close

Вход

Забыли?

вход по аккаунту

?

Craniometric variation of the Ainu An assessment of differential gene flow from Northeast Asia into Northern Japan Hokkaido.

код для вставкиСкачать
AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 137:283–293 (2008)
Craniometric Variation of the Ainu: An Assessment
of Differential Gene Flow From Northeast Asia Into
Northern Japan, Hokkaido
Tsunehiko Hanihara,1* Kohzo Yoshida,2 and Hajime Ishida3
1
Department of Anatomy and Biological Anthropology, Saga Medical School, Saga 849-8501, Japan
College of Humanities and Sciences, Nihon University, Tokyo 156-8550, Japan
3
Department of Anatomy, Faculty of Medicine, University of the Ryukyus, Okinawa 903-0215, Japan
2
KEY WORDS
Okhotsk culture people; admixture; simulative study; population structure; Jomon
ABSTRACT
In and after the latest Neolithic period
in Japan (B.P. 2,300 years), there were two distinct
waves of migration from eastern Asia. One is well known
as successive episodes in which indigenous inhabitants
of main-island Japan were intruded on by new arrivals
with advanced technology, and of a different genetic
stock. Another migration of people and culture, identified as the Okhotsk culture, reached the northeastern
part of Hokkaido. As opposed to main-island Japan, the
morphological continuity from the Neolithic to recent
inhabitants in Hokkaido (Ainu) is notable, so that the
evidence of admixture easily could have escaped notice.
In this study, the effects of gene flow from an outside
source on the pattern of among-group variation of Hokkaido Ainu are examined by means of two models. One
is the R-matrix model comparing observed and expected
craniometric variation for estimating differential external gene flow into a region. The other is a simple simulation model that estimates admixture in a population
with two parental populations. The two approaches give
similar results. The results suggest the possibility
of admixture between the migrants from Northeast
Asia, the Okhotsk culture people, and the indigenous
inhabitants in Hokkaido during the 5th to 12th centuries
A.D., at least in northeastern Hokkaido. Such gene flow
may have a certain degree of effect on the genetic
structure of recent Ainu. The findings further suggest
morphological heterogeneity in Northeast Asia during
the Holocene that has relevance for understanding
the morphological heterogeneity seen through time in
the New World. Am J Phys Anthropol 137:283–293,
2008. V 2008 Wiley-Liss, Inc.
The northernmost island of the Japanese Archipelago,
Hokkaido, is located in the subfrigid zone. The island
was not incorporated into Japan until the end of the
19th century. After the Palaeolithic age represented by
microblade industries and blade arrowhead cultures, the
prehistoric times of Hokkaido are divided into four periods based mainly on archaeological chronology; Jomon
(13,000–2,300 years B.P.), Epi-Jomon (2,300–1,300 years
B.P.), Satsumon (1,300–700 years B.P.), and Ainu culture
periods (700 years B.P. to recent). The Ainu, the indigenous inhabitants of Hokkaido, are an ethnic group with
a distinct Ainu culture (reviewed by Kodama, 1970;
Ishida, 1988, 1996; Hanihara et al., 1998; Fitzhugh and
Dubreuil, 1999). According to Kodama (1970), some
Europeans began to pay attention to the Ainu in Hokkaido since the middle of the 17th century because of
their unique, European-like features. The first scientific
report of the cranial morphology of the Ainu was performed by Busk (1867), and followed by Davis (1870).
Concerning the population history of the Japanese, the
‘‘dual structure model’’ (Hanihara, 1991) is generally
accepted. Modern Japanese have two primary origins:
the aboriginal Jomon people, Neolithic hunter-gatherers,
and migrants from the eastern Asian continent via the
Korean Peninsula to the southwestern part of Japan,
who brought rice agriculture and metalworking technologies. During the period from the end of the Jomon age to
the early historic age, or from 2,300 years B.P. to 1,300
years B.P., admixture between the indigenous Jomon
people and the incoming populations blurred the distinction between the two populations except for geographi-
cally isolated Hokkaido. The impact of the post-Jomon
migrants was unexpectedly large, so that a majority of
modern Japanese carry a large amount of eastern Asian
characteristics (Turner, 1987, 1990; Brace and Hunt,
1990; Lahr, 1995, 1996; Ossenberg et al., 2006).
In the northern part of the Japanese Archipelago,
another migration of people and culture, known as the
Okhotsk culture, from northeastern Asian continent,
during the 5th to 12th centuries A.D., is identified. The
Okhotsk culture spread from southern Sakhalin Island
and the northernmost part of Hokkaido through the
coastal region of the Sea of Okhotsk to eastern Hokkaido
(Ishida, 1988; Amano, 2003; Sato et al., 2007; Komesu
et al., 2008). The people of the Okhotsk culture have
physical and cultural affinities to not the contemporary
C 2008
V
WILEY-LISS, INC.
C
Grant sponsor: Japan Society for the Promotion of Science; Grant
numbers: 1850220, 18370099; Grant sponsors: Japan Fellowship for
Research in United Kingdom, Japan Society for the Promotion of
Science; Smithsonian Opportunities for Research and Study, Smithsonian Institution Fellowship Program.
*Correspondence to: Tsunehiko Hanihara, Department of Anatomy
and Biological Anthropology, Saga Medical School, Nabeshima, Saga
849-8501, Japan. E-mail: hanihara@cc.saga-u.ac.jp
Received 13 February 2008; accepted 22 April 2008
DOI 10.1002/ajpa.20869
Published online 9 July 2008 in Wiley InterScience
(www.interscience.wiley.com).
284
T. HANIHARA ET AL.
TABLE 1. Materials used and the brief information
Sample name
Recent Ainu
Sakhalin island
Kurile islands
Hokkaido
Sea of Okhotsk/Soya
Sea of Okhotsk/Abashiri
Southeast/KushiroNemuro
Southeast/Tokachi
Central/Hidaka
Southwest/Ishikari-Iburi
Southwest/Shiribeshi
Comparative samples
Jomon/Hokkaido
Jomon/East Japan
Epi-Jomon
Okhotsk culture people
Modern Japanese
Brief information
Sakaehama, Mohka, Honto villages, Sakhalin Island (Kyoto Univ., Tohoku Univ.)
