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The southern stream of Yamna migrants showed a later expansion from the Lower Danube and Tisza rivers to the southern tip of the Balkan peninsula, and its population is probably represented by the transition of Early Helladic II to Early Helladic III period ca. 2200 BC[Gimbutas 1977], coincident with the arrival of Minyan pottery style. Others have proposed a later date, the beginning of the Middle Helladic culture ca. 2000-1900 BC[Beekes 2011].
A recent study of Minoan samples from Crete (ca. 2900-1700 BC) and Mycenaean samples from mainland southern Greece (ca. 1700-1200 BC) have shown an introduction of CHG without EHG compared to Mesolithic samples, thus independent of a steppe invasion. This eastern influence is also found in Y-DNA haplogroups, all four of them J-M304, related to Bronze Age pastoralists of the Eurasian steppe. This eastern influence may have arrived from Anatolia, since Neolithic samples from central Anatolia at Tepecik-Çiftlik already show it some millennia earlier[Kilinc et al. 2016], and more recent samples from south-western Anatolian (ca. 2800-1800 BC) show it too.
Mycenaean samples show a ‘northern’ contribution, apart from the Neolithic farmer and ‘eastern’ ancestry shown by Minoan samples. This ‘eastern’ contribution may be interpreted in part as a steppe ancestry similar to that found in the Balkans in the third millennium, suggesting a rapid migration of Proto-Greek from the Balkans, although it is found only in samples from mainland Greece, and not in one sample from Crete[Lazaridis et al. 2017].
The introduction of millet and the horse in northern Greece seem to coincide, pointing to their introduction by horse breeding, millet-consuming cultures from the north or north-east, via river valleys leading to the Danube[Valamoti 2016].
The potential invasion and assimilation of R1b1a1a2a2-Z2103 settlers in Proto-Anatolian-speaking populations in Cernavodă III and Ezero cultures further confounds modern genetic studies, and more aDNA samples are needed to more clearly depict the expansion of both populations. Such a contact may have happened early during the southward migration, as suggested by Anatolian loanwords found in Greek.
The complexity of Y-DNA haplogroups found in the modern population of Greece bears witness to the thousands of years of European and Asian interaction in the formation of its peoples. While it is clear that Yamna ancestry does not represent a big part of its modern population in any study published to date, it is unclear how the ancient population was affected by the migration of peoples of R1b1a1a2a2-Z2103 lineages.
Analysis of modern Greek and Cretan lineages point to a Neolithic expansion of haplogroup R1b1a1a2-M269 in the region, which was found nearer to Italian than to Balkan lineages[King et al. 2008]. Analysis of Greek-Cypriot modern populations revealed the presence of R1b1a1a2a2-Z2103 lineages in easternmost and westernmost sides of the island, with central (R1b1a1a2a2c1a-Z2110) lineages appearing only in the east[Voskarides et al. 2016]. The early attestation of Mycenaean Greek in the island point to an early expansion of R1b1a1a2a2c1a-Z2110 lineages, but the early Anatolian influence over the island precludes a precise identification of their origin.
Most R1b1a1a2-M269 lineages in Greece are probably linked to the Bronze Age expansion associated with the Minyan pottery, in turn linked to an earlier Yamna expansion into south-eastern Europe. Most R1a1a1b2-Z93 lineages – whose proportion in Greece and in historically Greek Anatolia increases with latitude and in peripheral areas – are thus to be linked to the recent southern migration of Balkanic populations of R1a1a1b1-Z283 lineages (mainly Slavs, but also Aromanians, and Albanians), and to the western spread of Iranian peoples of R1a1a1b2-Z93 lineages[Heraclides et al. 2017].
A potentially older invasion of certain R1a1a1b2-Z93 lineages during the Bronze Age could be supported by the finding of haplogroup R1a1a1b2-Z93 in an individual from Merichleri, in Bronze Age Bulgaria, ca. 1690 BC[Mathieson et al. 2017].
On the controversial ethnicity and language of the Sea Peoples and the closely related Philistine question [Woudhuizen 2006][Maeir, Davis, and Hitchcock 2016][Middleton 2015], genetic research points to a mixture of steppe ancestry found in the Lebanese population that occurred ca. 1740-160 BC [Haber et al. 2017] – probably in the earlier part of that period, possibly still earlier than the estimated range –, showing a clear haplogroup turnover in the modern Levantine population, with new subclades R1b1a1a2a2-Z2103 potentially from early Balkan (IE) individuals, and R1a1a1b2-Z93 probably from eastern (Iranian) migrants.
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