Caucasus Hunter-Gatherer ancestry and Indo-Hittite
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The expansion of R1a1a1-M417 lineages may have disrupted the Early Proto-Indo-European R1b1a1a-P297 community thriving in east Europe. In this context, R1a-M417 lineages might have spoken Uralo-Yukaghir languages when arriving in the Forest Zone from the east, and Uralic could therefore be considered a superstratum over a Pre-Indo-European substratum. A Uralo-Yukaghir community spread over Eurasia is supported by the east-west direction of cultural innovations in the region, and by the finding of maximum Ancient North Eurasian ancestry in modern-day Kets, Mansi, Native Americans, Nganasans and Yukaghirs (Flegontov et al. 2016). On the other hand, R1a1a1-M417 lineages may have brought a Yukaghir superstratum to the Indo-Uralic spoken in the Forest Zone (an equivalent to Early Proto-Indo-European in this macro-family proposal) by R1b1a1a-P297 communities similar to the Narva samples, developing a Proto-Uralic-speaking community.
Given the early sample of R1a-M420 in the Mesolithic north Pontic area – and maybe the rare subclade R1a5-Z645 found in Estonia (Saag et al. 2017) –, it would also be possible, although unlikely given the available archaeological data, to propose that only pottery was adopted from the east, and that the migration of R1a-M420 subclades (including R1a1a1-M417) happened from the north-west Pontic area, with a back-migration of these lineages to the Baikal region.
In any case, a conservative view will be observed in this paper, assuming Proto-Uralic to be the language of hunter-gatherers in the Forest Zone, north of the Pontic Caspian steppe, at the same time as Middle Indo-European was spoken in the steppes in the transition to the Neolithic (Parpola 2012).
The arrival of N1a1-M46 (previously called N1c1) lineages into Northern Europe has been dated after 5000 BC (Ilumae et al. 2016), coinciding with the TMRCA of N1a1a1a-L708. However, the more recent formation (ca. 3800 BC) and TMRCA (ca. 2300 BC) of common European lineage N1a1a1a1a-L392 point to a late and stepped spread of these hunter-gatherer groups into the Forest Zone, that cannot be associated with the expansion of Pit-Comb Ware cultures. Cultural assimilation remains the best explanation at the moment for the shared Uralic languages of modern R1a1a1-M417 and N1a1-M46 communities. Eastern groups with N1a1a1a1a-L392 lineages may have brought with them the Altaic traits found in Uralic languages (Kortlandt 2010).
Even though an aDNA sample of haplogroup N1a-F1206 is found in the Forest Zone dated ca. 2500 BC at Serteya (Chekunova et al. 2014), it is tempting to place the mass migration of Siberian hunter-gatherer communities around the Urals later, with the expansion of the poorly understood Seima-Turbino phenomenon (which began ca. 2000 BC in East Asia), since it connected cultures from Mongolia to Finland. Three samples of haplogroup NO (xO) found in the Middle Bronze Age Okunev culture, and two samples later in the Chermuchek culture area (Hollard et al. 2014) may give support to this assumption.
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Saag, Lehti, Liivi Varul, Christiana Lyn Scheib, Jesper Stenderup, Morten E Allentoft, Lauri Saag, Luca Pagani, Maere Reidla, Kristiina Tambets, Ene Metspalu, Aivar Kriiska, Eske Willerslev, Toomas Kivisild, and Mait Metspalu. 2017. Extensive farming in Estonia started through a sex-biased migration from the Steppe. bioRxiv.