The Yamnaya Steppe Herders from Russia

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The Yamnaya Steppe Herders from Russia Jul 15, 2020 6:53:47 GMT

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Post by Admin on Jul 15, 2020 6:53:47 GMT

Early farmer ancestry in Fatyanovo Culture individuals
Then, we turned to the Bronze Age Fatyanovo Culture individuals and determined that their maternal
(subclades of mtDNA hg U5, U4, U2e, H, T, W, J, K, I and N1a) and paternal (chrY hg R1a-M417)
lineages (Table 1, Supplementary Fig. 5, Supplementary Data 2) were ones characteristic of CWC
individuals elsewhere in Europe13,14,29,32,28. Interestingly, in all individuals for which the chrY hg could
be determined with more depth (n=6), it was R1a2-Z93 (Table 1, Supplementary Data 2), a lineage now
spread in Central and South Asia, rather than the R1a1-Z283 lineage that is common in Europe38,39.

On the PCA, the Fatyanovo individuals (and the Estonian CWC individual) group together with many
European Late Neolithic/Bronze Age (LNBA) and Steppe Middle/Late Bronze Age (MLBA)
individuals on top of modern Northern and Eastern Europeans (Fig. 2A). This ancient cluster is shifted
towards Anatolian and European EF compared to Steppe Early/Middle Bronze Age (EMBA)
populations, including the Yamnaya. The same could be seen in ADMIXTURE analysis where the
Fatyanovo individuals are most similar to LNBA Steppe ancestry populations from Central Europe,
Scandinavia and the Eastern Baltic (Fig. 2B, Supplementary Fig. 1). These populations are composed
of the blue "WHG" and yellow "Khanty" component and two brown components maximized in HG
from the Caucasus and Iran, similarly to Yamnaya populations. However, the European LNBA
populations (including Fatyanovo) also display a green component most frequent in Anatolian and
European EF populations, which is not present in the Yamnaya from Russia.

Figure 2. Principal component and ADMIXTURE analyses’ results.

We compared the affinities of the Fatyanovo individuals to those of other populations using f3 and D
statistics and found that Fatyanovo shares more with European EF populations and modern Near
Easterners than Yamnaya_Samara does (Fig. 3C, Supplementary Fig. 2C, Supplementary Fig. 3C).
Importantly, this signal can also be seen when using either autosomal or X chromosome positions from
the Lazaridis et al. 201631 ancient dataset instead of the autosomal positions of the EBC-chipDB
(Supplementary Fig. 3A–B). However, when comparing Fatyanovo to Central_CWC, there were no
clear differences in their affinities to different ancient or modern population groups (Fig. 3D,
Supplementary Fig. 2D). Furthermore, we confirmed the presence of sex-biased admixture previously
seen in CWC individuals from Estonia, Poland and Germany40,41,32,42,43 in the Fatyanovo

(Supplementary Fig. 3D). This is also supported by the presence of mtDNA hg N1a in two Fatyanovo
individuals – a hg frequent in Linear Pottery Culture (LBK) early farmers, but not found in Yamnaya
individuals44,13,29.

Figure 3. Outgroup f3 statistics' results of comparisons with ancient populations.

Since the previous analyses suggested that the genetic makeup of the Fatyanovo Culture individuals
was a result of admixture between migrating Yamnaya individuals and contemporary European
populations, we used two complementary methods (qpAdm and ChromoPainter/NNLS) to determine
the most suitable proxies for the admixing populations and the mixing proportions. The model with the
highest p-value and the lowest standard errors for qpAdm had Yamnaya and Levant Neolithic (N) as
sources while the model with the smallest residuals for ChromoPainter/NNLS included also WHG, so
we are presenting both models for both analyses (Fig. 4A–B). We ran the analyses for Fatyanovo,
Central CWC and Baltic CWC and saw that in the results of the more complex model, some of the
Yamnaya ancestry from the simpler model got reassigned as WHG but the proportions of Levant N
ancestry remained similar (within 4%). We found that although the results from qpAdm and
ChromoPainter/NNLS differed somewhat, both showed more Levant N ancestry in Fatyanovo (~22%
and ~17%) than in Central/Baltic CWC (~11% and ~10%). What is more, this result is supported by
models where Fatyanovo and Central/Baltic CWC are modeled as a mixture of Levant N and
Baltic/Central CWC, respectively (Supplementary Fig. 4A).

