Out of Eden: The Peopling of the World (62 page)

BOOK: Out of Eden: The Peopling of the World
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27.
Middle Palaeolithic tradition
: Blackwell, B.A.B. et al. (1998) ‘ESR (Electron Spin Resonance) dating the Palaeolithic site at Tsagaan Agui, Mongolia’, Proceedings of the 31st International Symposium on Archaeometry, 27 April–1 May 1998, Budapest.
Upper Palaeolithic (microlithic) traditions
: Tang Hui Sheng (1995) ‘Lithic industries of the Qinghai-Tibetan Plateau’
The Artefact
18
: 3–11.

28.
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-Paleo-Indian Adaptations
(Plenum Press, New York) pp. 33–49.

29.
Cavalli-Sforza, L. et al. (1994)
The History and Geography of Human Genes
(Princetown University Press, Princetown) pp. 223–38.

30.
Chan Wing-hoi (1995) ‘Ordination names in Hakka genealogies: A religious practice and its decline’ in D. Faure and H. Siu (eds)
Down to Earth: The Territorial Bond in South China
(Stanford University Press, Stanford) pp. 65–82. The myth of a homogenous mix throughout China is still promoted for whatever reason (see e.g. Yuan-Chun Ding et al. (2000) ‘Population structure and history in East Asia’
Proceedings of the National Academy of Sciences USA
97
: 14003–6) and then demolished, sometimes by the same authors (see e.g. Yong-Gang Yao et al., op. cit.

31.
Su, B. et al. (1999) ‘Y-chromosome evidence for a northward migration of modern humans into East Asia during the last ice age’
American Journal of Human Genetics
65
: 1718–24.

32.
palaeoclimatological studies
: see the maps at
http://www.esd.ornl.gov/projects/qen/NEW_MAPS/eurasia1.gif
and
http://www.esd.ornl.gov/projects/qen/euras(2.gif
deep genetic diversity of south-west Central Asia
: see Fig. 2 in Metspalu, E. et al. (1999) ‘The Trans-Caucasus and the expansion of the Caucasoid-specific mitochondrial DNA’ in S.S. Papiha et al. (eds)
Genomic Diversity: Applications in Human Population Genetics
(Kluwer Academic/Plenum, New York) pp. 121–34. See specific maps on Jonathan Adams’ ESD ORNL reference website:
www.esd.ornl.gov/projects/qen/euras18k.gif
and
www.esd. ornl.gov/projects/qen/euras(2.gif
. See also the archaeological evidence presented in the next chapter.

33.
Groups A and X are first-generation from N/Nasreen, while B and F derive from Nasreen through Rohani. Groups C, D, E, G, and Z all derive directly from M/Manju (see
Figures 5.9
and 5.10). The first full description of Groups A to G all shown on one tree was by Torroni, A. et al. (1994) ‘Mitochondrial DNA analysis in Tibet: Implications for the origins of the Tibetan population and its adaptaton to high altitude’
American Journal of Physical Anthropology
93
: 189–99. For the first naming of Group X, see Forster, P. et al. (1996) ‘Origin and evolution of Native American mtDNA variation: A reappraisal’
American Journal of Human Genetics
59
: 935–45. A further group, Y, appears almost exclusively among the Nivkhs of Sakhalin Island in the north; it is a very young group, and therefore peripheral to this discussion (see below). For Groups Y and Z and the most recent published updates of the Asian mtDNA tree, see Kivisild, T. et al. (2002) ‘The emerging limbs and twigs of the East Asian mtDNA tree’
Molecular Biology and Evolution
19
(10): 1737–51; Yong-Gang Yao et al., op. cit.; and Fig. 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.

The first task is to identify distinct northern and southern genetic lines among Mongoloid populations. C and Z are both related M types (on newly identified parent group M8, Yong-Gang Yao et al., op. cit.), and are found almost entirely in North Asia (and C also in the Americas), and not in Southern Mongoloids. A, although common in north-east China, north-east Siberia, and North America (Kolman, C.J. et al. (1996) ‘Mitochondrial DNA analysis of Mongolian populations and implications for the origin of New World founders’
Genetics
142
: 1321–34) is also found at appreciable frequencies and diversity in south and south-west China, Tibet, and Xinjiang (Yong-Gang Yao et al., op. cit.), but is absent from India and West Eurasia (Kivisild, T. et al. (1999) ‘The place of the Indian mitochondrial DNA
variants in the global network of maternal lineages and the peopling of the Old World’ in S.S. Papiha et al. (eds)
Genomic Diversity: Applications in Human Population Genetics
(Kluwer Academic/Plenum, New York), pp. 135–52). So, a south-west China source, with movement up the Yangtzi to Tibet and thence to Central and North Asia, seems more likely for the A group origins than either an Indian or North Asian source. Y, a daughter of N, is found only in Northeast Asia (Stariovskaya, Y.B. et al. (1998) ‘mtDNA diversity in Chukchi and Siberian Eskimos: Implications for the genetic history of Ancient Beringia and the peopling of the New World’
American Journal of Human Genetics
63
: 1473–91; Torroni, A. et al. (1993) ‘mtDNA variation of aboriginal Siberians reveals distinct genetic affinities with Native Americans’
American Journal of Human Genetics
53
: 591–608). But Y has an age of only around 5,000 years, and is ultimately derived from a Southeast Asian N branch, N9. X, another daughter of N, is limited to Europeans and Americans (Brown, M. et al. (1998) ‘mtDNA Haplogroup X: An ancient link between Europe/Western Asia and North America?’
American Journal of Human Genetics
63
: 1852–61), and there is one reported instance in Siberians (see below). Although any of these three northern lines (C, X, and Z) could theoretically be characteristic Mongoloid genetic lines, they are absent from Southern Mongoloids, and also are found in other populations, thus making this less likely. Several more Asian clades found throughout China have recently been added: three are from M (M7, M9 and M10) and a fourth is from N (N9, estimated age 64,300 ± 20,000 years and ancestral to Group Y): see Yong-Gang Yao et al., op. cit.

The two Asian granddaughters of N (B and F) are the only known East Asian offspring of R/Rohani (except for a newly described but ancient clade, R9, estimated age 81,000 years ± 24,600years: see Yong-Gang Yao et al., op. cit., which has been shown to be ancestral to F in the south and thus has already been relabelled F4 in: Hill, C. et al. (2003) ‘Mitochondrial DNA variation in the Orang Asli of the Malay Peninsula’ (in preparation); See also
Fig. 5.6
). B and F have their greatest diversity and frequency in Southern Mongoloids of Southeast Asia (Torroni et al. (1994) op. cit.; Yong-Gang Yao et al., op. cit.). They are also both found north of the Himalayan barrier in Central Asia (Fig. 2 in Metspalu et al., op. cit), Mongolia (Kolman et al., op. cit.), and Tibet, Korea, and Japan (Torroni et al. (1994) op. cit.; Horai, S. et al. (1996) ‘mtDNA polymorphism in East Asian populations’
American Journal of Human Genetics
59
: 579–90). They are not, however, found farther north in Siberia in the Subarctic region. Of these southern lines, B is also found in America, although again below the Subarctic region (see
Chapter 7
). Since B and F are present on both sides of the Himalayas, they are potential candidate original Mongoloid lines, but there is still a question mark over whether they define all Mongoloids (F is absent from America).

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. .

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