Out of Eden: The Peopling of the World

by Oppenheimer, Stephen

  We have looked at three Asian lines from the north and three Southeast Asian probables, so that leaves D, E and G. These daughters of M/Manju are potentially related at their origin (since they are all M’s and share a mutation at the fast (unstable) site of 16362 – see Kivisild et al. (1999, 2002) op. cit.). D and G are widely distributed throughout the Mongoloid dispersal right down as far as Indo-China. Unique local versions of D are found at high rates throughout China, Japan, Mongolia, Tibet, Korea, Central and Northeast Asia, and the Americas (Torroni et al. (1994) op. cit.; Torroni, A. et al. (1993) ‘Asian affinities and continental radiation of the four founding Native American mtDNAs’ American Journal of Human Genetics 53: 563–90; Kolman et al., op. cit.; Kivisild et al. (1999, 2002) op. cit.). Although found in southern China, D is not a feature of Southeast Asia (Torroni et al. (1994) op. cit.). Unique versions of G are found in Central Asia (see Fig. 2 in Metspalu et al., op. cit.), Tibet (Torroni et al. (1994) op. cit.), the Ainu (Horai et al., op. cit.) and Northeast Asia (Torroni et al. (1994) op. cit). Group E has its own unique subgroup in Island Southeast Asia, and is also found in Tibet (Torroni’s types 83, 89, 94, 104, 106, 109, and 119 – misidentified due to the presence of RFLP site at base 16517, Torroni et al. (1994) op. cit.; Fig. 2 in Metspalu et al., op. cit.). A newly described clade, M7 (age 61,000 ± 20,000 years) has a broad East Asian coastal distribution, like D but, like E, extends down into Southeast Asia: see Kivisild et al. (2002) op. cit.; Yong-Gang Yao et al., op. cit.

  34. Data for Central Asia from Fig. 2 in Metspalu et al., op. cit.; for Tibet, Torroni et al. (1994) op. cit.; for Mongolia, Kolman et al., op. cit.; for China and Korea, Torroni et al. (1994) op. cit.

  35. age of four of these lines in Mongolia: Kivisild et al. (1999) op. cit. (note that there is the possibility of carried-over diversity affecting local age estimates in Mongolia). ages of the same lines farther south in China: Table 3 in Yong-Gang Yao et al., op. cit. Note that in Yong-Gang Yao et al. individual branches of D and G age at 51,000–60,000 years, and these two haplogroups coalesce with M9/E even earlier (Kivisild et al. (2002) op. cit.).

  36. Group B, with an estimated Asian age of about 75,000 years (74,600 ± 18,700 years, Yong-Gang Yao et al., op. cit.), achieves its highest frequencies in peoples of Southeast Asia, Oceania (excluding New Guinean highlanders and Australians), and the west Pacific coast.

  37. For B and F diversity and antiquity in the south, see Ballinger, S.W. et al. (1992) ‘Southeast Asian mitochondrial DNA analysis reveals genetic continuity of ancient Mongolid migrations’ Genetics 130: 139–52. Data also from Fucharoen, G. et al. (2001) ‘Mitochondrial DNA polymorphisms in Thailand’ Journal of Human Genetics 46: 115–25; and Oota, H. et al. ‘Extreme mtDNA homogeneity in continental Asian populations’ American Journal of Physical Anthropology 118: 146–53; see also Kivisild et al. (2002) op. cit.; Yong-Gang Yao et al., op. cit.: Groups B (estimated age 74,600 ± 18,700 years) and R9(F4) (estimated age 81,000 ± 24,600 years) which is ancestral to F in the south (see above in note 33). For the newly identified pre-F haplogroup now, by agreement re-classified as R9, see Hill, C. et al. op. cit. See Fig 5.5. The newly described M7 haplogroup mirrors B and F. M7 dates to 61,000 years, and its oldest branch, M7b, is a feature of Vietnamese populations – Kivisild et al. (2002) op. cit.

