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Ancient DNA analysis of human neolithic remains found in northeastern Siberia.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 126:458 – 462 (2005)
Ancient DNA Analysis of Human Neolithic Remains
Found in Northeastern Siberia
François-Xavier Ricaut,1,2* A. Fedoseeva,3 Christine Keyser-Tracqui,1 Eric Crubézy,2 and
Bertrand Ludes1,2
1
Laboratoire d’Anthropologie Moléculaire, Institut de Médecine Légale, 67085 Strasbourg, France
Laboratoire d’Anthropobiologie, Université Paul Sabatier, CNRS, UMR 8555, 3100 Toulouse, France
3
Department of Archeology and Ethnography, Yakutsk State University, Yakutsk 677891, Russia
2
KEY WORDS
ancient DNA; HV1 sequence; STRs; north Siberia
ABSTRACT
We successfully extracted DNA from a
bone sample of a Neolithic skeleton (dated 3,600 ⫾ 60
years BP) excavated in northeastern Yakutia (east Siberia). Ancient DNA was analyzed by autosomal STRs (short
tandem repeats) and by sequencing of the hypervariable
region I (HV1) of the mitochondrial DNA (mtDNA) control
region. The STR profile, the mitochondrial haplotype, and
The cultural, morphological, and genetic similarities observed between Siberian and Native American populations led to the consensus that the ancestral population(s) of Native Americans migrated
from Siberia through Beringia during the last glaciation (30,000 –12,000 years BP) (Cavalli-Sforza et
al., 1994; Crawford, 1998). Nevertheless, the numerous genetic studies analyzing the relationship between aboriginal Siberian and Native American
populations do not agree on the timing, source population(s), and number of waves of migration.
The difficulties in retracing the initial peopling of
the Americas arise from the many historic and demographic events that have occurred since the initial colonization, and which could have obscured the
ancestral Asian gene pool of Amerindians (Forster et
al., 2001; Malhi et al., 2002). Indeed, searching the
genetic traces of ancestral Siberian population(s) of
Native Americans among the Siberian modern gene
pool is a wager because the Siberian gene pool has
been affected by important changes since the first
migrants left for the Americas. The inhospitable
conditions of Siberia, in addition to a low human
density, favored a high degree of Siberian population isolation and genetic drift (Santos et al., 1999).
The depopulation of northern Asia during the last
glacial maximum (around 20,000 years ago) and its
repopulation at the end of the glacial phase 11,000
years ago (Forster et al., 2001) could also have
played an important role in these changes. More
recently, Russian colonization, which began in the
17th century, led to the extinction of several ethnic
groups and to a drastic reduction in the size and
©
2004 WILEY-LISS, INC.
the haplogroup determined were compared with those of
modern Eurasian and Native American populations. The
results showed the affinity of this ancient skeleton with
both east Siberian/Asian and Native American populations. Am J Phys Anthropol 126:458 – 462, 2005.
©
2004 Wiley-Liss, Inc.
genetic diversity of Siberian populations (Levin and
Potatov, 1964; Forsyth, 1996).
Only direct access to the gene pool of ancient aboriginal Siberian populations could clarify 1) northern Asian prehistory, 2) the ancestor-descendant
relationships between ancient and modern populations, and 3) the colonization of the New World,
independent of historical changes affecting the Siberian gene pool. Ancient DNA data from south Siberian populations were previously published (Clisson et al., 2002; Keyser-Tracqui et al., 2003; Ricaut
et al., 2003), but to the best of our knowledge, there
are no ancient DNA data of northeastern Siberians
available.
MATERIALS AND METHODS
During summer 1980, the “Prilenskoy” Russian
expedition discovered in northeastern Siberia (in the
Sakha Republic), near the Panteleikhe River (lower
Grant sponsor: French Department of Research; Grant sponsor:
ACI “Espaces et Territoires: Le complexe spatial Altaı̈-Baı̈kal. Plaque
tournante des flux géniques en Haute Asie de la période protohistorique à l’époque moderne.”
*Correspondence to: François-Xavier Ricaut, Department of Biological Anthropology, Leverhulme Centre of Human Evolutionary Studies, University of Cambridge, Downing Street, Cambridge CB23DZ,
UK. E-mail: fx.ricaut@infonie.fr
Received 23 July 2003; accepted 31 October 2003.