Lastly, we looked for closely related individuals in the Fatyanovo Culture sample-set using READ45.
There were no confirmed cases of 2nd degree or closer relatives (Supplementary Fig. 4B).

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The Yamnaya Steppe Herders from Russia Jul 15, 2020 21:35:17 GMT

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Phenotype informative allele frequency changes in Western Russia
We imputed the genotypes of 115 phenotype-informative positions connected to diet (carbohydrate,
lipid and vitamin metabolism), immunity (response to pathogens, autoimmune and other diseases) and
pigmentation (eye, hair and skin). We used previously published Eastern Baltic individuals for
comparison27,32,33. Here, we focus on variants associated with pigmentation (39 SNPs of the HIrisPlex-
S system), lactase persistence (rs4988235, rs182549; MCM6) and fatty acid metabolism (rs174570C;
FADS2–3) (Table 2), the latter three alleles showing a significant increase in frequency through time.
Although the results should be interpreted with due caution because of the small sample size, we inferred
that the examined WeRuHG individuals carried alleles connected to brown eyes, dark brown to black
hair and intermediate or dark skin pigmentation while around a third of the Fatyanovo individuals had
blue eyes and/or blond hair.

Furthermore, we infer that the frequency of the two alleles associated with
lactase persistence (rs4988235, rs182549; MCM6) is 0% in WeRuHG and ~10% in Fatyanovo Culture
individuals (similar to Eastern Baltic populations from the same time periods), but has a significant
increase to over 40% by the Late Bronze Age in the Eastern Baltic33. On the other hand, an allele
connected to an increase in cholesterol and a decrease in triglyceride levels in serum (rs174570C;
FADS2–3) significantly rises in frequency by the CWC period – from ~20% in Eastern Baltic and
Western Russian HG to ~50% in CWC individuals from the two regions. Interestingly, the frequency
of the allele increases further in the subsequent time periods in the Eastern Baltic.

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The Yamnaya Steppe Herders from Russia Jul 16, 2020 20:14:21 GMT

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Post by Admin on Jul 16, 2020 20:14:21 GMT

Discussion
After the Last Glacial Period, at the end of the 10th and the beginning of the 9th millennium BC, vast
areas in the Eastern Baltic, Finland and northern parts of Russia were populated relatively quickly by
hunter-gatherer groups46–49. Flint originating from several places in the Eastern Baltic and the European
part of Russia and the similarities in lithic and bone technologies and artefacts suggest the existence of
extensive social networks in the forest belt of Eastern and Northern Europe after it was populated50,51,48.
This has led to the hypothesis that the descendants of Paleolithic HG from Eastern Poland to the central
areas of European Russia took part in the process52,7,53 and remained connected to their origin, creating
a somewhat stretched social network8. However, these connections ceased after a few centuries, as can
be witnessed from the production of stone tools from mostly local materials, and new geographically
smaller social units appeared in the middle of the 9th millennium BC8. Ancient DNA studies of this
area have included Mesolithic individuals that are from the 8th millennium BC or younger and reveal
two genetic groups: WHG in the Eastern Baltic and EHG in Northwestern Russia14,29,27,32,28,30. However,
no human genomes from the settlement period have been published so far, leaving the genetic
ancestry/ancestries of the settlers up for discussion. The individual PES001 from around 10,700 cal BC
(probably somewhat affected by the freshwater reservoir effect) presented here with 4x coverage
provides evidence for EHG ancestry in Northwestern Russia close to the time it was populated. This, in
turn, raises the question of the ancestry of the first people of the Eastern Baltic, keeping in mind the
shared social network of the two areas at the time of settlement next to the presence of genetically
different groups later in time – a question hopefully answered by future studies.

