hungarian Conquerors were rich in Y-haplogroup N (Fóthi et al. 2020)

More on the association between Uralic expansions and Y-haplogroup N

Ancient DNA confirms the link between Y-haplogroup N and Uralic expansions