Hungarian mitogenomes similar to East and West Slavs, but genetic substratum predates their historic contacts

middle-age-hungarian

Whole mitochondrial genome diversity in two Hungarian populations, Malyarchuk et al. Mol Genet Genomics (2018).

Abstract:

Complete mitochondrial genomics is an effective tool for studying the demographic history of human populations, but there is still a deficit of mitogenomic data in European populations. In this paper, we present results of study of variability of 80 complete mitochondrial genomes in two Hungarian populations from eastern part of Hungary (Szeged and Debrecen areas). The genetic diversity of Hungarian mitogenomes is remarkably high, reaching 99.9% in a combined sample. According to the analysis of molecular variance (AMOVA), European populations showed a low, but statistically significant level of between-population differentiation (Fst = 0.61%, p = 0), and two Hungarian populations demonstrate lack of between-population differences. Phylogeographic analysis allowed us to identify 71 different mtDNA sub-clades in Hungarians, sixteen of which are novel. Analysis of ancestry-informative mtDNA sub-clades revealed a complex genetic structure associated with the genetic impact of populations from different parts of Eurasia, though the contribution from European populations is the most pronounced. At least 8% of ancestry-informative haplotypes found in Hungarians demonstrate similarity with East and West Slavic populations (sub-clades H1c23a, H2a1c1, J2b1a6, T2b25a1, U4a2e, K1c1j, and I1a1c), while the influence of Siberian populations is not so noticeable (sub-clades A12a, C4a1a, and probably U4b1a4).

Interesting excerpt:

Our analysis of ancestry-informative mtDNA sub-clades revealed a complex genetic structure associated with the genetic impact of populations from different parts of Europe. At least 8% of ancestry-informative haplotypes found in Hungarians demonstrate similarity with East (Russians and Ukrainians) and West (Poles and Slovaks) Slavic populations (sub-clades H1c23a, H2a1c1, J2b1a6, T2b25a1, U4a2e, K1c1j, and I1a1c). This observation is consistent with the results of mtDNA studies of medieval populations living in the Hungarian-Slavic contact zone of the Carpathian Basin in the 9th–12th centuries AD (Csákyová et al. 2016). Taken together, these data confirm earlier historical and archaeological reports on mixed populations of medieval Slavs and Magyars, based on the research into cemeteries discovered in Central Europe (Csősz et al. 2016; Csákyová et al. 2016). On the other hand, we cannot confirm the Hungarian-Slavic contacts using molecular dating of the identified mtDNA sub-clades, since their age exceeds the estimated time of the contact period and varies from 1.3 kya (for K1c1j) to 5.2 kya (for T2b25a1) (Figure S1). One of an issue may be sample size problem, because some haplotypes may be missed in the sampling, and this can lead to an overestimate of the age of the mtDNA sub-clade (Richards et al. 2000).

hungarian-mtdna-haplogroup-j
Figure S1. MDS plot based on Fst values calculated from complete mtDNA sequences for population samples from Europe. Stress value = 0.00078

However, it is known that the evolutionary ages of most mtDNA lineages specific to Eastern and Central Europeans correspond to approximately 4 kya (from 2.3 to 5.9 kya) (Malyarchuk et al. 2008, 2017; Mielnik-Sikorska et al. 2013; Översti et al. 2017), thus coinciding with the time of the Bronze Age expansion of Eastern Europeans in accordance with the Kurgan model established by archaeologists and paleogeneticists (Gimbutas 1971; Allentoft et al. 2015; Haak et al. 2015). Thus, similar haplotypes among Hungarians and Slavs and other European ethnic groups can be a reflection of the common genetic substratum which predates the formation of the most modern European populations. Therefore, mtDNA sub-clades H5a1m, T2a1c, and W3a1d1 (with the ages varying from 2.6 to 3.9 kya, based on complete mtDNA mutation rate), which are shared by Hungarians and Finno-Ugric peoples, such as Estonians and Finns, may testify these pan-European relationships (Figure S1). Another example is the sub-clade J2b1a6, which unites the mtDNA haplotypes of the ancient and modern population of Eastern and Central Europe from the Iron Age to the present (Figure S1).

Related:

First Hungarian ruling dynasty, the Árpáds, of Y-DNA haplogroup R1a

middle-ages-europe

Open access article DNA profiling of Hungarian King Béla III and other skeletal remains originating from the Royal Basilica of Székesfehérvár, Olasz, J., Seidenberg, V., Hummel, S. et al. Archaeol Anthropol Sci (2018).

