Some of the most interesting bits of the paper for me:
"T
he pre-Greek influence is most plausibly encapsulated by the following G2a haplogroups: U5*, PF3147*, L91, L293, P303*, and CTS342. Notably, most of these lineages occur in Anatolian ancient DNA specimens over 8200 years old [53]. In addition, some J2a lineages such as M67, M319, and J2b-M205 may also have contributed to the pre-Greek strata."
I know quite a few G2a southern Italians who carry those lineages. They'll be interested to see how long their paternal lineages may have been in place.
They link the G2a lineages they see with the Pre-Pottery Neolithic and the J2b possibly with Pottery Neolithic.
"Both genetic data from the literature and Syria (Chiaroni J, unpublished results) show a frequency peak of J2b-M205 in the Southern Levant in which the frequency decreases northwards with latitude (Pearson’s R 2 = 0.282, p value = 0.011). The J2b-M205 distribution coincides with the Neolithic crop package dissemination found in common between the Levant and Cyprus [5]. Also J2b-M12 Td estimates (Table 3) coupled with the J2b-M205 distribution overlap significantly with the Pottery Neolithic to Early Bronze Age pattern of settlements in Nicosia, Pafos, Limassol, and Kyreneia (chi-square = 11.29, p < .00084). This suggests the possibility that Cyprus experienced a later (Pottery Neolithic) immigration from the Southern Levant."
As to J2a:
"Regarding J2a-M410, the most common M530 sublineages are J2a-Z387 (4.9 %) linked with the distinctive six repeat allele at the DYS445 short tandem repeat locus proposed [
18] to represent a Neolithic expansion from Anatolia to Greece and Italy, a pattern similar to G2a-M406 [
30], at 4.9 % and J2a-Page55*(xM67,M319,Z387) at 5.2 %."
They also see an arrival of some of the yDna "E" lineages in the Neolithic:
"
E-V22 and E-M34 are common in the Southern Levant, Sicily, Algeria, and in Egypt and rare in Europe [27, 36]. These lineages, like J2b-M205, could mirror a Pottery Neolithic movement to Cyprus from the Southern Levant (Pearson R 2 coefficient of correlation of E- M34 to longitude: 0.164, p = 0.003)."
From this period until the Bronze Age, the population of Cyprus seems to have been relatively isolated. Then came the metal prospectors and traders of the Bronze Age.
"
Recent insights from ancient DNA studies suggest the spread of genes during the Bronze Age involved J2a-M67 individuals who appear in the Central European plains during the Late Bronze Age [86]. J2a-M67, proposed to represent both the Neolithic of Central Anatolia and the expansion of the Troia Maritime Culture in Northwestern Anatolia (13.5 %) [31], is also quite common in Cyprus (10.1 %). Testing the hypothesis that the origin of the Early Bronze Philia culture in Cyprus derives from Western/Northwest Anatolia, the distribution of J2a- M67 on Cyprus fits well (chi-square = 3.42, p < .032, one-tailed). J2a-Z489, present in Pafos and Northwest/Central Anatolia, may reflect Bronze Age immigration from Western Anatolia, the Philia phase, or mirror the Jewish population on the island from the Hellenistic/Roman Eras."
Their discussion of the yDna lineages is also very interesting, with a split between E-V13 and the others. They propose a movement of E-V13 later with a late Bronze Age expansion, an expansion that has previously been proposed. They don't tackle when the pre-EV13 arrived in the Balkans, although we know pre-E-V13 was there by around 4500 BC.
"
E-V13 is common in the Balkans and may mark some of the Greek demographic input to Cyprus from the Late Bronze Age through the Iron Age [79]. Network analysis of 46 E-V13 haplotypes (Additional file 10: Figure S5) shows a discrete clustering of 15 samples suggestive of a sub-haplogroup (encircled with an oval). This cluster is characterized by DYS437 = 15 repeats not seen in the Anatolian Greek population, or in the Provence samples [20]. The remaining 31 samples overlap with the Anatolian Greek E-V13 lineages."
This is what they have to say about the R1b Z2105 of Yamnaya fame.
" In Cyprus, Z2105 has the opposite distribution from J2a-M67 and J2b-M205, concentrating in the East/Northeast regions of Cyprus (chi-square = 5.01, p < .0256). The Late Bronze Age cities of Enkomi, Kition, and Hala Sultan Tekke, found in the Larnaka and Ammochostos districts, may have received immigrants from Hittite/Luwian Anatolia involved in the trade of the Late Bronze Age Eastern Mediterranean."
The Myceneans may also have carried this lineage. In terms of distinguishing the two strands, this is what they have to say:
"
Updating previous R1b-L23* data pertinent to Cyprus [18, 28, 56] reveals that while the presence of R1b-Z2105 in central and west Europe is minimal, conversely, it is informative in present-day Anatolia (10.2 %), Greece (7.0 %), Bulgaria (5.7 %), and Crete (3.1 %). While the emerging R1b-Z2105 substructure [64] has yet to be evaluated at population levels, one relevant sub-haplogroup defined by M589 is illuminating. The majority of Cypriot R1b-Z2105 lineages occur in both the westernmost and easternmost districts, Pafos and Ammochostos, respectively. Notably, of the two districts, the M589 sub-haplogroup occurs only in the east. The absence of M589 lineages in Crete and the Balkans and the presence in Asia Minor are compatible with a record of Late Bronze Age influences from Anatolia rather than from Mycenaean Greeks."
There's a lot of upstream Z2105 in southern Italy, particularly in Calabria. It would be interesting to see the subgroups there.