Mainly from Kunashiri Island (Univ. of Tokyo, Tohoku Univ.)
Esashi and Wakkanai (Univ. of Tokyo)
Abashiri, Monbetsu, Shari, and Toro (Univ. of Tokyo, Sapporo Medical Univ., Hokkaido Univ.)
Akkeshi and Shibetsu, Kushiro and Nemuro Subprefectures (Univ. of Tokyo, Sapporo Medical Univ.)
Hiroo and Ohtsu (Utsunai), Tokachi Subprefectures (Univ. of Tokyo)
Urakawa, Atsubetsu, Horoizumi, Mitsuishi, Saru, Shizunai, and Urakawa regions, Hidaka
Subprefecture (Univ. of Tokyo, Sapporo Medical Univ.)
Chitose and Usu, Iburi Subprefecture, and Sapporo, Ishikari Subprefecture (Univ. of Tokyo,
Sapporo Medical Univ.)
Otaru, Yoichi, and Iwanai, Shiribeshi Subprefecture (Univ. of Tokyo)
Usu-Moshiri, Kitakogane, Takasago, Irie (Abuta) Shell mounds; Motowa-Nish (Muroran),
Yakumo-Kotan (Oshima), Tenneru (Kushiro) Shimamaki (Shiribeshi) and other sites
(Tohoku Univ. Sapporo Medical Univ. Univ. of Tokyo)
Ebishima, Nakazawahama, Ohora, Hosoura (Iwate Prefecture); Sanganji (Fukushima Prefecture);
Sakai-Numazu, Hashimotogakoi, Satohama (Miyagi Prefecture); Kashidokoro (Akita Prefecture)
and Other sites: From Tohoku region of Eastern Japan (Univ. of Tokyo, National Museum of
Nature and Science, Tohoku Univ.)
Usu-Moshiri site (Tohoku Univ.)
5-12 century A.D. Hamanaka, Rebun Island; Ohmisaki, Soya; Moyoro, Abashiri; Susuya, Sakhalin
(Hokkaido Univ, Sapporo Medical Univ., Kyoto Univ.)
Mainly from Tohoku district, Northern part of Honshu, Japan (Tohoku Univ., The Univ. of Tokyo)
Epi-Jomon and Satsumon people, but to the populations
in the lower basin of the Amur River such as Nivkhi,
Orochs, Nanaians, Ulchs, Negidal, etc. (Yamaguchi,
1974; Ishida, 1994, 1995, 1996; Hudson, 2004; Shigematsu et al., 2004; Tajima et al., 2004; Sato et al., 2007;
Komesu et al., 2008).
Contrary to the large-scale migration and the subsequent rapid changes in the physical characteristics of
main-island Japanese, little genetic contribution of the
Okhotsk culture people to recent Ainu has been suggested (reviewed by Ishida, 1996; Hanihara et al., 1998).
It is often emphasized that the craniofacial and dental
features of the Native Hokkaido populations, the recent
Ainu, show remarkable similarities to those of skeletal
series of the Jomon people, suggesting overall post-Neolithic temporal and spatial population continuity in Hokkaido (Yamaguchi, 1974; Turner, 1987, 1990; Brace et
al., 1989; Brace and Hunt, 1990; Dodo and Ishida, 1990;
Kozintsev, 1990, 1992; Hanihara, 1991, 1998; Lahr, 1995,
1996; Dodo and Kawakubo, 2002; Ossenberg et al.,
2006). Recent morphological, genetic, and archaeological
studies suggest, however, the possible effect of Northeast
Asian admixture on the formation of physical and cultural characteristics of the Ainu (Horai et al., 1996;
Schurr et al., 1999; Amano, 2003; Shigematsu et al.,
2004; Tajima et al., 2004; Kondo, 2005; Sato et al., 2007;
Komesu et al., 2008).
Regarding the origins and affinities of the Ainu/Jomon
group, several competing hypotheses have been proposed
for nearly a half century: European origin (Howells,
1959; Kodama, 1970), Australian (Oceanian) origin
(Aroutiounov, 1968), Southeast Asian origin (Turner,
1987, 1990), or Northeast Asian origin (Nei, 1995;
Omoto, 1995; Omoto and Saitou, 1997). Today, it is often
suggested that the Ainu (and Jomon) are one of the most
obvious outliers in an eastern Asian context (Howells,
1986; Brace et al., 2001, 2006; Lahr, 1995, 1996; Hanihara et al., 1998; Hanihara, 2006). For example, Tajima
American Journal of Physical Anthropology
et al. (2004) find predominant distribution of the YAP1
lineages (Y-haplogroup D) among the Ainu, which are
sporadically distributed in Asia such as in Tibet, Andaman Islands, and the Japanese Archipelago. Using craniometric data, Brace et al. (1989) and Brace and Hunt
(1990) show that the Jomon/Ainu group falls into a cluster with Pacific populations. Jantz and Owsley (2001,
2003) indicate, on the other hand, a possible association
between the Ainu and early North Americans. These unusual population affinities of the Ainu may suggest their
complex population structure or evolutionary forces
(such as long-term external gene flow, genetic drift by
isolation, or natural selection). The seminal morphological studies carried out by Yamaguchi (1974, 1981) demonstrate, moreover, that the Ainu in Hokkaido as a
whole are not necessarily uniform.
Such knowledge raises the fundamental question of
whether the Ainu are the direct descendants of the
Jomon, and how the observed variation between the
Jomon and Ainu groups as well as within Ainu themselves should be interpreted in an evolutionary context.
Few studies have, however, specifically addressed these
issues.
