Out of Eden: The Peopling of the World (57 page)

17.
half of all western and northern European maternal lines
: Table 2 in Kivisild et al. (1999) op. cit.
The expansion of HV has been dated to 33,500 years ago
: Richards et al. (2000) op. cit.

18.
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 Publishers, New York 1999) pp. 121–134.

19.
Inos, after Seth’s son Enos
: This consensus haplogroup I (Inos) clan is uniquely identified by bi-allelic marker M170 (haplotypes 49–53) in Underhill et al., op. cit., is synonymous with groups Eur7 and 8 in Semino et al., op. cit., and is largely overlapping with Tyler-Smith’s Haplogroup 2 (Rosser et al., op. cit.). For an explanation of the nomenclature of I (Inos), see The Y Chromosome Consortium op. cit.
According to the Leicester-based geneticist Zoë Rosser
: Rosser et al., op. cit.
Ornella Semino and her colleagues
: Semino et al., op. cit.

20.
earliest changes from Mousterian
: 46,000–47,000 years ago – Bar-Yosef, O. (1994) ‘The contributions of Southwest Asia to the study of the origins of modern humans’ in M.H. Nitecki and D.V. Nitecki (eds)
Origins of Modern Humans
(Plenum Press, New York, 1994),
Chapter 2
. These are radiocarbon dates, and – as stated by Bar-Yosef – if correction were possible, could be much older (e.g. as much as 51,000 years ago – see comments on radiocarbon ceiling, note 7 above.) The rock shelter of Ksar ‘Aqil in southern Lebanon has also been suggested as a site of the earliest transition from Mousterian technology to Levantine Aurignacian, possibly (unfortunately the date of the transition of culture could only be inferred stratigraphically) between 50,000 and 52,000 years ago; see Mellars, P. and Tixier, J. (1989) ‘Radiocarbon-accelerator dating of Ksar ‘Aqil (Lebanon) and the chronology of the Upper Palaeolithic sequence in the Middle East’
Antiquity
63
: 761–8.
hiatus coincided with a climatic worsening
: The period spans two warm, wet interstadials, IS 12 and 11, with cold dry snaps preceding each of these. (IS numbering and dates according to Dansgaard op. cit) The first cold event coincided with Heinrich event No. 5. (See also Schultz et al., op. cit.)

21.
The progressively worsening cold spell spanned the whole period between (IS) 12 (43,500 years) and IS 8 (34,000 years), the latter IS signalling the onset of re-warming. (Interstadial numbering and dates according to Dansgaard op. cit.) The first cold event coincided with Heinrich event No. 5. See also Schultz et al., op. cit.

22.
For details of the Shanidar burial and the Baradostian culture, see Solecki, R. (1972)
Shanidar: The Humanity of Neanderthal Man
(Allen Lane, London).

23.
Table 6.1 in Gamble op. cit.

24.
Earliest Upper Palaeolithic, 33,000–45,000 years ago
: Table 6.5 in Gamble op. cit. Occupation peaks for this phase occur between 40,000 and 44,000 years ago (calibrated
¹⁴
C dates, ibid. p. 285).
Cultures taking off from around 30,000 years ago
: Table 6.5 ibid.
third phase of high occupation
: These (the Gravettian technocomplex) correspond with a third occupation peak (or Middle Upper Palaeolithic) between 25,000 and 29,000 years ago (uncalibrated dates, Fig. 6.5 ibid.).
33,000 years ago, for example at Kostenki
: ibid. pp. 287–92.

25.
cultural additions innovations of the Gravettian
: Gamble op. cit. pp. 287–92, esp. p. 290; Soffer, O. (1993) ‘Upper Paleolithic adaptations in Central and Eastern Europe and man-mammoth interactions’ in O. Soffer and N. Praslov (eds)
From Kostenki to Clovis: Upper Paleolithic-Paleo-Indian Adaptations
(Plenum, New York) pp. 31–49.
may also have represented an intrusion of peoples carrying the seeds of such cultural practices from eastern Europe
: Otte, M. (2000) ‘The history of European populations as seen by archaeology’ in C. Renfrew and K. Boyle (eds)
Archaeogenetics: DNA and the Population Prehistory of Europe
(MacDonald Institute for Archaeological Research, Cambridge) pp. 139–41.

26.
Torroni, A. et al. (2001) ‘A signal, from human mtDNA, of postglacial recolonization in Europe’
American Journal of Human Genetics
69
: 844–52.