  38. two common subgroups of B: The oldest in Mongolia (40,500 years), B1, is the dominant type throughout Southeast Asia, the Pacific, and the Americas. B1 is also the type found in Tibet – Kivisild et al. (1999) op. cit.; Torroni et al. (1994) op. cit. B1 and B2 are differentiated respectively by 16217 (now generally classified as B4) and 16243/16140 (now reclassified by Kivisild et al. (2002) op. cit. as B5b) in Kolman et al., op. cit. The younger of the two, B2 (33,500 years in Mongolia), features rather more than B1 on the west Pacific coast (Eastern China, Korea, Japan) – Horai, S. and Hayasaka, K. (1990) ‘Intraspecific nucleotide sequence differences in the major non-coding region of human mitochondrial DNA’ American Journal of Human Genetics 46: 828–42; Nishimaki, Y. et al. (1999) ‘Sequence polymorphism in the mtDNA HV1 region in Japanese and Chinese’ Legal Medicine 1: 238–49; Horai et al. (1996), op. cit; Seo, Y.B. et al. (1998) ‘Sequence polymorphism of mitochondrial DNA control region in Japanese’ Forensic Science International 97: 155–64. both B types during the Palaeolithic: Fig. 2 in Metspalu et al., op. cit.

  39. three partly related Manju lines D, E, and G: the most dominant of these, D at 44,500 years (Kivisild et al. (1999) op. cit.), is commonest in Siberia, and is present in South China and the whole of the Americas but not Southeast Asia, i.e. Sundaland (Torroni et al. (1994) op. cit.). D is also present at low rates in India (Kivisild et al. (1999) op. cit.). D and E are also common in western Central Asia (Fig. 2 in Metspalu et al., op. cit.; for E see Ballinger et al., op. cit). A variant of E known to occur in Korea is found commonly in Southeast Asia, raising the possibility that the ancestor of this group also ultimately came from the south. There is also a link between G and unique Semang M group M21c, again suggesting a possible ultimate southern source for this group (non-coding site 5108 in Hill et al., op. cit.). sister branches C and Z reach their highest rates: Kolman et al., op. cit.; Bamshad, M. et al. (1999) ‘Genetic evidence on the origins of Indian caste populations’ Genome Research 11: 994–1004; Yong-Gang Yao et al., op. cit.; Kivisild et al. (2002) op. cit. Groups C and Z in India, Turkey, Central Asia, and Mongolia: Kivisild et al. (1999) op. cit. (C only in India), Bamshad et al., op. cit. age of C in Mongolia: Kivisild et al. (1999) op. cit. .

  40. X: a single report from southern Siberia: Derenko, M.V. et al. (2001) ‘The presence of mitochondrial Haplogroup X in Altaians from South Siberia’ American Journal of Human Genetics 69: 237–41. up to 30,000 years old: Brown et al., op. cit.

  41. D/E, C, and F are the new consensus names respectively for YAP+, RPS4Y/M216, and M89 – see The Y Chromosome Consortium (2002) ‘A nomenclature system for the tree of human Y-chromosomal binary haplogroups’ Genome Research 12: 339–48.

  42. Eastern Indonesia, Australia, and New Guinea: Haplotype 48 to eastern Indonesia and New Guinea, and Haplotype 49 to Australia; around the Indo-Pacific coast to Japan: Haplotype 50; a unique progenitor Asian son: M217 or Haplotype 52, giving rise to Haplotypes 51 and 53 – all haplotypes in Underhill, P.A. et al. (2001) ‘The phylogeography of Y-chromosome binary haplotypes and the origins of modern human populations’ Annals of Human Genetics 65: 43–62. into Mongolia and Central Asia: Haplogroups 21–26 and 31–34, respectively, Karafet, T.M. et al. (1999) ‘Ancestral Asian source(s) of New World Y-chromosome founder haplotypes’ American Journal of Human Genetics 64: 817–31. into the Americas: ibid.; Underhill et al., op. cit.; Underhill, P.A. et al. (2000) ‘Y-chromosome sequence variation and the history of human populations’ Nature Genetics 26: 358–61.

  43. no further north than Korea: Karafet et al., op. cit. Mongolia and the Russian Altai: Data in ibid.; see also data in Bing Su et al. (1999) ‘Y-chromosome evidence for a northward migration of modern humans into Eastern Asia during the last ice age’ American Journal of Human Genetics 65: 1718–24.