Many morphological studies based mainly on classic
multivariate analyses have focused on the degree of similarity between populations. In these analyses, the
degree of similarity is assumed to be a reflection of population history rather than population structure (Howells,
1973, 1989, 1995; Lahr 1995, 1996; Brace and Hunt,
1990; Brace et al., 2001, 2006; Pietrusewsky, 2000; and
many others). However, biological distance between populations is seen as reflecting not only common ancestry,
but also genetic drift and/or gene flow. As pointed out by
Relethford (1991, 1994, 1996), it is difficult to determine
which factor or combination of factors is responsible for
a given set of biological distances.
The model developed by Relethford and Blangero
(1990) and applied in this study tests the deviations of
CRANIOMETRIC VARIATION OF THE AINU
285
Fig. 1. Geographical location of the cranial samples used in this study.
observed variation from an expected pattern of genetic
variation, which provide information about differential
gene flow from outside sources, genetic drift, population
size, and so forth (Relethford, 1991, 1994; Relethford and
Blangero, 1990; Relethford and Harpending, 1994;
Powell and Neves, 1999; Stojanowski, 2004; Hanihara,
2008). In this model, when considering long-term evolution on a local scale, a population having received
greater long-range gene flow within groups will show
greater variation than expected. On the other hand, a
population with less long-range gene flow will show less
variation than expected. This method provides an objecAmerican Journal of Physical Anthropology
286
T. HANIHARA ET AL.
TABLE 2. List of 34 craniofacial measurements
1. Maximum cranial length (GOL)
2. Nasion-opisthocranion (NOL)
3. Cranial base length (BNL)
4. Maximum cranial breadth (XCB)
5. Minimum frontal breadth (M9)
6. Maximum frontal breadth (XFB)
7. Biauricular breadth (AUB)
8. Biasterionic breadth (ASB)
9. Basion–bregma height (BBH)
10. Sagittal frontal arc (M26)
11. Saggital parietal arc (M27)
12. Saggital occipital arc (M28)
13. Nasion-bregma chord (FRC)
14. Bregma-lambda chord (PAC)
15. Lambda-opisthion chord (OCC)
16. Basion prosthion length (BPL)
17. Breadth between frontomalare temporale (M43)
18. Bizygomatic breadth (ZYB)
19. Middle facial breadth (M46)
20. Nasion prosthion height (NPH)
21. Interorbital breadth (DKB)
22. Orbital breadth (M51)
23. Orbital height (OBH)
24. Nasal breadth (NLB)
25. Nasal height (NLH)
26. Palate breadth (MAB)
27. Mastoid height (MDH)
28. Mastoid width (MDB)
29. Breadth between frontomalare orbitale—Frontal chord
(M43(1))
30. Frontal subtense (No 43c)
31. Minimum horizontal breadth of the nasalia (sc)—Simotic
chord (M57, WNB)
32. Simotic subtense (No 57a, SIS)
33. Breadth between zygomaxillare anterius—Zygomaxillary
chord (M46b, ZMB)
34. Zygomaxillary subtense (No 46c, SSS)
For additional description, see Howells (1973, 1989). M; Martin
number, and No; Bräuer (1988).
tive means for investigating differences in long-range
gene flow (Crawford and Devor, 1980; Relethford and
Blangero, 1990; Kondo, 2005).
The purpose of the present study is to examine the
effects of admixture on the patterns of among-group variation of Ainu by applying Relethford and Blangero’s
(1990) R-matrix approach and simple simulative study to
craniometric data. The findings provide, moreover, an
additional insight into the relationships between Ainu,
Northeast Asians, and Asian originated populations as
represented by New World populations, because the
Northeast Asian region and the time period (in and after
the late Pleistocene) of the people considered as the parental population of the Jomon/Ainu, are both relevant
to the peopling of the Americas.
MATERIALS AND METHODS
The cranial materials representing the Ainu are
derived from three major geographic regions: Hokkaido,
Sakhalin, and Kurile Islands. In this study, the cranial
series of the Hokkaido Ainu are divided into seven local
groups based mainly on the administrative divisions of
the subprefecture, which correspond almost exactly to
the watersheds (Kondo, 1995). Brief information on the
Ainu groups and the five comparative samples is given
in Table 1. The location of the samples used is shown in
American Journal of Physical Anthropology
TABLE 3. Number of samples for each sex and census sizes of
the local Ainu populations
Male
Female
Recent Ainu
Sakhalin island
37
Kurile islands
Kunashiri Island
7
Hokkaido
Coastal region along the Sea of Okhotsk
Soya
7
Abashiri
14
Southeast
Kushiro and Nemuro
9
Tokachi
15
Central
Hidaka
38
Southwest
Ishikari and Iburi
13
Shiribeshi
11
Camparative samples
Jomon/East Japan
50
Jomon/Hokkaido
32
Epi-Jomon
20
Okhotsk people
48
Modern Japanese
150
Population
size at
1875
19
2,372
9
64
8
8
358
951
5
6
2,040
1,475
33
5,236
6
12
3,868
799
43
13
8
35
68
–
–
–
–
–
Figure 1. Phenotypic variation and biological distance
were assessed using 34 craniofacial measurements
(Table 2). The sample sizes for both sexes are listed in
Table 3.
First, to examine patterns of phenotypic variation
with reference to population structure of the Ainu, we
performed Relethford and Blangero’s (1990) R-matrix
analysis, restricting our focus to nine local Ainu groups
(Table 3). To conduct this analysis, an estimate of average heritability for craniofacial traits of h2 5 0.55
obtained by Devor (1987) and followed by several other
studies (Relethford, 1994, 2002, 2004; Relethford and
Harpending 1994; Donnelly and Konigsberg, 1998;
Powell and Neves, 1999; Kondo, 2005), as well as census
population size for the Ainu given in Table 3, were used.
Recently, Carson (2006) published new estimates for heritabilities of 33 craniometric dimensions, showing that
heritability rates of metric cranial features vary a great
deal depending of the measurement being used. Twentyfour out of the 33 variables are the same as those used
here, and the mean heritability rate estimated for the 24
measurements is calculated to be 0.39. Based on such
estimation, we used additionally the distinct value of
heritability of h2 5 0.40. The census data for the Ainu
were surveyed by one of the present authors (K.Y.)