DOI 10.1002/ajpa.20257
Published online 13 August 2004 in Wiley InterScience (www.
interscience.wiley.com).
459
GENETIC ANALYSIS OF A NEOLITHIC SIBERIAN
Kolyma River basin), a frozen Neolithic grave. This
tomb (radiocarbon-dated at 3,600 ⫾ 60 years BP by
classic analysis of bone artifacts associated with the
skeleton) contained a very well-preserved human
skeleton of a young woman (20 –25 years old) with
indications of Asian-ancestry skull traits (Gokhmana and Tomtosovoy, 1983). Based on archaeological data, this woman could not be affiliated with
known recent or ancient Siberian ethnic groups
(Kistenev, 1992).
During each step of sample preparation (abrasion,
extractions, and amplifications), standard precautions were taken to minimize the risk of contamination and to detect any potential contamination. The
outer surface of the ancient bone fragment was removed to almost 3– 4-mm depth to eliminate possible surface contamination. Extraction and amplification were performed in a dedicated ancient DNA
laboratory, routinely sterilized by different treatments (DNAse away威, bleach and ultraviolet light
irradiation at 254 nm), wearing full body protective
clothing, and using dedicated equipment and reagents. Extraction and amplification blanks were
used as negative controls, and mitotypes and genetic
profiles of all persons involved in processing samples
were determined (data not shown) and compared to
results obtained from the ancient bone sample.
Moreover, DNA was extracted at least three times,
and at least three PCR amplifications were made
from each extract to assess the reproducibility of
results.
In this study, a femoral bone fragment from this
ancient female skeleton was used to extract ancient
DNA according to published protocols (Fily et al.,
1998). The aqueous phase was purified, using a
CleanMix Kit (Talent, France), and concentrated to
40 ␮l, using Microcon威-30 filters (Millipore, France).
Mitochondrial DNA analyses were performed on hypervariable region 1 of the mtDNA control region
(HV1). This region was divided into two subregions
(a and b) amplified with two sets of overlapping
primer pairs, L15989/H16239 5⬘-CCCAAAGCTAAGATTCTAAT-3⬘/5⬘-TGGCTTTGGAGTTGCAGTTG-3⬘
and L16190/H16410 5⬘-CCCCATGCTTACAAGCAAGT-3⬘/5⬘-GAGGATGGTGGTCAAGGGAC-3⬘.
DNA amplifications were performed in 50 ␮l of reaction mixture containing 2– 6 ␮l of the ancient DNA
extract, 10 mM Tris HCL (pH 8.3), 50 mM KCL, 1.5
mM MgCl2, 1 mg/ml BSA, 200 ␮M of each dNTP, 0.25
␮M of each primer, and 2.0 unit of Taq Gold Star
(Eurogentec). Cycling parameters were 94°C for 10
min, followed by 38 cycles of 94°C for 30 sec, 48°C for
30 sec for HV1a or 30 sec at 51°C for HV1b, 72°C for 45
sec, and 72°C for 5 min. Amplification products were
checked on a 1% agarose gel and purified with Microcon威-PCR filters (Millipore). Sequence reactions were
performed on each strand, with the same primers as
those used for PCR amplification, by means of the ABI
Prism BigDye威 Terminator cycle sequencing Ready
Reaction Kit (PE Applied Biosystems), according to
the manufacturer’s instructions. Sequence reaction
TABLE 1. mtDNA sequence (between bases 16015–16391) of
ancient Siberian sample
Polymorphic positions1
Sample
16223
16298
16327
CRS
Ancient Siberian
C
T
T
C
C
T
1
Numbered according to published Cambridge Reference Sequence (Anderson et al., 1981).
products were analyzed on the ABI Prism 3100 automatic sequencer (PE Applied Biosystems).
Autosomal short tandem repeats (STRs) were amplified using the AmpFlSTR威 Profiler Plus威 Kit (PE
Applied Biosystems). Nine STRs (D3S1358, vWA,
FGA, D8S1179, D21S11, D18S51, D5S818, D13S317,
and D7S820) and the amelogenin locus (determining
the individual’s sex) were simultaneously amplified.
Each amplification was carried out in 10 ␮l of a
reaction mixture containing 3.82 ␮l PCR reaction
mix, 2 ␮l primer set, 0.182 ␮l AmpliTaq Gold威 (PE
Applied Biosystems), and 1– 4 ␮l of the DNA extract.