Abstract

A few decades after the collapse of the Avar Khaganate (c. 822 AD), Hungarian invaders conquered the Carpathian Basin (c. 862–895 AD). The first Hungarian ruling dynasty, the Árpáds played an important role in European history during the Middle Ages. King Béla III (1172–1196) was one of the most significant rulers of the dynasty. He also consolidated Hungarian dominance over the Northern Balkans. The provostry church of the Virgin Mary (commonly known as the Royal Basilica of Székesfehérvár) played a prominent role as a coronation church and burial place of medieval Hungarian kings. The basilica’s building and graves had been destroyed over the centuries. The only royal graves that remained intact were those of King Béla III and his first spouse, Anna of Antioch. These graves were discovered in 1848. We defined the autosomal STR (short tandem repeat) fingerprints of the royal couple and eight additional individuals (two females and six males) found in the Royal Basilica. These results revealed no evidence of first-degree relationship between any of the investigated individuals. Y-chromosomal STR profiles were also established for all the male skeletons. Based upon the Y-chromosomal data, one male skeleton showed an obvious patrilineal relationship to King Béla III. A database search uncovered an existing Y-chromosomal haplotype, which had a single-repeat difference compared to that of King Béla. It was discovered in a person living in an area close to Hungary. This current male line is probably related paternally to the Árpád Dynasty. The control region of the mitochondrial DNA was determined in the royal couple and in the remains of the inferred relative. The mitochondrial results excluded sibling relationship between the King and the patrilineal relative. In summary, we successfully defined a Y-chromosomal profile of King Béla III, which can serve as a reference for the identification of further remains and disputed living descendants of the Árpád Dynasty. Among the examined skeletons, we discovered an Árpád member, whose exact affiliation, however, has not yet been established.

The Árpad Dynasty

The Árpád Dynasty (c. 850–1301 AD) played an important role in European history during the Middle Ages (Hóman 1940-1943). The first Great Prince Álmos organised the monarchic state in the northern region of the Black Sea c. 850. A few decades after the collapse of the Avar Khaganate (c. 822 AD), Álmos and his son Árpád conquered the Carpathian Basin (c. 862–895 AD) (Szőke 2014). During the conquest, Hungarian invaders, together with Turkic-speaking Kabars assimilated the Avars and Slavonic groups (Szádeczky-Kardoss 1990). Thus, most of the population in the Carpathian Basin originated from the Hun-Turkic cultural community of the Eurasian Steppe and was accompanied by Slavonic and German-speaking groups (László 1996). The origin of Hungarians is still controversial, and this paper cannot cover this complex subject. The Hungarian Great Principality represented the Eurasian steppe empires in Central Europe from c. 862 until 1000. Saint Stephen I, the last Great Prince (997–1000) and first King (1000–1038) of Hungary re-organised this early Hungarian state as a Christian kingdom. Saint Stephen received the royal crown from the Pope and joined the post-Roman Christian political system and cultural commonwealth of Latin Europe (Pohl 2003; Szabados 2011). Hungary remained an independent state between the German and Byzantine empires (Makk 1989). King Béla III (1172–1196) was one of the most significant rulers of the dynasty. He was the second son of King Géza II (1141–1162) and Queen Euphrosyne, the daughter of Mstislav I (1125–1132), the Great Prince of Kiev. Through the mediation of Byzantine Emperor Manuel I Komnenos, Béla married Anna of Châtillon from Antioch (1150–1184), the half-sister of the Emperor’s wife in 1170. After Manuel’s death, King Béla consolidated Hungarian dominance over the Northern Balkans.

The provostry church of the Virgin Mary (commonly known as the Royal Basilica of Székesfehérvár) was built by Saint Stephen I at the beginning of the eleventh century. The basilica played a prominent role as a church of coronation and as the main burial place of Hungarian kings in the Middle Ages. Fifteen kings, several queens, princes and princesses and clerical and secular dignitaries were buried there over five centuries (Engel 1987)

king-bela-iii
The five graves excavated by János Érdy. Drawn by János Varsányi (1848). Originally published by Érdy (1853). I: remains of Béla III; II: remains of Anna of Antioch; III: a male skeleton whose identity with II/52 is questioned; IV: the skeleton of an expectant female, only foetal bones remained; V: a crushed skeleton, it has not been preserved.