(Yoshida, 2006, 2007). In the present study, census population sizes of the Ainu in 1875 are adopted because information on the Ainu used in this study consists largely
of that collected by Y. Koganei in the late 19th century
(Koganei, 1893). The R-matrix analysis was performed
separately by sex and pooling both sexes. Before the
male and female samples were pooled, the Z-score standardization was performed within each sex separately.
This is a common method for removing sex-related size
variation (Relethford, 1994; Kondo, 2005).
To analyze and visualize the pattern of similarity
between recent Ainu groups and the neighboring populations, Torgerson’s (1952) metric multidimensional scaling
method was applied to the distance matrix transformed
from R-matrix (Relethford and Harpending, 1994).
287
CRANIOMETRIC VARIATION OF THE AINU
TABLE 4. Nine local Ainu samples: genetic distances to the centroid (rii ), as well as observed, expected, and residual variances computed using an average heritability of h2 5 0.40 and 0.55
rii
Sample name
Sakhalin
Kurile Islands
Soya
Abashiri
Kushiro-Nemuro
Tokachi
Hidaka
Ishikari-Iburi
Shiribeshi
Sex
0.40
0.55
Observed
variance
M
F
M1F
M
F
M1F
M
F
M1F
M
F
M1F
M
F
M1F
M
F
M1F
M
F
M1F
M
F
M1F
M
F
M1F
0.1140
0.2087
0.1326
0.2333
0.3651
0.1574
0.2082
0.1762
0.0940
0.1564
0.2121
0.1238
0.1344
0.2808
0.1305
0.1166
0.1924
0.0741
0.0516
0.0863
0.0553
0.0365
0.0866
0.0512
0.0795
0.2486
0.0790
0.0858
0.1609
0.0907
0.1757
0.2813
0.1336
0.1568
0.1358
0.0788
0.1178
0.1634
0.0950
0.1012
0.2164
0.1018
0.0878
0.1483
0.0588
0.0388
0.0665
0.0344
0.0275
0.0668
0.0307
0.0599
0.1916
0.0718
1.1286
1.0636
1.1193
0.8451
1.0258
0.9492
1.3236
1.3962
1.2909
1.0664
1.2891
1.1192
0.9845
0.9232
0.9329
0.9994
0.8662
0.9490
0.9712
1.2097
1.1017
1.0447
0.9155
0.9832
1.2625
0.9977
1.1384
The next part of the analysis deals with the estimation
of admixture. The analysis is based on two assumptions.
The first is that the hybrids tend to show morphologically median characteristics between the parental populations. The relationship between cranial morphology
and genetic relationships is complex, and we are not convinced that we know enough about the genetic and epigenetic mechanisms underlying it to assume a median
phenotype for hybrids. However, several comparative
studies support, or at least do not deny, the argument
that hybrids tend to be morphologically median (Hanihara, 1987; Hanihara and Hanihara, 1989; Howells,
1989; Brace et al., 1989; Pietrusewsky, 2000; Hallgrı́msson et al., 2004). The second is that the Epi-Jomon and
Okhotsk culture people represent the two possible parental populations from which the recent Ainu groups with
greater observed variation than expected were drawn.
The admixture analysis was performed using the following equations (Hanihara, 1987):
x1 ¼ mðp1 a þ p2 bÞ;
ð1Þ
where x1 is an estimated measurement for the hybrids, a
is an actual measurement of the Epi-Jomon population,
b is that of the Okhotsk culture people, p1 is a rate of
admixture for the former population, and p2 is that for
the latter population (p1 1 p2 5 1). This model includes
the rate of microevolution or temporal morphological
change (m) that took place without admixture. A microevolution rate during the time period under consideration
is computed by
m ¼ c=a;
ð2Þ
where c is an actual measurement for the recent Ainu
without admixture.
Expected variance
Residual variance
S.E.
0.40
0.55
0.40
0.55
0.40
0.55
1.0531
1.0541
1.0260
0.9113
0.8458
0.9967
0.9410
1.0975
1.0717
1.0026
1.0497
1.0365
1.0288
0.9582
1.0285
1.0499
1.0758
1.0952
1.1272
1.2172
1.1175
1.1451
1.2168
1.1223
1.0941
1.0010
1.0894
1.0493
1.0550
1.0351
0.9462
0.9035
0.9862
0.9678
1.0865
1.0485
1.0126
1.0518
1.0302
1.0316
0.9852
1.0224
1.0470
1.0708
1.0714
1.1032
1.1736
1.0991
1.1162
1.1733
1.1033
1.0791
1.0164
1.0565
0.0756
0.0094
0.0933
20.0662
0.1799
20.0475
0.3826
0.2987
0.2192
0.0638
0.2394
0.0827
20.0443
20.0350
20.0957
20.0506
20.2096
20.1462
20.1561
20.0076
20.0158
20.1004
20.3013
20.1391
0.1685
20.0033
0.0490
0.0793
0.0086
0.0842
20.1011
0.1222
20.0370
0.3558
0.3097
0.2424
0.0538
0.2373
0.0890
20.0471
20.0620
20.0895
20.0476
20.2046
20.1223
20.1321
0.0360
0.0026
20.0715
20.2578
20.1201
0.1834
20.0187
0.0819
0.0479
0.0973
0.0471
0.5373
0.3450
0.2214
0.7494
0.7274
0.2760
0.0962
0.7117
0.0933
0.2338
0.3740
0.1533
0.1712
0.6128
0.1207
0.0660
0.1429
0.0456
0.1621
0.5299
0.1648
0.4751
0.2008
0.1643
0.0479
0.0973
0.0481
0.5378
0.3452
0.2172
0.7489
0.7275
0.2823
0.0962
0.7116
0.0942
0.2338
0.3743
0.1486
0.1711
0.6127
0.1163
0.0659
0.1432
0.0460
0.1620
0.5294
0.1615
0.4753
0.2009
0.1658
To simulate the rate of admixture, measurements for
the hypothetical populations are estimated using Eq. (1)
by changing p1 and p2 (admixture proportions of the two
parental populations), then the distance coefficients
measured by Mahalanobis D2 between the Ainu samples
with possible long-range external gene flow (possible
hybrid populations) and the hypothetical populations are
computed. This procedure may suggest the rate of
admixture in the Ainu populations.
