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Slowly, but surely, the diversity of genome-wide SNP samples available to the public is increasing. One of the most interesting and valuable sets, released courtesy of the recently published Yunusbayev et al. study, comes from the Mordovian Republic, in the Russian Federation. It includes the results for 15 people from the closely related Erzya and Moksha ethnic groups (10 and 4 samples respectively, and one mixed individual).

So why are these Mordovian Republic samples so important? Because the Erzya and Moksha have a long history in the Middle Volga region of Russia, which is a key zone for those interested in the histories of the Eurasian steppe and forest steppe. Their presence in the area as a distinct Volgaic Finnic entity probably dates back to around 500 BC, but various groups who contributed to their ethnogenesis were likely here much earlier. We can infer this from the Erzya and Moksha languages. They belong to the Finno-Ugric (and thus Uralic) family of tongues, but show ancient influences from Indo-Iranian, Baltic, and possibly even proto-Tocharian. What this suggests is that their genomes should provide strong clues to the various population movements through this region, possibly all the way from the metal ages to historic times.

Based on a variety of analyses I've carried out with these samples to date, this certainly appears to be the case. But the results give a strong impression of mostly one way traffic across Mordovia - from the west to east. Below is an MDS featuring the Erzya and Moksha from Yunusbayev et al., plus a single Erzya from my Eurogenes Ancestry Project (Erzya1), against samples from the North Caucasus, Chuvashia and North and East Europe. It shows clearly where they stand in terms of biogeographic affinities. Not only do they cluster tightly with the North and East Europeans (PL = Poland, RU = Russia, SE = Sweden, UA = Ukraine), but they're very clearly separated from the North Caucasus samples, and even their immediate neighbors on the Volga, the Chuvash.

These results are obviously totally out of whack with geography. In fact, it appears that the Swedes are a little closer to the North Caucasus than the Erzya and Moksha are across the more significant dimension 1. This might seem bizarre at first, but not if we consider that there weren't any significant post-Neolithic migrations from the south (ie. from West Asia/Middle East) directly to the Middle Volga region. It's not just my own work showing this, but also the aforementioned Yunusbayev et al. study, which focused on the apparent lack of gene flow between the Caucasus and Eastern Europe. And it's what some old school archeologists have been saying for years.

From the Eneolithic [Copper] period to the Early Iron Age there were no invasions of population from the Near East or Iran in the steppes of Eurasia. All migrations, though numerous, occurred within the steppe and forest-steppe zones.

Elena E. Kuz'mina, James P. Mallory (Editor), The Origin of the Indo-Iranians, Vol. 3, Barnes & Noble, 2007

But aren't Scandinavians pure Europeans from way back, you might ask? No they're not. In fact, my data and ancient DNA results suggest that Europe, including Northern Europe, was overwhelmingly populated from the south and southeast, way post Ice Age, and Eastern Europe mostly from Central Europe. Hence, the cline of West Asian/Middle Eastern influence across the continent generally runs like this: Southern/Southeastern Europe > Western Europe > North/Central Europe > Eastern Europe. And this is why Swedes have higher West Asian/Middle Eastern affinity than the Erzya and Moksha. In fact, groups like the French from Western Europe proper, have significantly more West Asian/Middle Eastern influence than the two Mordovian groups. This was illustrated with a PCA by Yunusbayev et al. (see here).

Obviously, the Erzya and Moksha do carry influence from the east of the Volga. This shows up on the MDS plot along dimension 2, with the samples from Mordovia placed slightly closer to the Chuvash than most of the other North and Eastern Europeans. My ADMIXTURE analyses also indicate about 6% of North/East Asian ancestry, which is 5-6% higher than in most North/Central European groups (see here).

This isn't surprising, considering the number of invasions from the east that have passed through or near the Middle Volga on their way to Europe, including various Turkic and Altaic onslaughts during the middle ages. What is startling, however, is how much of the older, western-like genetic ancestry the Erzya and Moksha have manged to retain. This didn't happen across the border in what is now Chuvashia, where language replacement to Turkic seems to have been accompanied by significant gene flow from Siberia. Thus, modern Chuvashs carry around 20-25% North/East Asian admixture.