Cycling parameters were 94°C for 11 min, followed
by 37 cycles of 94°C for 1 min, 59°C for 1 min, and
72°C for 1 min, and a final delay of 45 min at 60°C.
Both nuclear and mtDNA amplification products
were analyzed on an ABI Prism 3100 automatic
sequencer (PE Applied Biosystems).
RESULTS AND DISCUSSION
The mtDNA HV1 region analysis of the Yakut
specimen resulted in the recovery of a 377-base pair
(bp) fragment (nucleotide positions 16015–16391 of
the Cambridge Reference Sequence (CRS); Anderson et al., 1981), which was confirmed on both
strands (Table 1). This sequence was compared to
the CRS: there was a total of three variable nucleotide positions (np), all corresponding to transitions.
These mutations at np 16223 C-to-T, 16298 T-to-C,
and 16327 C-to-T correspond to substitutions which
are characteristic of the founding HV1 sequence for
haplogroup C in Siberian and Asian populations
(Torroni et al., 1993).
Analysis of the ancient Siberian haplogroup and
haplotype distribution in Eurasian populations
showed that they are both widespread in modern
Asian populations. They were most frequently found
in east and south Siberian populations (Tuvinian,
Buryat, Yakut, Evens, Koryak, and Chukchi) (Derenko and Shields, 1997; Starikovskaya et al., 1998;
Schurr et al., 1999; Pakendorf et al., 2003; Derenko
et al., 2003), and then to a lesser extent in east Asian
populations (Mongol, Chinese, and Korean) (Kolman
et al., 1996; Lee et al., 1997; Yao et al., 2002). The
above distribution and the presence in an ancient
northeast Siberian skeleton of an HV1 sequence presenting mutations at nucleotide positions 16223,
16298, and 16327 are in accordance with genetic
studies of modern populations which assert that this
HV1 haplotype is the founding HV1 sequence for
haplogroup C in Siberian and Asian populations
460
F.-X. RICAUT ET AL.
TABLE 2. Autosomal STR results obtained with Profiler Plus kit from ancient DNA sample1
Extraction
A
B
C
Consensus
genotype
1
Amelo
genine
D8S1179
D21S11
D7S820
D3S1358
D13S317
vWA
D18S51
D5S818
FGA
XX
XX
XX
XX
XX
XX
XX
XX
XX
14/14
14/14
14/(15)
14/14
(13)/14
14/14
14/14
14/14
14/14
31/(31)
31/32.2
(32.2)/32.2
31/32.2
31/32.2
31/(31)
31/32.2
(29)/31/32.2
31/32.2
8/8
8/8
8/(11)
8/(9)
8/(11)
8/8
8/(11)
15/15
(14)/15
15/(18)
15/15
15/(16)
15/15
15/15
15/15
15/15
8/9
8/9
8/9(11)
8/9
8/9
8/9
8/9
8/9
8/9
18/18
17/(18)
18/18
18/18
17/(18)
18/18
18/18
18/18
18/18
15/(17)
15/15
15/15
15/15
15/(17)
15/15
(14)/15
(14)/15/(17)
15/(?)
11/12
11/12
11/12
11/12
11/12
(10)/11/(13)
11/12
11/12
11/12
22/(23)
(20/24)
22/(23)
22/(24)
22/(23/24)
(23/25)
22/(24)
22/(22)
22/(?)
8/(?)
Alleles which could not be detected in at least five different amplifications are in parentheses.
(Torroni et al., 1993). Moreover, the fact that all east
Siberian mainland populations shared this ancient
haplotype with a relatively high frequency, without
restriction to any linguistic or ethnic group, 1) could
either be explained by a common ancestral population or by the effect of a significant genetic drift
favored by the high degree of Siberian population
isolation, and 2) allows us to postulate that the
Neolithic woman studied here might be considered
an ancestor of present-day inhabitants of east Siberia.
Moreover, haplogroup C (to which the ancient east
Siberian individual belongs) is considered one of
four major founding Native American haplogroups
(Torroni et al., 1993), and was also widespread in
Native American populations (Merriwether et al.,
1995; Malhi et al., 2002). However, the ancient east
Siberian haplotype was only to be found in two
southern Native Americans (one modern Chilean
individual, Horai et al., 1993; and one ancient Colombian mummy, Monsalve et al., 1996) and eight
northern Native Americans (seven ancient Norris
Farms Oneota individuals, Stone and Stoneking,
1998; and one modern Ponca individual, Malhi et al.,
2002). As suggested by Malhi et al. (2002), the share
of this haplotype in east Siberian/Asian and Native
American populations could be due to either convergence or common ancestry.
