The Copper Age began in Bulgaria ca. 5200-5000 BC, and Old European copper-trade network included the Pontic-Caspian steppe societies after ca. 4600 BC. The revolution of herding, travel, and raiding – and thus the change in the steppe – came with horseback riding, appearing ca. 4800 in early Khvalynsk, and spreading south- and eastward. The early Sredni Stog culture began about 4400 BC, and it seems that people from the east Pontic Caspian steppe (related to early Khvalynsk) brought a new culture[Anthony 2007], and probably also their Middle Indo-European language.
Within this new culture, a new elite group associated with the Suvorovo-Novodanilovka complex[Anthony 2007] was involved in raiding and trading with the lower Danube valley during the Trypillian B1 period, before and during the collapse of Old Europe. Settlements of Suvorovo-Novodanilovka chiefs have been found along the lower Danube, and sites of the posterior Cernavodă I culture seem to represent the assimilation of migrants from the steppes, therefore linked to Anthony’s first expansion from the Pontic-Caspian steppes into southern Europe ca. 4200-4000 BC[Anthony 2007][Anthony 2013].
Three samples dated ca. 5150 BC are found in early Khvalynsk, one of R1b1a-L754 (probably M269), one of R1a1-M459, and one of Q1a-F903 lineage. While the R1b1a-L754 sample was reported as from a high-status burial, similar to high-status individuals buried under kurgans in later Yamna graves, and therefore founder of an elite group of patrilineally-related families, the R1a1-M459 individual shows scarce decoration and his lineage is not found in later high-status Yamna graves[Mathieson et al. 2015 Supplementary materials]. A sample of the Mesolithic Dnieper-Donets culture (predating Sredni Stog in the same region), dated to a similar time and belonging to haplogroup R1a-M420[Jones et al. 2017], points – together with the diversity found in the Khvalynsk II cemetery – to a time preceding or coinciding with the successful expansion of R1b1a1a2-M269 lineages.
Two individuals of haplogroups R-M207 and R1b1a-L754 (dated ca. 4500 BC) in Smyadovo and one of haplogroup R1-M173 (dated ca. 4460 BC) in Varna I cemetery[Mathieson et al. 2017] are not proven to correspond to R1b1a1a2-M269 lineages, and dates are slightly early for the mass migration proposed by Anthony[Anthony 2007], so they could be assumed to correspond to old Balkan R1b1a1a-P297 branches. However, the region shows a discontinuity in R1b1a1a-P297 lineages in the Balkans after the arrival of Middle East Neolithic farmers (of G2-P15 lineages), which represent around half of more than 30 Y-DNA samples in the period from 6000 BC to 4500 BC[Mathieson et al. 2017]. The fact that haplogroup R-M207 is not found in later Balkan samples either (until the second Yamna expansion) is also significant.
Admixture analysis shows steppe-related ancestry (defined as a mixture of European and Caucasus hunter-gatherer similar to individuals from Yamna) in these samples, contrasting with the sex-biased resurgence of hunter-gatherer ancestry in central Europe and Iberia during the Middle Neolithic period – also supported by haplogroup replacement[Mathieson et al. 2017].
The older origin of haplogroup R1b1a1a2-M269 (ca. 11300 BC) compared to a later TMRCA (ca. 4300 BC) for the subclades survived in the modern population, coinciding with the successful spread of basal R1b1a1a2a-L23* (formed ca. 4300 BC, TMRCA ca. 4200 BC), point to an expansion occurred around this time period, probably from some eastern clans of Pontic-Caspian herders that developed the Sredni Stog culture in the west, and turned into Suvorovo-Novodanilovka chiefs and south-eastern European settlers. Both lineages are found in the Balkans, Central Europe, and Armenia[Myres et al. 2011][Herrera et al. 2012], and their expansion is therefore to be associated with the split of Proto-Anatolian[Kortlandt 1990][Ringe 2006] from a common Middle Proto-Indo-European language[Tischler and Oettinger 1989][Lehrman 1996][Melchert 1998].
The modern distribution of R1b1a1a2-M269 haplogroup in the Balkans and Anatolia (not reaching the Armenian highlands) points to the posterior migration of R1b1a1a2-M269 lineages with Anatolian languages. Its modern peak around Kosovo can be explained by posterior founder effects that might have happened during any expansion of peoples in the region in the past four thousand years, and which can tentatively be assigned to a recent Albanian expansion. Its modern distribution in the Alps and in ancient Tyrrhenia might point to an eastern route of the Suvorovo-Novodanilovka settlers of eastern Hungary, hence giving support to the theories describing Etruscan as an Anatolian branch[Adrados 1989][Adrados 1994]. On the other hand, it could well be a sign of independent back and forth migrations between the Adriatic Sea and the Italian Peninsula.
Modern distribution of R1b1a1a2-M269 (xL23) lineages, adapted from Richard Rocca (2012).
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