27.
perfectly preserved Caucasoid mummies
: The point about the 3,000-year-old Caucasoid mummies on the Silk Road is merely to emphasise that there is no
a priori
reason to assume that the first populations of this part of Central Asia were Mongoloid: Barber, E.W. (2000)
The Mummies of Urumchi
(Pan, London); Mallory, J.P. and Mair, V. (2000)
The Tarim Mummies: Ancient China and the Mystery of the Earliest Peoples from the West
(Thames & Hudson, London).
two sites in the Russian Altai
: Otte, M. and Derevianko, A. (2001) ‘The Aurignacian in Altai’
Antiquity 75
: 44–8; Goebel, T. et al. (1993) ‘Dating the Middle-to-Upper-Palaeolithic transition at Kara Bom’
Current Anthropology
34
: 452–8; Goebel, T. and Aksenov, M. (1995) ‘Accelerator radiocarbon dating of the intial Upper Palaeolithic in Southeast Siberia’
Antiquity
69
: 349–57.

28.
Half of these consist of HV stock
: Metspalu et al., op. cit.; Kivisild et al. (1999) op. cit.
recent eastward European emigration
: Comas, D. et al. (1998) ‘Trading genes along the Silk Road; mitochondrial DNA sequences and the origin of Central Asian populations’
Molecular Biology and Evolution
13
: 1067–77.
absent from Central Asia
: although there are matches for most European H founders.
most of the other ‘west Eurasian Nasreen lines’
: U, J, and T haplogroups – U2i, U7, U5a, 1a, U4, U1, and K; see Fig. 2 in Metspalu et al., op. cit.; Kivisild et al. (1999) op. cit.
HV could have originally come from South Asia
: and J1, U1, and T.

29.
Dates of branches vary enormously
: e.g. Kivisild (2003b) op. cit.; Forster, P. et al. (2000) ‘A short tandem repeat-based phylogeny for the human Y chromosome’
American Journal of Human Genetics
67
: 182–96.
careful analysis of founder lines and mtDNA dating
: Richards et al. (2000) op. cit.; Metspalu et al., op. cit.; Kivisild et al. (1999) op. cit.

30.
Underhill et al., op. cit.; Semino et al., op. cit.

31.
The root haplotype 87 defined by M9 in Underhill et al., op. cit. This clade has been reclassified as K/Krishna (while M89 is F/Seth): see The Y Chromosome Consortium op. cit.

32.
the Oxford-based geneticist
: Zergal, T. et al. (1997) ‘Genetic relationships of Asians and Northern Europeans, revealed by Y-chromosomal DNA analysis’
American Journal of Human Genetics
60
: 1174–83.
his genetic father and grandfather
: haplotypes 87 and 71 respectively, defined by M9 and M89 in Underhill et al., op. cit.
a migration from Central Asia
: possibly only since the last glaciation – see Rosser et al., op. cit.; Semino et al., op. cit.

33.
Hungarians achieve the highest frequency
: followed closely by Poland, the Ukraine, and Russia at 56, 54, and 47%, respectively: Semino et al., op. cit.; Rosser et al., op. cit.
especially the Altai
: 52%, Quintana-Murci et al., op. cit.; Underhill et al., op. cit.

34.
the ultimate origin of M17
: Kivisild et al. (2003a,b) op. cit.
highest rates and diversity of the M17 line in Pakistan, India, and eastern Iran, and low rates in the Caucasus
: respectively 32, 20, 31, and 2% – Quintana-Murci et al., op. cit.; Rosser et al., op. cit. For a tabulation of the relative diversity of M17 (R1a) showing the highest diversity in South Asia, particularly Iran, see Table 5 in Kivisild et al. (2003a) op. cit.
36,000 years
: measured at 35,700 years – Kivisild (2003b) op. cit.; other methods of calculation yield much lower estimates.

35.
movement from the east to the west 30,000 years ago
: Semino et al., op. cit. Phylogenetic analysis produces a realistic age, for M17 in Europe, of 27,000 years, see: Kivisild (2003b) op. cit. In contrast, Quintana-Murci et al. (op. cit.) tentatively suggests that the expansion into Europe started only around 5,000 years ago with the advent of farming.
Ruslan
: M173 root type, i.e. Haplotype 104, in Underhill et al., op. cit. Reclassified as ‘R’ in: The Y Chromosome Consortium (2002) op. cit.

36.
according to one study
: M173/ht 104, in Underhill et al., op. cit.
Ruslan’s genetic father, P
: Haplotype 111 in ibid.; reclassified as P in The Y Chromosome Consortium op. cit.