  44. 12,127 Asians and Pacific islanders: M89 (defines Consensus group F/Seth) in Ke, Y. et al. (2001) ‘African origin of modern humans in East Asia: A tale of 12,000 Y chromosomes’ Science 292: 1151–2. This paper incidentally also shows that all these 12,127 Asians share the M168 Out-of-Africa Adam mutation as do 99.9% of other non-Africans. as several geneticists have suggested: The view that Seth came as a later exodus from Africa is elaborated in Underhill et al. (2001) op. cit. For Seth types, in relict beachcomber populations: See (a) M89 (consensus Group F) 23% and M95 (consensus Group O) 65% respectively in Orang Asli aboriginals of the Malay Peninsula – see data in Bing Su et al. (2000) ‘Polynesian origins: Insights from the Y chromosome’ Proceedings of the National Academy of Sciences USA 97: 8225–8; (b) M9 (consensus Group K including Subgroups O, L, and P) 100% in Greater Andamans, while Asian YAP+ (Cain or consensus Group D) 100% in other Andaman Islanders – see data in
Thangaraj, K. et al. (2002) ‘Genetic affinities of the Andaman Islanders, a vanishing human population’ Current Biology (published online 26 November); (c) M9, 98% in Australoid tribal groups in India: Chenchus and Koyas – see data in Kivisild et al. (2002) op. cit.; (d) M9, 35% in Australians and 94% in New Guinea Highlanders – see data in Kayser, M. et al. (2001) ‘Independent histories of human Y chromosomes from Melanesia and Australia’ American Journal of Human Genetics 68: 173–90.

  45. Seth represents a quarter of all Indian Y chromosomes and his sons most of the rest: For Seth see Fig. 1 and Table 1 in Hammer, M.F. et al. (2001) ‘Hierarchical patterns of global human Y-chromosome diversity’ Molecular Biology and Evolution 18(7): 1189–1203; for representatives of Seth’s sons, Haplotypes 19–24, see Fig. 1, ibid.

  46. Hammer et al. (2001), op. cit., Haplotypes 20–23, see also Fig. 1, ibid. See also Chapters 3 and 4. Consensus group F/Seth accounts for G–R in The Y Chromosome Consortium (2002) op. cit.

  47. 40 per cent of Y-chromosome types: Underhill et al. (2001) op. cit.; Hammer et al., (2001) op. cit. suggests that he was born in India very soon after the initial out-of-Africa dispersal: One estimate (using the ‘Phylogenetic method’) of the age of Krishna’s immediate ancestor line M89 (Seth) is 88,000 years; the age of a sub-branch (M17) of Polo in India is estimated at 51,200 years using this method. M17 later moved to Central Asia and Europe – see Table 3 in Kivisild, T. (2003) ‘Genetics of the language and farming spread in India’ in P. Bellwood and C. Renfrew (eds) Examining the Farming/Language Dispersal Hypothesis (McDonald Institute for Archaeological Research, Cambridge) pp. 215–22. Several [sons] are local to Pakistan and India: Haplotypes 90 and 91 in haplogroup defined by M11, in Underhill et al. (2000) op. cit.; Haplotypes 90 and 91 defined by M147 and M70, inibid. another is found only in Melanesia: Haplotypes 94–97 in haplogroup defined by M4G/M5T/M9G, in Kayser et al., op. cit. and in Capelli, C. et al. (2001) ‘A predominantly indigenous paternal heritage for the Austronesian-speaking peoples of insular Southeast Asia and Oceania’ American Journal of Human Genetics 68: 432–43. another (TAT) is exclusive to Central Asia: TAT, Haplotype 92, Underhill et al. (2000) op. cit.

  48. M175 or Consensus type O . . . Ho: M175 branch in Underhill et al. (2000, 2001) op. cit. O/Ho corresponds to Haplogroup O in The Y Chromosome Consortium (2002) op. cit. Ho splits easily into three branches . . .; One remained in southern China, Indo-China and Southeast Asia: M95; southern China . . . concentrating on Taiwan: M119; Japan, Korea, and Northeast Asia: M122 – see data in Bing Su et al. (1999, 2000) op. cit.; Karafet et al., op. cit.; Underhill et al. (2000) op. cit.

  49. the other major Asian son of Krishna – Polo: the M45 branch in Underhill et al. (2000) op. cit. P/Polo corresponds to Haplogroup P in: The Y Chromosome Consortium (2002) op. cit. Kets and Selkups: Karafet et al., op. cit.

  50. Soffer, O. and Praslov, N.D. (eds) (1993) From Kostenki to Clovis: Upper Paleolithic – Paleo-Indian Adaptations (Plenum Press, New York).

