The genomic ancestry of the Scandinavian Battle Axe Culture people and their relation to the broader Corded Ware horizon

 https://royalsocietypublishing.org/doi/10.1098/rspb.2019.1528

We generated and analysed genome sequence shotgun data from 11 individuals originating in northeastern Europe datedto 3300–1660 cal BCE (electronic supplementary material,table S1). Five individuals were excavated from CWC contexts:two from Obłaczkowo, Poland, one from Karlova, Estonia,and two from the CWC-related BAC burial Bergsgraven in Linköping, Sweden. The six additional individuals werefrom other archaeological contexts in Sweden: five from megalithic burial structures primarily associated with Funnel BeakerCulture (FBC) (two from Rössberga in Västergötland and threefrom Öllsjö in Scania) and one from a Pitted Ware Culture(PWC) context (Ajvide on Gotland). Radiocarbon dating showed that the three individuals from the Öllsjö megalithic tomb derived from later burials, where oll007 (2860–2500 calBCE) overlaps with the time interval of the BAC, and oll009and oll010 (1930–1650 cal BCE) fall within the Scandinavian Late Neolithic and Early Bronze Age (table 1; electronic supplementary material, table S1 and figure S2). Genome-wide sequence coverages range from 0.1 to 3.2×, and the sequence data for all individuals exhibit characteristic properties ofancient DNA: short fragment size and cytosine deamination at the ends of fragments (e.g. [24]) (table 1; electronic sup-plementary material, figure S5). Estimates of mitochondrialcontamination [25] were low, less than 2% for all 11 individuals, as was the estimated nuclear contamination on theX-chromosome in males [26,27] (less than 1.2%) (table 1; electronic supplementary material, table S3). Five individuals were genetically determined to be males, and six were females,based on the fraction of sequence fragments mapping to thedifferent sex chromosomes [28] (table 1)

The other individuals who were contemporary with BAC but had unclear cultural contexts, and who were buried in the Ölljsö megalith constructed many hundred years earlier (oll007), or found as a stray find (Ölsund) [9], show the same genetic profile as individuals from typical BAC contexts in other parts of Sweden

The best fitting two-source model was Funnelbeaker (FBC) + Yamnaya (YAM) (p = 0.86), while YAM + PWC (Pitted Ware Culture) would also fit the data (although trending toward low p-values; p = 0.07). These observations suggest that—to our statistical resolution—a direct PWC contribution to BAC is not needed in a model, but actual PWC admixture might have been small or there may have been indirect PWC contributions through PWC first mixing with FBC [34] who later contributed ancestry to BAC. Notably, using only the CWC population as the single source for BAC was consistent with the data in all cases (p > 0.05, except when using CWC-associated individuals from Latvia, CWC_LV).  The BAC groups fit as a sister group to the CWC-associated group from Estonia (CWC_EE, electronic supplementary material, figure S8) but not as a sister group to the CWC groups from Poland (CWC_PL, figure 3) or Lithuania (CWC_LT, electronic supplementary material) (|Z| > 3), indicating some differences in ancestry between these CWC groups and BAC. 

Funnelbeaker tombs were reused by the Battle Axe Culture.

The Scandinavian Middle Neolithic megalithic tombs are associated with the FBC. However, their reuse, indicated by artefacts common to the BAC and later periods, has been noted [17]. The oll007 individual, buried in the FBC-associated Öllsjö megalithic tomb, but radiocarbon dated to the time period of the BAC, is genetically very similar to individuals from BAC contexts (e.g. Bergsgraven and Viby). Thus, although archaeologically the reuse of megalithic tombs was assumed earlier [17], our study may be the first direct link (using genetics) showing that indeed FBC-associated megalithic tombs were used as burial places also for the people of the BAC. This could possibly also extend to the Danish Single Grave Culture (SGC) [49], as RISE61 [2], a male buried in the Kyndeløse passage grave and with a radiocarbon date overlapping with the BAC/CWC/SGC time period, also displays some steppe ancestry. 

 

Supplementary information: https://royalsocietypublishing.org/action/downloadSupplement?doi=10.1098%2Frspb.2019.1528&file=rspb20191528supp1.pdf

 

More here: https://onlinelibrary.wiley.com/doi/10.1002/ajpa.24079

 

 

The Fatyanovo were swarthy & not lactose tolerant

 https://www.science.org/doi/10.1126/sciadv.abd6535

The presented genome-wide data are derived from 3 Stone Age HGs (WeRuHG; 10,800 to 4250 cal BCE, BER001, KAR001, PES001) and 26 Bronze Age Fatyanovo Culture farmers from western Russia (Fatyanovo; 2900 to 2050 cal BCE) and 1 Corded Ware Culture individual from Estonia (EstCWC; 2850 to 2500 cal BCE)

In the case of radiocarbon dating, it is possible that fish from rivers and lakes consumed by Stone Age fisher-hunter-gatherers may cause a notable reservoir effect. This means that the radiocarbon dates obtained from the human bones and teeth can be hundreds but not thousands of years older than the actual time these people lived (33). Unfortunately, we do not yet have data to estimate the size of the reservoir effect for each specific case. Then, we turned to the Bronze Age Fatyanovo Culture individuals and determined that they carry 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, fig. S1, and tables S2 to S4) that have also been found in CWC individuals elsewhere in Europe (14–16, 18, 27). In all individuals for which the chrY hg could be determined with sufficient depth (n = 6), it is R1a2-Z93. 

The Fatyanovo are the most likely candidates for pre-Proto-Indo-Iranians, representing the earliest stage Pre-Proto-IIr splitting up from what would be Proto-Balto-Slavic. 

We estimated the time of admixture for Yamnaya and EF populations to form the Fatyanovo Culture population using DATES (37) as 13 ± 2 generations for Yamnaya Samara + Globular Amphora and 19 ± 5 generations for Yamnaya Samara + Trypillia. If a generation time of 25 years and the average calibrated date of the Fatyanovo individuals (~2600 cal BCE) are used, this equates to the admixture happening ~3100 to 2900 BCE. 

The confidence intervals (CIs) were extremely wide with Trypillia, but the chrX data showed 40 to 53% Globular Amphora ancestry in Fatyanovo, in contrast with the 32 to 36% estimated using autosomal data. The sex-biased admixture is also supported by the presence of mtDNA hg N1a in two Fatyanovo individuals—an hg frequent in Linear Pottery Culture (LBK) EFs but not found in Yamnaya individuals so far 

The Fatyanovo were formed as a result of the process of Yamnaya male invaders taking European farmer women as wives, resultant genome being 2/3 Yamnaya and 1/3 EEF. The chrX data is based on a horribly low 45k snps so I will take it with a grain of salt, but the Y-DNA makes it somewhat clear still.

From the phenotype prediction results, we see that only 4% of Fatyanovo samples had blonde/dark blonde hair, while 96% had dark hair and 80% had brown eyes. 60% of the samples had an intermediate skin tone, and the remaining 40% had a combination of mixed skin tone, unpredictable intermediate and just straight up dark skin. Lactase persistence was only 17%, clearly showing that it had already undergone selection from Steppe EMBA times. A gene for high serum cholesterol was 83% but drops to barely 13% by the Estonian Middle Ages, showing strong negative selection for serum cholesterol possibly.

Outgroups used in the study:  Mota, Ust-Ishim, Kostenki14, GoyetQ116, Vestonice16, MA1, AfontovaGora3, ElMiron, Villabruna, WHG, EHG, CHG, Iran_N, Natufian, Levant_N, and Anatolia_N