RESULTS
Table 4 gives the rii, the average dispersion of the
samples around the genetic centroid, as well as observed,
expected, and residual variance calculated using average
heritabilities of h2 5 0.55 and 0.40, separately by sex
and pooled, for the nine local samples of the Ainu from
three main geographic regions (Hokkaido, Sakhalin, and
Kurile Islands). The plot of observed variance versus
genetic distance from the centroid (rii) for the male,
female, and sex-combined samples with the expected
regression line based on h2 5 0.55 is shown in Figure 2.
Of course, the rii values and the related parameters
depend on the specific choice of average heritability
used. However, the differences presented here are fairly
small and do not dramatically affect the results (as
pointed out by Roseman and Weaver, 2004). The most
noticeable outlier to the expected regression line is the
Soya sample in three datasets, followed by the male
Shiribeshi and the female Abashiri samples. These samples exhibit greater phenotypic variation than expected,
indicating greater than average gene flow from outside
the region.
To avoid a potential influence of nonrandom sampling
effects caused by small sample sizes that may artificially
American Journal of Physical Anthropology
288
T. HANIHARA ET AL.
Fig. 2. Plot of observed phenotypic variance versus genetic
distance from the centroid (rii) for the nine local Ainu samples.
The line indicates the expected regression line derived from
Relethford and Blangero’s (1990) model.
bias between-group variation, the seven local Hokkaido
Ainu series were grouped into four regional clusters (Table 3). The results of the analysis using combined datasets are reported in Table 5 and Figure 3. The sample
from the coastal region along the Sea of Okhotsk and
the male Sakhalin Ainu sample are plotted above the
expected regression line, suggesting greater than average admixture with outside populations.
Given estimates of the differential gene flow from outside sources, possible patterns of external migration and
admixture are further examined using the six Ainu
samples and the five comparative series (Table 1). The
two-dimensional expression of multidimensional scaling
applied to the distance matrix converted from R-matrix
(Relethford and Harpending, 1994) is shown in Figure 4.
All figures express about 90% of the total variance.
American Journal of Physical Anthropology
Three major constellations are evident in the three diagrams. The Jomon and Epi-Jomon samples form a relatively tight cluster in one corner of the diagram. This
cluster borders a more diffused recent Ainu grouping.
The sample of modern Japanese forms the third constellation. The Okhotsk sample forms a separate cluster in
the three scattergrams. As a whole, the six Ainu groups
appear as a mid-morphology between Jomon/Epi-Jomon
and modern Japanese. However, the Ainu samples from
Sakhalin Island and the coastal region of the Sea of
Okhotsk are more northern-like than are the other Ainu
series, suggesting admixture with Okhotsk culture
people.
On the basis of these results shown in Figure 4, we
estimated admixture for the recent Ainu from the coastal
region along the Sea of Okhotsk under the assumption
that the Epi-Jomon and the Okhotsk culture people were
the two parental populations. The microevolution rate
during the period from Epi-Jomon to recent Ainu was
computed by Eq. (2) using recent Ainu samples from
Southwest Hokkaido for males and from central Hokkaido for females. As shown in Figures 2 and 3, these
two samples show the smallest rii and the least residual
variance, indicating the least deviation from Relethford
and Blangero’s (1990) model, which assumes constant
effective population sizes and symmetric numbers of
migrants between pairs of populations. This may allow
us to regard the two Ainu samples as indigenous lineages without a large impact from gene flow from outside
sources or genetic drift.
Table 6 shows the results of the simulation. The male
Soya sample shows the greatest similarity to the hypothetical population using measurements computed with
the assumption that the rates of admixture were 0.87 for
the indigenous lineage (Epi-Jomon population) and 0.13
for the migrant lineage (Okhotsk culture people). On the
other hand, the male Abashiri sample shows the greatest
similarity to the hypothetical population with the admixture rate of 0.98 for the indigenous lineage and 0.02 for
the migrant lineage. The female samples show somewhat
higher admixture rates than the male samples (Table 6).
The same analysis was performed pooling the two regional samples. The combined sample exhibits the greatest similarity to the hypothetical population with the
admixture rate of indigenous lineage : migrant lineage
5 0.97 : 0.03 and 0.76 : 0.24 in males and females,
respectively.
In Table 6, moreover, Mahalanobis’ D2 distances
between the best fit admixtures estimations and Ainu
samples as well as the mean distance among Ainu
groups are presented to see how close the admixture
estimations are to the real data of recent Ainu. In the
female samples, the distances between hypothetical populations with best fit admixture estimations and the real
Ainu samples are smaller than those among the mean
Ainu groups compared. While the patterning of the two
distances in the male samples differs from that found in
the female results, this presentation suggests the fit of
the simulation model to admixture analysis in morphological variation.
DISCUSSION
It is often emphasized that all cranial diversity of the
recent Ainu in Hokkaido could be accommodated in one
single human biological stock, the Neolithic Jomon lineage (Dodo and Kawakubo, 2002). Much of this debate
TABLE 5. Six regional Ainu samples: genetic distances to the centroid (rii ), as well as observed, expected, and residual variances
computed using an average heritability of h2 5 0.40 and 0.55
rii
Sample name
Sakhalin
Kurile Islands
Sea of Okhotsk
Southeast
Central
Southwest
Sex
0.40
0.55
Observed
variance
M
F
M1F
M
F
M1F
M
F
M1F
M
F
M1F
M
F
M1F
M
F
M1F
0.1167
0.2317
0.1285
0.2441
0.3060
0.1715
0.1060
0.1085
0.0740
0.0486
0.1071
0.0368
0.0548
0.0834
0.0398
0.0152
0.1083
0.0316
0.0872
0.1760
0.0956
0.1824
0.2326
0.1275
0.0792
0.0825
0.0550
0.0363
0.0814
0.0274
0.0410
0.0634
0.0296
0.0114
0.0823
0.0235
1.0991
1.0162
1.0777
0.8256
0.9705
0.8994
1.0949
1.2815
1.1511
1.0056
0.8576
0.9615
0.9449
1.1592
1.0584
1.0938
0.9621
1.0428
Fig. 3. Plot of observed phenotypic variance versus genetic
distance from the centroid (rii) for the six regional Ainu samples. The line indicates the expected regression line.