37.
Semino et al., op. cit. It should be noted that the coalescent estimate for M17, M173, and M172 at M89 (Underhill haplotype 71) in India may be as old as 88,300 years – estimated by phylogenetic analysis, see Kivisild (2003b) op. cit.

Chapter 4

1.
For a full discussion of the phylogeographic approach and founder analysis see Richards, M. et al. (1998) ‘Phylogeography of mitochondrial DNA in western Europe’
American Journal of Human Genetics
67
: 241–60; Richards, M. et al. (2000) ‘Tracing European founder lineages in the Near Eastern mtDNA pool’
American Journal of Human Genetics
67
: 1251–76.

2.
much more recent African admixture
: Richards, M. et al. (2003) ‘Extensive female-mediated gene flow from sub-Saharan Africa into Near Eastern Arab populations’
American Journal of Human Genetics
72
: 1058–64.
a number of other South Asian aboriginal groups
: India also has so-called proto-Asian groups, such as Maria Gond, Khonda Dora, and Kattuniaken. Some of these are probably, like the Munda/Mundari groups, more recent east-west re-entrants from Indo-China: see Figs 14 and 21 in Oppenheimer, S.J. (1998)
Eden in the East
(Weidenfeld & Nicolson, London). For an anthropological/genetic description of the Indian aboriginal groups see also Watkins, W.S. et al. (1999) ‘Multiple origins of the mtDNA
9-bp deletion in populations of South India’
American Journal of Physical Anthropology
109
: 147–58; Watkins, W.S. et al. (2001) ‘Patterns of ancestral human diversity: An analysis of Aluinsertion and restriction-site polymorphisms’
American Journal of Human Genetics
68
: 738–52.
beach-settling ancestors from Africa
: Kivisild. T. et al. (2003) ‘The genetic heritage of the earliest settlers persists both in Indian tribal and caste populations’
American Journal of Human Genetics
72
: 313–32.

3.
two ancient and unique Indian Manju clans, M2 and M4
: for these groups in Andamanese, see Endicott, P. et al. (2003) ‘The genetic origins of the Andaman Islanders’
American Journal of Human Genetics
72
: 178–84; Thangaraj, K. et al. (2003) ‘Genetic affinities of the Andaman Islanders, a vanishing human population’
Current Biology
published online, 26 November 2002.
commonest mtDNA component among the Indian aboriginal groups
: Kivisild et al., op. cit.
On the paternal side
: for Y types in Andamanese, see Thangaraj et al., op. cit.

4.
Pre-F1: Hill, C. et al. (2003) ‘Mitochondrial DNA variation in the Orang Asli of the Malay Peninsula’ (in preparation). For similar Pre-F1 haplotypes in Andamanese and Nicobars, see Andaman haplotypes 9 and 10 and Nicobar haplotype 1 in: Thangaraj et al., op.cit.; Prasad, B.V. et al. (2001). ‘Mitochondrial DNA variation in Nicobarese Islanders’
Human Biology
73
: 715–25.

5.
Nasreen and Manju mtDNA lines in New Guinea and Australia (see also Fig. 8.2 in Appendix 1): New Guinea has three main mtDNA clades, labelled PNG 1–3 in Redd, A.J. and Stoneking, M. (1999) ‘Peopling of Sahul: mtDNA variation in Aboriginal Australian and Papua New Guinean populations’
American Journal of Human Genetics
65
: 808–28. The name labels of these three clades in the above paper corresponding to the nomenclature used in this book (Table 8.1 in Appendix 1) are as follows. PNG1 = Haplogroup B (of R haplogroup); PNG2 = local subgroup (of R haplogroup, also Group P 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); PNG3 = Q in Forster et al. (2003) op. cit., and M11 in Richards, M. and Macaulay, V. (2000) ‘Genetic data and colonization of Europe: Genealogies and founders’ in C. Renfrew and K. Boyle (eds)
Archaeogenetics: DNA and the Population Prehistory of Europe
(McDonald Institute for Archaeological Research, Cambridge) pp. 139–41, Fig. 14.1. These three clades can also be confirmed unambiguously using typing data from several other publications, e.g. PNG2 = Haplotypes 6–8 and PNG3 = Haplotype 22, in Ingman, M. et al. (2000) ‘Mitochondrial genome variation and the origin of modern humans’
Nature
408
: 708–13; PNG1 = Haplogroup 1 and PNG3 = Haplogroup 2, in Sykes, B. et al. (1995) ‘The origins of the Polynesians: An intepretation from mitochondrial lineage analysis’
American Journal of Human Genetics
57
: 1463–75. For further Papuan details, see also Forster et al. (2003) op. cit.