  Chapter 6

  1. from the Russian Altai . . . through Lake Baikal in southern Siberia to the Aldan River in the east: Klein, R.G. (1999) The Human Career: Human Biological and Cultural Origins (Chicago University Press) – there is an excellent map of Upper Palaeolithic sites on p. 536. For the Ikhine II, Ust’ Mil’ (eastern Siberia) and Malaia Syia (Altai) sites, see Velichko, A.A. and Kurenkova, E.I. (1990) ‘Environmental conditions and human occupation of northern Eurasia during the Late Valdai’; in C. Gamble and O. Soffer (eds) The World at 18,000 bp, Vol. 1 (Unwin Hyman, London) pp. 254–65; Goebel, T. et al. (1993) ‘Dating the Middle-to-Upper-Palaeolithic transition at Kara Bom’ Current Anthropology 34: 452–8. For the Lake Baikal area, see Goebel, T. and Aksenov, M. (1995) ‘Accelerator radiocarbon dating of the initial Upper Palaeolithic in southeast Siberia’ Antiquity 69: 349–57. the Arctic Circle was penetrated: Pavlov, P. et al. (2001) ‘Human presence in the European Arctic nearly 40,000 years ago’, Nature 413: 64–7; see also Velichko and Kurenkova op. cit. on a northern bend of the Yellow River: at Salawasu/Shuidonggou, Chen, C. and Olsen, J.W. (1990) ‘China at the Last Glacial Maximum’ in C. Gamble and O. Soffer (eds) The World at 18,000 bp, Vol. 1 (Unwin Hyman, London) pp. 276–95.

  2. first flowering of the mammoth culture: Soffer, O. (1993) ‘Upper Paleolithic adaptations in Central and Eastern Europe and man–mammoth interactions’ in O. Soffer and N.D. Praslov (eds) From Kostenki to Clovis: Upper Paleolithic Adaptations (Plenum, New York) pp. 31–49. first possible evidence of Mongoloid features: Alekseev, V. (1998) ‘The physical specificities of Paleolithic hominids in Siberia’ in A.P. Derev’anko (ed.) The Paleolithic of Siberia: New Discoveries and Interpretations (University of Illinois Press, Urbana) pp. 329–35.

  3. See e.g. the description in Oppenheimer, S.J. (1998) Eden in the East: The Drowned Continent of Southeast Asia (Weidenfeld & Nicolson, London) pp. 23–7.

  4. Zhang, D.D. and Li, S.H. (2002) ‘Optical dating of Tibetan human hand- and footprints: An implication for the palaeoenvironment of the last glaciation of the Tibetan Plateau’ Geophysical Research Letters 29 (published online DOI: 10.1029/2001GL013749).

  5. The first [refuge] . . . characterized by . . . the Solutrean culture: Otte, M. (1990) ‘The northwestern European Plain around 18,000 bp’; Chapter 3, Gamble, C. and Soffer, O. (eds) (1990) The World at 18,000 bp, Vol. 1 (Unwin Hyman, London), pp. 61–5; for more details of Solutrean in SW Europe see also Chapters 2, 4–6. pp. 40–169. other southern refuges . . . described more generally as Epi-Gravettian: Otte, M. (1990) in Gamble and Soffer op. cit. second refuge area was Italy: ibid.; Mussi, M. (1990) ‘Continuity and change in Italy at the Last Glacial Maximum’ in Gamble and Soffer op. cit. pp. 126–43. third was the Ukraine: Gamble and Soffer op. cit, and Chapters 3, 7, 10–12; Soffer (1993) op. cit. Two other regions of Central Europe: Kozlowski, J.K. (1990) ‘Northern Central Europe c.18,000 bp’ in Gamble and Soffer op. cit. pp. 204–27.

  6. Soffer (1993) op. cit.

  7. Torroni, A. et al. (1998) ‘mtDNA analysis reveals a major late Paleolithic population expansion from Southwestern to Northeastern Europe’ American Journal of Human Genetics 62: 1137–52; Torroni, A. et al. (2001) ‘A signal, from human mtDNA, of postglacial recolonization in Europe’ American Journal of Human Genetics 69: 844–52.