Expected variance
0.40
0.55
1.0083
0.9418
0.9888
0.8628
0.8506
0.9401
1.0205
1.0928
1.0507
1.0861
1.0945
1.0929
1.0790
1.1235
1.0895
1.1241
1.0930
1.0988
1.0143
0.9666
1.0037
0.9085
0.9003
0.9683
1.0231
1.0764
1.0487
1.0708
1.0776
1.0794
1.0656
1.0987
1.0769
1.0985
1.0765
1.0837
Residual variance
0.40
0.55
0.0908
0.0744
0.0888
20.0372
0.1198
20.0407
0.0744
0.1888
0.1004
20.0805
20.2369
20.1314
20.1340
0.0357
20.0311
20.0304
20.1309
20.0559
0.0848
0.0496
0.0739
20.0828
0.0702
20.0689
0.0718
0.2051
0.1023
20.0652
20.2200
20.1179
20.1207
0.0604
20.0185
20.0047
20.1144
20.0408
S.E.
0.40
0.55
0.0451
0.0951
0.0449
0.5306
0.2848
0.2118
0.0721
0.2758
0.0590
0.0669
0.1960
0.0571
0.0664
0.1387
0.0444
0.0996
0.1112
0.0558
0.0451
0.0951
0.0449
0.5313
0.2850
0.2121
0.0721
0.2760
0.0590
0.0669
0.1958
0.0570
0.0663
0.1389
0.0443
0.0996
0.1111
0.0557
Fig. 4. Two dimensional scattergrams drawn by multidimensional scaling.
290
T. HANIHARA ET AL.
TABLE 6. The rate of admixture between the two parental populations (Epi-Jomon and Okhotsk people) for the hypothetical
population who show the largest similarity to the Ainu samples from the Okhotsk Sea coast, and Mahalanobis’ D2 between the Ainu
samples from the Okhotsk Sea coast and the hypothetical populations as well as mean Mahalanobis’ D2 among Ainu groups
Epi-Jomon
Okhotsk people
Mahalanobis’ D2
from hypothetical
population
Soya
M
F
0.87
0.66
0.13
0.34
7.73
4.99
6.88
7.00
Abashiri
M
F
0.98
0.93
0.02
0.07
7.96
3.67
7.03
7.06
Coastal region of the Sea
of Okhotsk (Soya 1 Abashiri)
M
F
0.97
0.76
0.03
0.24
7.60
5.72
7.65
7.01
relies on a putative singularity of the dental and cranial
morphology of the Jomon and Ainu (Howells, 1986;
Turner, 1987, 1990; Brace et al., 1989; Brace and Hunt,
1990; Dodo and Ishida, 1990; Kozintsev, 1990, 1992;
Ishida, 1995; Lahr, 1995, 1996), and the fact that the
craniometric features of Okhotsk culture people are very
different from those prevalent today among the Ainu
(Yamaguchi, 1981; Ishida, 1988, 1994). On the other
hand, several comparative studies based on nonmetric
cranial traits and ancient mitochondrial DNA analysis
suggest that the Okhotsk culture people introduced
northern genetic elements to the Ainu (Ishida, 1988,
1996; Hanihara et al., 1998; Ishida and Kondo, 1999;
Shigematsu et al., 2004; Sato et al., 2007; Komesu et al.,
2008).
It is generally accepted that small effective population
size would decrease genetic and phenotypic (human
craniometric) variation of a population resulting from
genetic drift, founder’s effects, bottlenecks, etc. (Relethford and Harpending, 1994; Relethford, 2001). Despite the small population sizes of the recent Ainu
groups from the coastal region of the Sea of Okhotsk
(Table 3), the present examination shows a systematic
pattern of positive deviation from an expected pattern of
genetic variation in these samples. Such results suggest
an indispensable external gene flow into northeastern
Hokkaido. The simple simulative study for migration
applied to craniometric data further suggests that the
migrants during Epi-Jomon and early Satsumon periods
from Northeast Asia into Hokkaido, the Okhotsk culture
people, had an impact on the formation of craniometric
characteristics of recent Ainu at least in northeastern
Hokkaido, where the Okhotsk culture once thrived. It is
a matter of additional interest that the estimated admixture rates are somewhat higher for females than for
males (Table 6). At the simplest level, we do not necessarily rule out the possibility of sampling error based on
the relatively small sample sizes or inappropriate parameter estimation in the simulation model. However,
the Ainu showed closer affinities with Northeast Asians
in mtDNA sequence types (maternal lineage) than in
Y-haplogroups (paternal lineage) (Tajima et al., 2004).
Tajima et al. (2004) found the shared mtDNA types
between the Ainu and Northeast Asians, but no Ainuspecific Y-haplogroups in Northeast Asians, suggesting
maternal contacts between them, or at least unidirectional migration over time, from Northeast Asia to
Hokkaido.
Several comparative studies of mtDNA and Y-chromosome variations have suggested a higher female than
male migration rate due to the widespread practice of
patrilocality (Seielstad et al., 1998; Stoneking, 1998;
American Journal of Physical Anthropology
Mean D2 distance
among Ainu
groups
Oota et al., 2001). Recent investigations indicate, on the
other hand, that different patterns of genetic structure
between mtDNA and Y-chromosome are not shaped by
the higher migration rate among females than among
males but by a shorter matrilineal generation interval,
at least on a global scale (Helgason et al., 2003; Wilder
et al., 2004; Shriver, 2005).
