which had strong cultural influence on the "Epi-Jōmon" of southern Hokkaido and northern Honshu, but that the Ainu people themselves formed from the combination of both ancient groups. Additionally he notes that the historical distribution of Ainu dialects and its specific vocabulary correspond to the distribution of the maritime Okhotsk culture. A 2021 study confirmed that the Hokkaido Jōmon population formed from "Terminal Upper-Paleolithic people" (TUP people) indigenous to Northern Eurasia and from proper Jōmon people, which arrived from Honshu about 15,000 BC. The Ainu in turn originated from the Hokkaido Epi-Jōmon and from the Okhotsk people in Hokkaido. Another study in 2021 (Sato et al.) analyzed the indigenous populations of northern Japan and the Russian Far East. They concluded that Siberia and northern Japan was populated by two distinct waves: Genetics Paternal lineages Genetic testing has shown that the Ainu belong mainly to Y-DNA haplogroup D-M55 (D1a2) and C-M217. Y DNA haplogroup D M55 is found throughout the Japanese Archipelago, but with very high frequencies among the Ainu of Hokkaidō in the far north, and to a lesser extent among the Ryukyuans in the Ryukyu Islands of the far south. Recently it was confirmed that the Japanese branch of haplogroup D M55 is distinct and isolated from other D branches for more than 53,000 years. Several studies (Hammer et al. 2006, Shinoda 2008, Matsumoto 2009, Cabrera et al. 2018) suggest that haplogroup D originated somewhere in Central Asia. According to Hammer et al., the ancestral haplogroup D originated between Tibet and the Altai mountains. He suggests that there were multiple waves into Eastern Eurasia. A study by Tajima et al. (2004) suggest that fourteen out of sixteen Ainu (or 87.5%) belong to YAP+ lineages (Y-haplogroups D-M55* and D-M125), with 13/16 (81.3%) belonging to D-M55 and 1/16 (6.25%) belonging to D-M125 (the latter is much more typical of mainland Japanese males than Ainu). The presence of Haplogroup C M217 in the Ainu suggest a degree of genetic admixture with the Nivkhs. Two out of a sample of sixteen Ainu men (or 12.5%) belong to , which is the most common Y chromosome haplogroup among the indigenous populations of Siberia and Mongolia. Hammer et al. (2006) found that one in a sample of four (or 25%) Ainu men belonged to haplogroup C M217. Maternal lineages Based on analysis of one sample of 51 modern Ainu, their mtDNA lineages consist mainly of