Red circle, tower landmark, Fleury sur Orne: Long Burrow

Ancient H2 Timeline: Migration Mediterranean routes of the Neolithic expansion
Mediterranean route into Western Europe, the Iberian Peninsula and ultimately, Ireland.
Rohrlach, A.B., Papac, L., Childebayeva, A. et al. Using Y-chromosome capture enrichment to resolve haplogroup H2 shows new evidence for a two-path Neolithic expansion to Western Europe. Sci Rep 11, 15005 (2021).
H2 in Fleury-sur-Orne, dated 4700-4300 BC (Monuments, male elite of the Cerny culture, Neolithic)
H2 North of Kilkenny, dated 3600-3400 BC
H2 Nakht-Ankh and Khnum-Nakht, Egypt 12th Dynasty, Middle Kingdom dates of 1985-1773 BC

https://www.pnas.org/doi/pdf/10.1073/pnas.2120786119
Ancient DNA gives new insights into a Norman Neolithic
monumental cemetery dedicated to male elites
Nine page report with charts and photos.
The Middle Neolithic in western Europe is characterized by monumental funerary
structures, known as megaliths, along the Atlantic fac¸ade. The first manifestations of
this phenomenon occurred in modern-day France with the long mounds of the Cerny
culture. Here, we present genome-wide data from the fifth-millennium BCE site of
Fleury-sur-Orne in Normandy (France), famous for its impressively long monuments
built for selected individuals. The site encompasses 32 monuments of variable sizes,
containing the burials of 19 individuals from the Neolithic period. To address who was
buried at the site, we generated genome-wide data for 14 individuals, of whom 13 are
males, completing previously published data [M. Rivollat et al., Sci. Adv. 6, eaaz5344
(2020)]. Population genetic and Y chromosome analyses show that the Fleury-sur-Orne
group fits within western European Neolithic genetic diversity and that the arrival of a
new group is detected after 4,000 calibrated BCE. The results of analyzing uniparentally inherited markers and an overall low number of long runs of homozygosity suggest
a patrilineal group practicing female exogamy. We find two pairs of individuals to be
father and son, buried together in the same monument/grave. No other biological relationship can link monuments together, suggesting that each monument was dedicated
to a genetically independent lineage. The combined data and documented father–son
line of descent suggest a male-mediated transmission of sociopolitical authority. However, a single female buried with an arrowhead, otherwise considered a symbol of power
of the male elite of the Cerny culture, questions a strictly biological sex bias in the burial
rites of this otherwise “masculine” monumental cemetery

2 path expansion of Hap H
https://en.wikipedia.org/wiki/Haplogroup_H_(Y-DNA)
https://www.nature.com/articles/s41598-021-94491-z
As a case study, we focus on H2, a haplogroup associated with a critical event in European human history: the Neolithic transition. By disentangling the evolutionary history of this haplogroup, we further elucidate the two separate paths by which early farmers expanded from Anatolia and the Near East to western Europe.

We highlight the improved performance obtained via YMCA by analysing the Y-chromosomal haplogroup H2 (H-P96), a low-frequency YHG that is associated with early farmers during the Neolithic transition in Western Eurasia. We curated a data set of 46 previously published individuals (45 ancient and 1 modern), and 49 newly YMCA-sequenced individuals (all ancient). We show that our current understanding of H2, which is based largely on modern H2 samples (n = 20), is inconsistent with the ancient diversity of our H2 individuals. In resolving this ancient haplogroup, we can show two distinct migration paths along the Mediterranean and Danube for Neolithic groups from Anatolia to Western Europe, ultimately resulting in the Mediterranean-derived groups also reaching Britain and Ireland.

The YHG H (H-L901) is thought to have formed in South Asia approximately ~ 48 kya34. Three subsequent sub-haplogroups, H1 (H-M69), H2 (H-P96) and H3 (H-Z5857), appear to have quickly formed over the following 4000 years. H1 and H3 have estimated formation times of ~ 44.3 kya, however, H2 is estimated to have formed slightly earlier at ~ 45.6 kya https://www.yfull.com].

Haplogroups H1 and H3 are still found in frequencies as high as 20% in South Asia35, but in extremely low frequencies in Europe, with H1 only being found associated with the spread of the Romani people ~ 900 ya. Conversely, H2 has been present in Western Eurasia since at least 10 kya36, and is strongly linked with the spread of agriculture37,38, but is found at no higher than 0.2% frequency in modern-day western European populations. In contrast, H2 was more common in Neolithic groups, and has been found to have constituted between 1.5 and 9% of the observed Y haplogroups, with the exception of the highly related individuals from Rivollat et al. 2020, for which H2 was ~ 30%15,19,24,39,28,36,38,40,41.

With the arrival of Steppe-related ancestry ~ 5 kya, incoming YHGs such as R1a and R1b would largely replace many of the older, “Neolithic” YHGs, such as G2, T1a, and H219, and although H2 was never found in particularly high frequencies among Neolithic individuals, we expect that its diversity was also greatly reduced, and many sub-lineages were potentially lost altogether.

To test whether our YMCA could improve the haplotyping quality to a point which would allow us to also draw phylogeographic inferences, we made use of newly collated collection of prehistoric ancient human DNA data and selected individuals, who were tentatively assigned to YHG H2. We generated new data for n = 49 individuals, and merged this with n = 46 published Y-chromosomal genomes (see Tables S1 and S2). While H2 is commonly found alongside the more dominant Neolithic YHG G2a (G-Z38302)24,37, it is precisely the low frequency of H2 which is of interest here. The relative scarcity of H2 individuals, especially compared to the relatively high frequency of the accompanying G2a individuals, allows us to better track the ‘genealogical history’ and thus potential dispersal routes as we would expect a stronger effect of lineage sorting and therefore a higher chance of observing geographic patterns. In this particular case, we could trace expanding Neolithic farmers from Anatolia to Western Europe through the use of unique markers associated with H2 individuals and test whether we can genetically discern the proposed so-called “Danubian or inland'' and “Mediterranean'' routes of the Neolithic expansion42, which had recently also found support by genomic signals from the nuclear genome38.

**When we plotted all of the samples in our study on a map of Europe, a phylogeographic pattern clearly emerged (Fig. 3B. The H2d individuals are all found along the so-called inland/Danubian route into central Europe, and all but one of the H2m individuals are found along the so-called Mediterranean route into Western Europe, the Iberian Peninsula and ultimately, Ireland. The solitary H2m individual (LEU019) found in central Germany is dated to the Late Neolithic/Early Bronze Age context, postdating the Neolithic expansion by 2000–3000 years. Archaeological and mtDNA evidence of an eastward expansion of Middle/Late Neolithic groups such as Michelsberg43,44,45 could potentially explain this single geographically outlying observation.

However, we were also confident in the quality of our samples as H2 is a very rare modern haplogroup, with only 19 individuals being downstream of H2-P96 on YFull at the time of this publication. Hence, if any of our samples had been contaminated by a male source, it would be readily noticeable in bam pileups as derived alleles for another haplogroup, which means these samples would not have been identified as H2, and hence would not be in the study.

Web Page Updated: https://pbase.com/daveb/flr_h2