The different pattern of variation depending on sex
presented here may be an additional morphological support for a genetically suggested sex-mediated migration
model such as patrilocality and/or shorter average generation times for females than males.
These findings suggest that both the quantitative and
qualitative morphological analyses, as well as genetic
studies, generate essentially convergent results regarding possible admixture between the ancestors of recent
Ainu and the Okhotsk culture people in the post-Jomon
periods. The different rates of genetic drift and external
gene flow may be possible factors responsible for the
morphological differentiation and diversification of
Jomon and Ainu as well as among Ainu themselves.
The results obtained show that the Sakhalin Ainu
sample shows larger craniometric variation than
expected and is much like the Okhotsk culture people.
However, the Sakhalin Ainu migrated from Hokkaido to
the southern part of Sakhalin Island during the 14th to
15th century A.D., long after the Okhotsk culture had
disappeared (Ohyi, 1985; Ishida and Kida, 1991; Fitzhugh and Dubreuil, 1999). This indicates that the northern-like features of the Sakhalin Ainu may be attributed
to more recent admixture with indigenous groups of
Sakhalin Island such as Oroch, Gilyak, Nivkh, etc.,
rather than the direct influence of the Okhotsk culture
people (Kozintsev, 1990; Turner, 1990; Ishida and Kida,
1991; Fitzhugh and Dubreuil, 1999).
Taking these into account, the results of the present
study provide an additional support for the previous suggestion (e.g. Shigematsu et al., 2004; Komesu et al.,
2008) that the genetic characteristics of the recent Ainu
do not necessarily reflect those of the Jomon population,
as often hypothesized in genetic studies (Hammer and
Horai, 1995; Nei, 1995; Omoto, 1995; Omoto and Saitou,
1997).
Finally, the craniofacial morphological shift of the
Ainu to Northeast Asians illuminates the relationships
between Paleo-Indians and Northeast Asians again.
Many studies of ancient population groups arriving in
the New World indicate that they had a distinct morphological pattern from those characterizing Native Americans and Northeast Asians today, their supposed descendants and sister groups, respectively (Neves and
Pucciarelli, 1991; Gonzáletz-José et al., 2001, 2005). The
CRANIOMETRIC VARIATION OF THE AINU
Paleo-Indians from Central and South America present
craniofacial features similar to recent Australians and
Melanesians (Gonzáletz-José et al., 2003, 2005; Neves
and Hubbe 2005; Sardi et al., 2005; Hubbe et al., 2007;
Neves et al., 2007a, b). On the other hand, morphological
affinities between Ainu and Paleo-Indians from North
America have been suggested (Lahr, 1995; Brace et al.,
2001; Jantz and Owsley, 2001, 2003).
The present results indicate some morphological association of the Ainu with northeastern Asian continental
populations as suggested from genetic studies (Horai
et al., 1996; Omoto and Saitou, 1997; Schurr et al., 1999;
Underhill et al., 2001; Tajima et al., 2004; Sato et al.,
2007). It has been often suggested, moreover, that the
Jomon/Ainu group retains some morphological characteristics attributed to those of the upper Palaeolithic inhabitants of the eastern Asian region (Yamaguchi, 1982;
Lahr, 1995, 1996; Hanihara et al., 1998; Brace et al.,
2001). This possible interpopulation relationship between
Jomon/Ainu and early Northeast Asians sheds light on a
model in which Northeast Asian descendants with more
generalized craniofacial features entered the New World
continent in the late Pleistocene and early Holocene
(Neves and Pucciarelli, 1991; Merriwether et al., 1995;
Bonatto and Salzano, 1997; Kozintsev, 1999; Brace et al.,
2001; Schurr, 2004; Hanihara, 2006; Fagundes et al.,
2008).
CONCLUSIONS
Gene flow is one of many potential historical factors
affecting human variation. In the present study, we
examined the potential effects of gene flow (admixture)
on the pattern of among-group variation of Hokkaido
Ainu populations using the R-matrix method. The findings suggest that the Ainu from the coastal region along
the Sea of Okhotsk cannot be regarded as a homogenous
population descended from Jomon and Epi-Jomon lineage. The Okhotsk culture people who have physical and
cultural affinities with Northeast Asians have contributed indelibly to the unique cultural and genetic characteristics of the recent Ainu. The possibility of sex-mediated migration, higher female gene flow, was suggested
by a simple simulation study for admixture. This may be
attributed to patrilocality, shorter average generation
times for females than males, and/or asymmetric migration over time, from Northeast Asia to Hokkaido. Taking
the recent suggestion of the morphological affinities
between the Paleo-Indians from North America and the
Ainu into account, the present findings shed light on a
model in which Northeast Asian antecedents with more
generalized craniofacial features similar to the prehistoric Jomon people and the recent Ainu entered the New
World continent in late Pleistocene and early Holocene
times.
ACKNOWLEDGMENTS
We wish to express our sincere thanks to T. Amano of
Hokkaido University Museum, Sapporo; M. Matsumura
of the Department of Anatomy, Sapporo Medical University, Sapporo; Y. Dodo of the Department of Nursing,
Hokkaido Bunkyo University, Eniwa; G. Suwa of the
Department of Anthropology, University Museum, The
University of Tokyo, Tokyo; H. Baba and Y. Mizoguchi of
the Department of Anthropology, National Museum of
Nature and Science, Tokyo; K. Katayama of the Depart-
291
ment of Zoology, Kyoto University, Kyoto; for their kind
permission to study the materials under their care.
LITERATURE CITED
Amano T. 2003. What is the Okhotsk culture? In: Nomura T,
Utagawa Y, editors. Epi-Jomon and Okhotsk culture. Sapporo:
Hokkaido Sinbun-sha. p 110–133 (in Japanese).
Aroutiounov S. 1968. Ainu problem viewed by Russian and
Soviet Science. Proc VIIIth Interntl Cong Anthropol Ethnol
Sci,Vol III, p 107–109.
Bonatto SL, Salzano FM. 1997. Diversity and age of the four
major mtDNA haplogroups, and their implications for the peopling of the New World. Am J Hum Genet 61:1413–1423.