The genetic typing by megaplex amplifications
provided, after a combination of different amplification results, an incomplete consensus STR profile
(Table 2). Indeed, to reduce ancient DNA STR genotyping errors and ensure the reliability of results,
only the amplified products from each locus that
were reproducible in at least five different amplification reactions were considered authentic
(Schmerer et al., 1999). This strategy allowed identification of artifact alleles and false-homozygosity
resulting from sporadic contaminations or amplification artifacts (such as shadow bands and allelic
dropout) which were caused by a lack of DNA quality and/or quantity (Schmerer et al., 1999; Hummel
et al., 2000). Of the nine autosomal loci tested, only
six (vWA, D21S11, D8S1179, D5S818, D13S317, and
D3S1358) and the amelogenin locus gave clearly
reproducible results in five different amplification
reactions, and can be considered authentic (Table 2).
For three longer STR loci (D7S820, 258 –294 bp;
D18S51, 273–341 bp; and FGA, 219 –267 bp), the
degree of reproducibility of results was not sufficient
to provide a reliable profile and to assess with certainty the homozygous or heterozygous nature of the
genotype obtained.
The results obtained from AmpFlSTR威 Profiler
Plus威 Kit (PE Applied Biosystems) indicate that our
results were not influenced by contamination. Indeed, 1) the amplified products were reproducible
from multiple extractions and amplifications, 2) STR
autosomal amplification success correlated negatively with the length of the amplicons, 3) the morphological and molecular sex determinations of the
Siberian skeleton were in accordance with each
other, and 4) the ancient Siberian STR profile (even
incomplete) was never found to correspond to someone involved in processing samples. The nuclear
data obtained in this study attest to the authenticity
of the DNA extract and amplified products, and
prove that ancient DNA molecules were preserved in
the sample bone. The HV1 sequence obtained, which
was fully reproducible from multiple amplifications
and extractions and was different from that of the
French and Yakut staff, can thus be considered authentic.
An assignment method, based on analysis of allelic frequencies of the six STR loci considered in the
consensus genotype, was performed to investigate
the population affinities of the ancient skeleton, by
determining the populations in which the STR profile from the ancient skeleton was most likely to
occur (Cornuet et al., 1999). The probability of observing an individual with the ancient skeleton STR
profile was 10 times higher in Native Alaskan (Athabaskan, Inupiat, and Yupik; Budowle et al., 2002),
Native West Canadian (Salesman and Saskatchewan; Budowle et al., 2001), and East Asian (Vietnamese, Korean, Japanese, and Chinese; Budowle
et al., 2001) populations than in other Native American populations located to the south (Apache, Navajo, Northern Ontario, and Puna of Argentina; Budowle et al., 2001; Albeza et al., 2002) and European
populations (Russian, Polish, Austrian, Spanish,
and Greek; Kornienko et al., 2002; Pawlowski and
GENETIC ANALYSIS OF A NEOLITHIC SIBERIAN
Maciejewska, 2000; Neuhuber et al., 1999; Gusmao
et al., 2000; Sanchez-Diz et al., 2002). In spite of the
absence of publications presenting STR data from
Siberian populations usable in this study, the assignment-method results indicated that populations
living in regions neighboring northeastern Siberia,
i.e., extreme North America and east Asia, presented the highest affinity with the ancient skeleton
STR profile.
CONCLUSIONS
The ancient DNA analysis of a Neolithic Siberian
human bone sample allowed us to report on the most
ancient nuclear profile and mtDNA sequence of
north Siberia. Comparison of the ancient HV1 sequence and autosomal STR profile with modern Eurasian and Native American populations 1) supports
the hypothesis that the HV1 sequence harboring the
16223T, 16298C, and 16327T motifs could be the
founding lineage for haplogroup C in Asian populations, and 2) indicates that this ancient skeleton was
linked by its maternal lineage and nuclear DNA to
both east Siberian/Asian and Native American populations.