The formation of Fatyanovo Culture is one of the main factors that affected the population, culture and
lifestyle of the previously hunter/fisher-gatherer culture of the Eastern European forest belt. The
Fatyanovo Culture people were the first farmers in the area and the arrival of the culture has been
associated with migration17,21. This is supported by our results as the Stone Age HG and the Bronze
Age Fatyanovo individuals are genetically clearly distinguishable. The sample size of our HG is
admittedly low, but the three individuals form a genetically homogenous group with previously reported
EHG individuals23–28 and the newly reported PES001 is the highest coverage whole-genome sequenced
EHG individual so far, providing a valuable resource for future studies. What is more, the Fatyanovo
Culture individuals (similarly to other CWC people) have mostly Steppe ancestry, but also some EF
ancestry which was not present in the area before and thus excludes the northward migration of
Yamnaya Culture people with only Steppe ancestry as the source of Fatyanovo Culture population. The
strongest connections for Fatyanovo Culture in archaeological material can be seen with the Middle
Dnieper Culture20,54 spread in present-day Belarus and Ukraine55,56. Importantly, the territory of what
is now Ukraine is where the most eastern individuals with European EF ancestry and the most western
Yamnaya Culture individuals are from based on published genomic data26,57 (Fig. 1, Supplementary
Data 1). Furthermore, archaeological finds show that LBK reached Western Ukraine around 5,300 BC58
and the Yamnaya Culture (burial mounds) arrived in Southeastern Europe around 3,000 BC and spread
further as far as Romania, Bulgaria, Serbia and Hungary59. These findings suggest present-day Ukraine
as the possible origin of the migration leading to the formation of the Fatyanovo Culture and of the
Corded Ware cultures in general.

The exact timing of and processes involved in the emergence of the Fatyanovo Culture in European
Russia and the local processes following it have also remained unclear. Until recently, the Fatyanovo
Culture was thought to have developed later than other CWC groups and over a longer period of
time17,20. However, radiocarbon dates published last year21 and the 25 new dates presented here point
towards a fast process, similar in time to CWC people reaching the Eastern Baltic and southern
Fennoscandia60–62. Importantly, the archaeological cultures are clearly differentiated between the areas.
What is more, it has been suggested that the Fatyanovo Culture people admixed with the local Volosovo
Culture HG after their arrival in European Russia6,17,36. Our results do not support this as they do not
reveal more HG ancestry in the Fatyanovo people compared to other CWC groups or any visible change
in ancestry proportions during the period covered by our samples (2,900–2,050 BC).

Finally, allele frequency changes in Western Russia and the Eastern Baltic revealed similar patterns in
both areas. Interestingly, the frequencies of alleles in the MCM6 and FADS2–3 genes, which have been
hypothesized to have increased due to dietary changes from the Neolithic onwards63,29,34, had significant
changes in frequency at different times – the MCM6 alleles by the CWC period and the FADS2–3 allele
after it. This points toward a more complex scenario for the onset of strong selection on these alleles
than just the arrival of farming, as has already been suggested previously13,29,34,64,28,33

Last Edit: Jul 16, 2020 20:14:57 GMT by Admin

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The Yamnaya Steppe Herders from Russia Nov 6, 2020 22:05:57 GMT

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Post by Admin on Nov 6, 2020 22:05:57 GMT

Published in the high-ranking journal PLOS ONE, their research was the result of innovative methodology. To determine snow cover and vegetation cycles, crucial to the survival of Bronze Age people and their flocks, they examined both satellite imagery and archaeological evidence, as well as interviewing modern-day herders.

In conjunction with researchers from the Institute of Archaeology, Chinese Academy of Social Sciences, Beijing, they then used these data to create a model of how the landscape was used more than 3,500 years ago.

"This detailed model of how Bronze Age people capitalised on the resources in their environment helps greatly in understanding the Prehistoric Silk Road," said lead author Dr. Peter Jia.

"For example, our ethnographic studies—interviews with local herders—have explained why certain locations were and still are chosen throughout the seasons: for the presence of early and late grass, optimal grazing potential in summer, and the absence of snow cover in the winter.

"Analysis of satellite imagery, together with the results of archaeological surveys and excavations, has enabled us to test these anecdotal accounts and prove their accuracy."

Dangers on the Steppe: why seasonal migration matters

The Eurasian steppe/mountain zone is a harsh environment. The main adaption to this arid landscape happened in the Bronze Age with the introduction of domesticated animals. But still today, it remains a place with inherent dangers to the livelihoods of people. Too much snow in winter and the animals cannot find enough food, dying by the hundreds in what locals call a 'white disaster'. Too little snow and there is not enough water for humans and animals, the feared 'black disaster'. Managing the landscape through seasonal migration is key to survival and maintenance of the economic system based on livestock.