Another interesting fact revealed by the Yunusbayev et al. dataset was the high level of light pigmentation amongst the Erzya and Moksha. I discovered this when plotting allele frequencies from various SNPs linked to fair hair and light eyes in Europeans (see here). The combined samples from Mordovia came out as one of the fairest groups. The result was confirmed by a reader, who sent me data from a pigmentation survey of Northeastern Europe, showing the Erzya as fairer than ethnic Russians, and the Moksha at about the same level as the Russians. A quick search via Google revealed the following two random representatives of the Erzya nation. I don't know the original source for the first image, while the second comes from Wikipedia.

I'm not sure whether those two look like the average Erzya, or even if they're representative of a significant minority of the population. The latter is more likely. In any case, it's interesting that two random hits off Google backed up the genetic analyses of the reference samples from Mordovia. The relatively light pigmentation of such individuals, from near the Samara Bend, certainly fits with the notion of an expansion of a predominantly light eyed and fair haired population from this area during the Bronze and Iron Ages. Of course, I'm referring here to the Andronovo tribes.

According to C-14 [carbon] dating, the Andronovo and Timber-grave cultures formed at the shift of the 3rd and 2nd millennium BC on the common base of Eastern European cultures. The main components were the steppe cultures of Poltavka, late Catacomb (to far lesser degree), and the Abashevo culture of the forest steppe zone. The latter culture belonged to the Corded Ware group of cultures of Central Europe.


The subsequent development of the Timber-grave culture in the Pontic Caspian steppes was determined by the large-scale participation of the Abashevo culture in the Don region and the post-Catacomb Multi-roller Ware culture in the Ukraine.


The close genetic affinity of the Andronovo and the Timber-grave culture tribes became even closer thanks to regular contacts, especially because of the flow of tin from Kazakhstan and of copper ( as well as of finished products) from the Urals. The contact zone lay between the Volga and the Urals. The Andronovan influence reached the Dnieper.

Elena E. Kuz'mina, James P. Mallory (Editor), The Origin of the Indo-Iranians, Vol. 3, Barnes & Noble, 2007

Our autosomal, Y-chromosomal and mitochondrial DNA analyses reveal that whereas few specimens seem to be related matrilineally or patrilineally, nearly all subjects belong to haplogroup R1a1-M17 which is thought to mark the eastward migration of the early Indo-Europeans. Our results also confirm that at the Bronze and Iron Ages, south Siberia was a region of overwhelmingly predominant European settlement, suggesting an eastward migration of Kurgan people across the Russo-Kazakh steppe. Finally, our data indicate that at the Bronze and Iron Age timeframe, south Siberians were blue (or green)-eyed, fair-skinned and light-haired people and that they might have played a role in the early development of the Tarim Basin civilization.

Christine Keyser et al.,
Ancient DNA provides new insights into the history of south Siberia..., Human Genetics, Saturday, May 16, 2009, doi: 10.1007/s00439-009-0683-0

The publically available datasets I used in this blog entry are found at the links below. My analysis can be reproduced with the PLINK software, found here.

HGDP-CEPH Human Genome Diversity Cell Line Panel

Yunusbayev et al. dataset

By the way, I'm interested in the types of R1a1a1 carried by Erzya and Moksha individuals. For instance, are they showing R1a1a1i (Z280+), R1a1a1g (M458+), or perhaps R1a1a1h (Z93+)?

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Ответы на эту тему форума

жалко английский не знаю((

Поэтому дал ссылку на мои комментарии.

Забавно, что шведы ближе к эрзе, чем соседи чуваши.

хм) финно-угры и скандинавы, это уже круто.

пургине, вы проводите анализы по днк?

Purgine said:

Забавно, что шведы ближе к эрзе, чем соседи чуваши.

Если хотите сделать анализ, то пишите в ЛС. Эрзянам и мокшанам советую вступить в Erzya-Moksha DNA Project



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