Discussion

There were three R1a and two R1b statistically predicted Y haplogroups among the male skeletons (Table 3). These are the most frequent and second most frequent haplogroups (25.6 and 18.1% respectively) in the present Hungarian population (Völgyi et al. 2009). King Béla III was inferred to belong to haplogroup R1a. The R1a Y haplogroup relates paternally to more than 10% of men in a wide geographic area from South Asia to Central Eastern Europe and South Siberia (Underhill et al. 2010). It is the most frequent haplogroup in various populations speaking Slavic, Indo-Iranian, Dravidian, Turkic and Finno-Ugric languages (Underhill et al. 2010).

Kinship analysis

The autosomal STR results contradicted the paternity between King Béla III and II/52. The mitochondrial sequence results excluded siblingship, too. Apart from that, we also tested the hypothesis for siblingship versus non-relationship based on the autosomal STR results using “Familias 3”. The LR (likelihood ratio) for the alternative hypothesis was found to be 7.67, which was inconclusive. Testing the hypothesis for a grandfather-grandson (or uncle-nephew) relationship versus non-relationship resulted in an LR of 5.44, which corresponds to a probability of 84.46% (assuming a prior probability of 50%). This result is indecisive for the hypothesis.

Honfoglalas
The Hungarian conquest of the Carpathian Basin, by Fz22 at Wikipedia.

So, the first Hungarian dynasty, which one can safely say were one of the ruling clans among Hungarian conquerors, a group of Ugric speakers that invaded the Carpathian basin from the steppe in the 9th c. (stemming originally from North-Eastern Europe) were of R1a lineages.

Who could have thought, right?

Related:

Admixture of Srubna and Huns in Hungarian conquerors

hungarian-conqueror-migrations

New preprint at BioRxiv, Mitogenomic data indicate admixture components of Asian Hun and Srubnaya origin in the Hungarian Conquerors, by Neparáczki et al. (2018), at BioRxiv.

Abstract (emphasis mine):

It has been widely accepted that the Finno-Ugric Hungarian language, originated from proto Uralic people, was brought into the Carpathian Basin by the Hungarian Conquerors. From the middle of the 19th century this view prevailed against the deep-rooted Hungarian Hun tradition, maintained in folk memory as well as in Hungarian and foreign written medieval sources, which claimed that Hungarians were kinsfolk of the Huns. In order to shed light on the genetic origin of the Conquerors we sequenced 102 mitogenomes from early Conqueror cemeteries and compared them to sequences of all available databases. We applied novel population genetic algorithms, named Shared Haplogroup Distance and MITOMIX, to reveal past admixture of maternal lineages. Phylogenetic and population genetic analysis indicated that more than one third of the Conqueror maternal lineages were derived from Central-Inner Asia and their most probable ultimate sources were the Asian Huns. The rest of the lineages most likely originated from the Bronze Age Potapovka-Poltavka-Srubnaya cultures of the Pontic-Caspian steppe, which area was part of the later European Hun empire. Our data give support to the Hungarian Hun tradition and provides indirect evidence for the genetic connection between Asian and European Huns. Available data imply that the Conquerors did not have a major contribution to the gene pool of the Carpathian Basin, raising doubts about the Conqueror origin of Hungarian language.

hungarian-conqueror-mtdna
“Comparison of major Hg distributions from modern and ancient populations. Asian main Hg-s are designated with brackets. Major Hg distribution of Conqueror samples from this study are very similar to that of other 91 Conquerors taken from previous studies [11,12]. Scythians and ancient Xiongnus show similar Hg composition to the bracketed Asian fraction of the Conqueror samples, but Hg B is present just in Xiongnus. Modern Hungarians have very small Asian components pointing at small contribution from the Conquerors. Of the 289 modern Hungarian mitogenomes 272 are published in [29]. Scythian Hg-s are from [48,49,55,59,71–74]. Xiongnu Hg-s are from [66–69].”

Just recently another article contributed to a similar idea. I already talked about the Bronze Age R1a-z93 sample with high steppe ancestry found in the Balkans, and its likely origin in an expansion of the Srubna or a related culture. No truce, therefore, for those looking for autochthonous continuity anywhere in Europe.