  8. the post-glacial dates of expansion of V: Torroni et al. (1998) op. cit. The high frequency of V in the Saami is thought to be a founder effect. See also Richards, M. et al. (2000) ‘Tracing European founder lineages in the Near Eastern mtDNA pool’ American Journal of Human Genetics 67: 1251–76. Pre-V found further east (and Trans-Caucasus) . . . and older: Torroni et al. (2001) op. cit. Exactly the same [geographic] pattern . . . for Ruslan: Semino, O. et al. (2000) ‘The genetic legacy of Paleolithic Homo sapiens sapiens in extant Europeans: A Y-chromosome perspective’ Science 290: 1155–9. Note in this context, Ruslan in Semino is ‘Eu 18’ – i.e. M45/M173 without the further M17 (or R without R1a1 in consensus nomenclature)

  9. persisting preglacial mtDNA lines: Table 5 in Richards et al., op. cit. Note the LGM partition comes between lines 4 (Middle Upper Palaeolithic) and 3 (Late Upper Palaeolithic) of the table. a feature of the Ukraine refuge: ibid.

  10. Table 4 in Richards et al. (2000) op. cit.

  11. they still mark a clear genetic boundary: Stefan, M. et al. (2001) ‘Y-chromosome analysis reveals a sharp genetic boundary in the Carpathian region’ European Journal of Human Genetics 9: 27–33. M17 is still found at high frequencies: Later post-glacial expansions into that region could have had the same effect – Semino et al., op. cit. (M17 is Eu 19 in Semino et al.) In the consensus nomenclature, M17 would now be called R1a1 – see The Y Chromosome Consortium (2002) ‘A nomenclature system for the tree of human Y-chromosomal binary haplogroups’ Genome Research 12: 339–48.

  12. population of southern Central Asia . . . severely reduced: Davis, R.S. (1990) ‘Central Asian hunter-gatherers at the Last Glacial Maximum’ in Gamble and Soffer op. cit. pp. 267–75; but se
e also signs of life in Tibet at the LGM in Zhang and Li op. cit. human activity north even of the permafrost line . . . Afontova Gora . . . scattered archaeological sites: Velichko and Kurenkova op. cit.

  13. Table 2 in Forster, P. et al. (2003) ‘Asian and Papuan mtDNA evolution’ in P. Bellwood and C. Renfrew (eds) Examining the Farming/Language Dispersal Hypothesis (McDonald Institute for Archaeological Research, Cambridge) pp. 89–98.

  14. [?] no Mongoloid types . . . in East Asia until around 7,000–10,000 years ago: Brown, P. (1999) ‘The first modern East Asians? Another look at Upper Cave 101, Liujiang and Minatogawa 1’ in K. Omoto (ed.) Interdisciplinary Perspectives on the Origins of the Japanese (International Research Center for Japanese Studies, Kyoto) pp. 105–30. [?]none in Southeast Asia until well after that: Bellwood, P. (1997) Prehistory of the Indo-Malaysian Archipelago revised edn (University of Hawaii Press, Honolulu pp. 70-95).

  15. B and F . . . great local antiquity in the south: See discussion in Chapter 5. For pre-glacial age of B in Southern Mongoloids, see: Taiwan (B4a: 30,500 years) in Table 1, Richards, M. et al. (1998) ‘MtDNA suggests Polynesian origins in Eastern Indonesia’ American Journal of Human Genetics 63: 1234–6. See also discussion and summary re Southern origin of B and R9, and estimated ages of B (and sub-groups) and R9 (F4) in Table 3Yong-Gang Yao et al. (2002) ‘Phylogeographic differentiation of mitochondrial DNA in Han Chinese’ American Journal of Human Genetics 70: 635–51.

  16. Oppenheimer op. cit.

  17. Ibid.; see also note 22 below and Bellwood op. cit.

  18. Largest population expansion in ISEA . . . arrival of the Metal Age: The metal age arrived much later in Island Southeast Asia than on the mainland – see Higham, C. (1996) The Bronze Age of Southeast Asia (Cambridge University Press) pp. 301–4. When rice agriculture greatly expanded: Paz, V. (2003) ‘Island Southeast Asia: Spread or friction zone?’ in P. Bellwood and C. Renfrew (eds) Examining the Farming/Language Dispersal Hypothesis (McDonald Institute for Archaeological Research, Cambridge) pp. 275–86. And Bulbeck F.D. (2002) ‘Recent Insights on the Chronology and Ceramics of the Kalumpang Site Complex, South Sulawesi, Indonesia’ Indo-Pacific Prehistory Association Bulletin 22 (Vol 6): 83–99.