The expansion of Balkan Indo-European
Later samples of R1b1a1a2a2-Z2103 subclades suggest a genetic continuity of early Khvalynsk population in the pastoral groups within Volga–Ural–North Caucasian steppe: early Yamna in Ekaterinovka ca. 2840 BC, late Yamna in Temrta IV ca. 2760 BC, and two early samples from the Poltavka culture in Lopatino II ca. 2770 and in Kutuluk ca. 2680 BC (Mathieson et al. 2015).
Eastern groups – if only because of their location within Yamna – may have followed the southern stream somehow later compared to western R1b1a1a2a1-L51 lineages, possibly through the settlement of Kovachevo-Troyanovo. The sample at Beli Manastir of the Vučedol culture (see above) shows how different groups of different lineages might have been involved in the Yamna migration to south-eastern Europe.
R1b1a1a2a2-Z2103 lineages appear in modern populations (Figure 11) as a Balkan group – mainly of R1b1a1a2a2c-Z2106 lineages, like the sample found in Stalingrad Quarry dated ca. 2670 –, with a central group dominated by R1b1a1a2a2c1a-Z2110 found in Greece and Central Europe; an Armenian group – of R1b1a1a2a2a-L584 lineages – found mainly in the Armenian highlands; and a northern group – of R1b1a1a2a2b-L277.1 lineage, possibly related to the Vučedol sample ca. 2775 BC and the Bell Beaker sample from Szigetszentmikklós ca. 2330 BC (see above).
The modern distribution of R1b1a1a2a2-Z2103 subclades around the Balkans gives therefore support to the existence of a Paleo-Balkan Sprachbund or dialect continuum (Kortlandt 2003).
Myres, N. M., S. Rootsi, A. A. Lin, M. Jarve, R. J. King, I. Kutuev, V. M. Cabrera, E. K. Khusnutdinova, A. Pshenichnov, B. Yunusbayev, O. Balanovsky, E. Balanovska, P. Rudan, M. Baldovic, R. J. Herrera, J. Chiaroni, J. Di Cristofaro, R. Villems, T. Kivisild, and P. A. Underhill. 2011. A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe. Eur J Hum Genet 19 (1):95-101.
Kortlandt, F. . 2003. Armeniaca. Comparative notes. Ann Arbor: Caravan books.
Mathieson, I., I. Lazaridis, N. Rohland, S. Mallick, N. Patterson, S. A. Roodenberg, E. Harney, K. Stewardson, D. Fernandes, M. Novak, K. Sirak, C. Gamba, E. R. Jones, B. Llamas, S. Dryomov, J. Pickrell, J. L. Arsuaga, J. M. de Castro, E. Carbonell, F. Gerritsen, A. Khokhlov, P. Kuznetsov, M. Lozano, H. Meller, O. Mochalov, V. Moiseyev, M. A. Guerra, J. Roodenberg, J. M. Verges, J. Krause, A. Cooper, K. W. Alt, D. Brown, D. Anthony, C. Lalueza-Fox, W. Haak, R. Pinhasi, and D. Reich. 2015. Genome-wide patterns of selection in 230 ancient Eurasians. Nature 528 (7583):499-503.