LITERATURE CITED
Albeza MV, Picornell A, Acreche N, Tomas C, Castro JA, Ramon
MM. 2002. Genetic variability at 14 STR loci in the Puna
population of northwestern Argentina. Int J Legal Med 116:
126 –132.
Anderson S, Bankier AT, Barrell BG, de Bruijn MH, Coulson AR,
Drouin J, Eperon IC, Nierlich DP, Roe BA, Sanger F, Schreier
PH, Smith AJ, Staden R, Young IG. 1981. Sequence and organization of the human mitochondrial genome. Nature 290:457–
465.
Budowle B, Shea B, Niezgoda S, Chakraborty R. 2001. CODIS
STR loci data from 41 sample populations. J Forensic Sci 46:
453– 489.
Budowle B, Chidambaram A, Strickland L, Beheim CW, Taft GM,
Chakraborty R. 2002. Population studies on three Native
Alaska population groups using STR loci. Forensic Sci Int 129:
51–57.
Cavalli-Sforza LL, Menozzi P, Piazza A. 1994. The history and
geography of human genes. Princeton: Princeton University
Press.
Clisson I, Keyser C, Francfort HP, Crubézy E, Samashev Z, Ludes
B. 2002. Genetic analysis of human remains from a double
inhumation in a frozen kurgan in Kazakhstan (Berel site, early
3rd century BC). Int J Legal Med 116:304 –308.
Cornuet JM, Piry S, Luikart G, Estoup A, Solignac M. 1999. New
methods employing multilocus genotypes to select or exclude
populations as origins of individuals. Genetics 153:1989 –2000.
Crawford MH. 1998. The origins of Native Americans: evidence
from anthropological genetics. Cambridge: Cambridge University Press.
Derenko MV, Shields GF. 1997. Mitochondrial DNA sequence
diversity in three north Asian aboriginal population groups.
Mol Biol (Mosk) 31:665– 669.
Derenko MV, Grzybowski T, Malyarchuk BA, Dambueva IK,
Denisova GA, Czarny J, Dorzhu CM, Kakpakov VT, MiscickaSliwka D, Wozniak M, Zakharov IA. 2003. Diversity of mitochondrial DNA lineages in south Siberia. Ann Hum Genet
67:391– 411.
Fily ML, Crubézy E, Courtaud P, Keyser C, Ebrard D, Ludes B.
1998. Paleogenetic analysis of the skeletons from the sepulchral cave of Elzarreko Karbia (Bronze Age, Basque Country).
C R Acad Sci [III] 321:79 – 85.
461
Forster P, Torroni A, Renfrew C, Röhl A. 2001. Phylogenetic star
contraction applied to Asian and Papuan mtDNA evolution.
Mol Biol Evol 18:1864 –1881.
Forsyth J. 1996. A history of the peoples of Siberia. Cambridge:
Cambridge University Press.
Gokhmana I, Tomtosovoy L. 1992. Anthropological search of the
Guiring and Roguinka Neolithic tombs [in Russian]. In:
Mokchavov U, editor. Archaeological search in Yakoutia. Novossibirsk: Novossibirsk Science Press. p 105–124.
Gusmao L, Sanchez-Diz P, Alves C, Lareu MV, Carracedo A,
Amorim A. 2000. Genetic diversity of nine STRs in two northwest Iberian populations: Galicia and northern Portugal. Int J
Legal Med 114:109 –113.
Horai S, Kondo R, Nakagawa-Hattori Y, Hayashi S, Sonoda S,
Tajima K. 1993. Peopling of the Americas founded by four
major lineages of mitochondrial DNA. Mol Biol Evol 10:23– 47.
Hummel S, Bramanti B, Schultes T, Kahle M, Haffner S, Herrmann B. 2000. Megaplex DNA typing can provide a strong
indication of the authenticity of ancient DNA amplifications by
clearly recognizing any possibility of modern contamination.
Anthropol Anz 58:15–21.
Keyser-Tracqui C, Crubézy E, Ludes B. 2003. Nuclear and mitochondrial analysis of a 2000-year-old necropolis in the Egyin
Gol Valley (Mongolia). Am J Hum Genet 73:247–260.
Kistenev S. 1992. Neolithic inhumation of Rodinkskoye [in
Russian]. In: Mokchavov U, editor. Archaeological search in
Yakoutia. Novossibirsk: Novossibirsk Science Press. p 68 –
83.