We are seeing how multiple migrations shaped the history of the Carpathian basin (and its complex genetic structure) – and of Europe in general -, often from the Pontic-Caspian steppe. That is clear from many different prehistorical and historical times, such as the expansions of Suvorovo-Novodanilovka, Yamna, Srubna, Thraco-Cimmerians, Sarmatians, Scythians, Huns,…

About the linguistic interpretations based on genetics contained in the paper (Hungarian language as a legacy of Huns), well, you know my stance regarding the Yamnaya ancestral concept (and the wrong linguistic interpretations derived from it, which many sadly keep to this day), and genetics in general to solve language questions

This is yet another example of how (what some people would call) “scientific data” is useless without sound anthropological models.

Featured image, from the article: “Hypothetic origin and migration route of different components of the Hungarian Conquerors. Bluish line frames the Eurasian steppe zone, within which all presumptive ancestors of the Conquerors were found. Yellow area designates the Xiongnu Empire at its zenith from which area the East Eurasian lineages originated. Phylogeographical distribution of modern East Eurasian sequence matches (Fig. 1) well correspond to this territory, especially considering that Yakuts, Evenks and Evens lived more south in the past [108], and European Tatars also originated from this area. Regions where Asian and European Scythian remains were found are labeled green, pink is the presumptive range of the Srubnaya culture. Migrants of Xiongnu origin most likely incorporated descendants of these groups. The map was created using QGIS 2.18.4[109]”.

Article available under a CC-BY-NC-ND 4.0 International license.

Discovered via Razib Khan.

See also:

Modern Hungarian mtDNA more similar to ancient Europeans than to Hungarian conquerors

middle-ages-europe

New preprint at BioRxiv, MITOMIX, an Algorithm to Reconstruct Population Admixture Histories Indicates Ancient European Ancestry of Modern Hungarians, by Maroti et al. (2018).

hungarian-shared-mtDNA
The estimated age distribution of the shared mt Hgs between Hungarians (Hun), the best hypothetical admix (mixFreq) and the populations contributing to this admix: Belgian/Dutch (BeN), Danish (Dan), Basque (Bsq), Croatian/Serbian (CrS), Baltic Late Bronze Age culture (BalBA), Bell Beaker culture (BellB), Slovakian (Slo). The numbers in parentheses indicate the contributions to the best hypothetical admix.

Abstract (emphasis mine)

By making use of the increasing number of available mitogenomes we propose a novel population genetic distance metric, named Shared Haplogroup Distance (SHD). Unlike FST, SHD is a true mathematical distance that complies with all metric axioms, which enables our new algorithm (MITOMIX) to detect population-level admixture based on SHD minimum optimization. In order to demonstrate the effectiveness of our methodology we analyzed the relation of 62 modern and 25 ancient Eurasian human populations, and compared our results with the most widely used FST calculation. We also sequenced and performed an in-depth analysis of 272 modern Hungarian mtDNA genomes to shed light on the genetic composition of modern Hungarians. MITOMIX analysis showed that in general admixture occurred between neighboring populations, but in some cases it also indicated admixture with migrating populations. SHD and MITOMIX analysis comply with known genetic data and shows that in case of closely related and/or admixing populations, SHD gives more realistic results and provides better resolution than FST. Our results suggest that the majority of modern Hungarian maternal lineages have Late Neolith/Bronze Age European origins (partially shared also with modern Danish, Belgian/Dutch and Basque populations), and a smaller fraction originates from surrounding (Serbian, Croatian, Slovakian, Romanian) populations. However only a minor genetic contribution (<3%) was identified from the IXth Hungarian Conquerors whom are deemed to have brought Hungarians to the Carpathian Basin. Our analysis shows that SHD and MITOMIX can augment previous methods by providing novel insights into past population processes.

hungarian-hierarchic-cluster
Unrooted hierarchic cluster of modern and archaic populations based on the SHD matrix.

It is interesting to keep receiving data as to how language does not correlate well with Genomics, whether admixture or haplogroups, even though it is already known to happen in regions such as Anatolia, the Baltic, South-Eastern or Northern Europe.

Thorough anthropological models of migration or cultural diffusion are necessary for a proper interpretation of genetic data. There is no shortcut to that.

hungarian-mtdna
Co-occurrence of Hungarian Bronze Age mt Hgs Distribution of mt Hgs found in Hungarian Bronze Age archaic samples in the analyzed populations. The fixation dates are based on Behar et al [6].

Images made available under a CC-BY-NC-ND 4.0 International license.
See also:

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