Pioneering methodology

The strength of the study lies in its interdisciplinary approach, combining cutting-edge satellite technology with ethnographic and archaeological fieldwork.

Establishing growth cycles for vegetation of grazing lands and estimating the snow depth using satellite imagery allowed the researchers to assess the suitability of different parts of the mountains for herding in different seasons. Comparing these data with the accounts of local Mongolian and Kazakh pastoralists, they found them to be closely matching.

"Archaeology is one of the few fields that provides insights into how humans have interacted with the environment in the past," said co-author, Dr. Gino Caspari.

"With exacerbating environmental conditions worldwide, it is crucial to analyse this history.

"This task requires us to connect academic disciplines and cooperate internationally. Our study is a good example of this."

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The Yamnaya Steppe Herders from Russia Nov 7, 2020 6:09:56 GMT

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Post by Admin on Nov 7, 2020 6:09:56 GMT

Bronze Age pastoralists in what is now southern Russia apparently covered shorter distances than previously thought. It is believed that the Indo-European languages may have originated from this region, and these findings raise new questions about how technical and agricultural innovations spread to Europe. An international research team, with the participation of the University of Basel, has published a paper on this topic.

During the Bronze Age (ca. 3900-1000 BCE), herders and their families moved across the slopes of the Caucasus and the steppes to the north, taking their sheep, goats and cattle with them. It is believed that the Indo-Germanic groups, who brought the Indo-European languages and technical innovations such as wagons, domestic horses and metal weapons to Europe, may have originated from this region.

Until now, experts assumed that this transfer of technology was based on the long-distance migrations and trade contacts of these mobile pastoral communities, and that this mobility connected the Middle East with Europe. An international research team, with the participation of the University of Basel, has now questioned whether these communities did actually travel over such long distances. They published their study in the journal Plos One.

Nutrition reveals low levels of mobility

The researchers reconstructed the diet of the Bronze Age pastoral societies in order to draw conclusions about their migration. Their analysis was based on skeletal remains from burial mounds and flat grave cemeteries on the plateaus of the Caucasus and the steppes bordering to the north. "These human bones and teeth are archaeological treasures," says the study's author Professor Kurt Alt, visiting professor at the University of Basel and professor at Danube Private University in Krems. "They are fundamental resources for gaining a deeper understanding of economic strategies, the mobility patterns associated with them and social differentiation."

The research team analyzed the isotopic composition of carbon and nitrogen in bone collagen from the skeletal remains of 150 people, taken from eight sites. The finds date back to a period from about 5000 to about 500 BCE. In addition, the scientists compared this data with the isotope ratios in the bone collagen of 50 animals, as well as with the local vegetation of that time. The isotope ratios in bone collagen reflect the isotope ratios in the main foodstuffs that a person eats.

As it turns out, the diets of these groups were mainly based on the foodstuffs within the landscapes where their remains were found. "The communities apparently remained within their respective ecological areas and did not switch between the steppes, forest steppes or higher regions," explains Sandra Pichler from the Department of Environmental Sciences at the University of Basel, co-author of the study. According to the isotope analysis, meat, milk and dairy products formed a large part of these individuals' basic diets, but they were supplemented by wild plants, too. It was not until the end of the Bronze Age that their diets began to be based more on cultivated cereals, with millet presumably the main crop in this regard.

Technology transfer by word of mouth

"This study's findings imply that Caucasian communities were not highly mobile and did not undertake large-scale migrations, suggesting that the revolutionary technical innovations of the 4th and 3rd millennium BCE, such as wagons or metal weapons, were transmitted in other ways."

If the pastoral communities of the time only moved across shorter distances, technologies could have been passed on from one group to the next transmitting the knowledge of metal weapons, the processing of bronze and the domestication of horses into Europe by word of mouth.

More information: Corina Knipper et al, Diet and subsistence in Bronze Age pastoral communities from the southern Russian steppes and the North Caucasus, PLOS ONE (2020). DOI: 10.1371/journal.pone.0239861