Kolman CJ, Sambuughin N, Bermingham E. 1996. Mitochondrial
DNA analysis of Mongolian populations and implications for
the origin of New World founders. Genetics 142:1321–1334.
Kornienko IV, Vodolazhsky DI, Ivanov PL. 2002. Genetic variation of the nine profiler plus loci in Russians. Int J Legal Med
116:309 –311.
Lee SD, Shin CH, Kim KB, Lee YS, Lee JB. 1997. Sequence
variation of mitochondrial region DNA control region in Koreans. Forensic Sci Int 87:99 –116.
Levin MG, Potatov LP. 1964. The peoples of Siberia. Chicago:
Chicago University Press.
Malhi RS, Eshleman JA, Greenberg JA, Weiss DA, Shook BAS,
Kaestle FA, Lorenz JG, Kemp BM, Johnson JR, Smith DG.
2002. The structure of diversity within New World mitochondrial DNA haplogroups: implications for the prehistory of
North America. Am J Hum Genet 70:905–919.
Merriwether DA, Rothhammer F, Ferrell RE. 1995. Distribution
of the four founding lineage haplotypes in Native Americans
suggests a single wave of migration for the New World. Am J
Phys Anthropol 98:411– 430.
Monsalve MV, Cardenas F, Guhl F, Delaney AD, Devine DV.
1996. Phylogenetic analysis of mtDNA lineages in South American mummies. Ann Hum Genet 60:293–303.
Neuhuber F, Radacher M, Meisner N, Tutsch-Bauer E. 1999.
Nine STR markers plus amelogenin (AmpFlSTR profiler plus):
a forensic study in an Austrian population. Int J Legal Med
113:60 – 62.
Pakendorf B, Wiebe V, Tarskaia LA, Spitsyn VA, Soodyall H,
Rodewald A, Stoneking M. 2003. Mitochondrial DNA evidence
for admixed origins of central Siberian populations. Am J Phys
Anthropol 120:211–224.
Pawlowski R, Maciejewska A. 2000. Forensic validation of a multiplex containing nine STRs-population genetics in northern
Poland. Int J Legal Med 114:45– 49.
Ricaut FX, Keyser-Tracqui C, Cammaert L, Crubézy E, Ludes B.
2003. Genetic analysis and ethnic affinities from two ScythoSiberian skeletons. Am J Phys Anthropol 123:351–360.
Sanchez-Diz P, Lareu MV, Brion M, Skitsa I, Carracedo A. 2002.
STR data for the AmpFlSTR Profiler Plus kit loci from Greece.
Forensic Sci Int 126:265–266.
Santos FR, Pandya A, Tyler-Smith C, Pena SDJ, Schanfield M,
Leonard WR, Osipova L, Crawford MH, Mitchell RJ. 1999. The
central Siberian origin for Native American Y chromosome.
Am J Hum Genet 64:619 – 628.
Schmerer WM, Hummel S, Herrmann B. 1999. Optimized DNA
extraction to improve reproducibility of short tandem repeat
462
F.-X. RICAUT ET AL.
genotyping with highly degraded DNA target. Electrophoresis
20:1712–1716.
Schurr TG, Subernik RI, Starikovskaya YB, Wallace DC. 1999.
Mitochondrial DNA variation in Koryaks and Itel’men: population
replacement in the Okhotsk Sea-Bering Sea region during the
Neolithic. Am J Phys Anthropol 108:1–39.
Starikovskaya YB, Subernik RI, Schurr TG, Kogelnik AM, Wallace
DC. 1998. mtDNA diversity in Chukchi and Siberian Eskimos:
implications for the genetic history of ancient Beringia and the
peopling of the New World. Am J Hum Genet 63:1473–1491.
Stone AC, Stoneking M. 1998. MtDNA analysis of a prehistoric
Oneota population: implications for the peopling of the New
World. Am J Hum Genet 62:1153–1170.
Torroni A, Schurr TG, Cabell MF, Brown MD, Neel JV, Larsen M,
Smith DG, Vullo CM, Wallace DC. 1993. Asian affinities and
continental radiation of the four founding Native American
mtDNAs. Am J Hum Genet 53:563–590.
Yao YG, Kong QP, Bandelt HJ, Kivisild T, Zhang YP. 2002.
Phylogeographic differenciation of mitochondrial DNA in Han
Chinese. Am J Hum Genet 70:635– 651.
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