Brace CL, Hunt KD. 1990. A nonracial craniofacial perspective
on human variation: A(ustralia) to Z(uni). Am J Phys Anthropol 82:341–360.
Brace CL, Brace ML, Leonard WL. 1989. Reflections on the face
of Japan: a multivariate craniofacial and odontometric perspective. Am J Phys Anthropol 78:93–113.
Brace CL, Nelson AR, Seguchi N, Oe H, Sering L, Pan Q, Li Y,
Tumen D. 2001. Old World sources of the first New World
human inhabitants: a comparative craniofacial view. Proc
Natl Acad Sci USA 98:10017–10022.
Brace CL, Seguchi N, Quintyn CB, Fox SC, Nelson AR, Manolis
SK. 2006. The questionable contribution of the Neolithic and
the Bronze age to European craniofacial from. Proc Natl Acad
Sci USA 103:242–247.
Bräuer G. 1988. Osteometrie: a. Kraniometrie. In: Knußmann
R, editor. Anthropologie: Handbuch der Vergleichenden Biologie des Menschen, Band I. Stuttgart: Gustav Fisher. p 160–
192.
Busk G. 1867. Description of an Ainu skull. Trans Ethnol Soc
NS 6:109–112.
Carson EA. 2006. Maximum likelihood estimation of human craniometric heritabilities. Am J Phys Anthropol 131:169–180.
Crawford MH, Devor EJ. 1980. Population structure and admixture in transplanted Tlaxcaltecan populations. Am J Phys
Anthropol 52:485–490.
Davis JB. 1870. Description of the skeleton of an Ainu woman,
and of three skulls of men of the same race. Mem Anthropol
Soc 3:21–40.
Devor EJ. 1987. Transmission of human craniofacial dimensions. J Craniofacial Genet Dev Biol 7:95–106.
Dodo Y, Ishida H. 1990. Population history of Japan as viewed
from cranial nonmetric variation. J Anthropol Soc Nippon
98:269–287.
Dodo Y, Kawakubo Y. 2002. Cranial affinities of the Epi-Jomon
inhabitants in Hokkaido. Japan. Anthropol Sci 110:1–32.
Donnelly SM, Konigsberg LW. 1998. Interpretation of population
structure when group structure is unknown. Am J Phys
Anthropol Suppl 26:106.
Fagundes NJR, Kanitz R, Eckert R, Valls ACS, Bogo MR, Salzano FM, Smith DG, Silva WA II, Zago MA, Ribeiro-dos-Santos AK, Santos SEB, Petzl-Erler ML, Bonatto SL. 2008. Mitochondrial population genomics supports a single pre-Clovis
origin with a coastal route for the peopling of the Americas.
Am J Hum Genet 82:583–592.
Fitzhugh WW, Dubreuil CO. 1999. Ainu: spirit of a northern
people. Washington DC: National Museum of Natural History.
González-José R, Dahinten SL, Luis MA, Hernández M, Pucciaelli
M. 2001. Craniometric variation and the settlement of the
Americas: testing hypotheses by means of R-matrix and matrix
correlation analyses. Am J Phys Anthropol 116:154–165.
González-José R, González-Martin A, Hernández M, Pucciarelli
HM, Sardi M, Rosales A, Molen SV. 2003. Craniometric evidence for Palaeoamerican survival in Baja California. Nature
425:62–65.
González-José R, Neves W, Lahr MM, González S, Pucciarelli H,
Martı́nez MH, Correal G. 2005. Late Pleistocene/Holocene craniofacial morphology in Mesoamerican Paleoindians: implica-
American Journal of Physical Anthropology
292
T. HANIHARA ET AL.
tions for the peopling of the New World. Am J Phys Anthropol
128:772–780.
Hallgrı́msson B, Donnabháin ÓB, Walters GB, Cooper DML,
Gubjartsson D, Stefánsson K. 2004. Composition of the founding population of Iceland: biological distance and morphological variation in early historic Atlantic Europe. Am J Phys
Anthropol 124:257–274.
Hammer MF, Horai S. 1995. Y chromosomal DNA variation and
the peopling of Japan. Am J Hum Genet 46:115–125.
Hanihara K. 1987. Estimation of the number of early migrants
to Japan: a simulative study. J Anthropol Soc Nippon 95:391–
403.
Hanihara K. 1991. Dual structure model for the population history of the Japanese. Jpn Rev 2:1–33.
Hanihara K. 1998. Reanalysis of local variations in the Ainu
crania. Anthropol Sci 106(Suppl):1–15.
Hanihara T. 2006. Interpretation of craniofacial variation and
diversification of East and Southeast Asians. In: Oxenham
M, Tayles N, editors. Bioarchaeology of Southeast Asia. Cambridge: Cambridge University Press. p 91–111.
Hanihara T. 2008. Morphological variation of major human populations based on nonmetric dental traits. Am J Phys Anthropol, in press. DOI 10.1002/ajpa.20792.
Hanihara K, Hanihara T. 1989. Multivariate analysis of tooth
crown morphology in Japanese-American F1 hybrids. Hum
Evol 4:417–427.
Hanihara T, Ishida H, Dodo Y. 1998. Place of the Hokkaido
Ainu (northern Japan) among circumpolar and other peoples of the world: a comparison of the frequency variations
of discrete cranial traits. Int J Circumpolar Health 57:257–
275.
Helgason A, Hrafnkelsson B, Gulcher JR, Ward R, Stefánnson
K. 2003. A populationwide coalescent analysis of Icelandic
matrilineal and patrilineal genealogies: evidence for a faster
evolutionary rate of mtDNA lineages than Y chromosomes.
Am J Hum Genet 72:1370–1388.
Horai S, Murayama K, Hayasaka K, Matsubayashi S, Hattori Y,
Fucharoen G, Harihara S, Park KS, Omoto K, Pan IH. 1996.
mtDNA polymorphism in East Asian populations, with special
reference to the peopling of Japan. Am J Hum Genet 59:579–
590.
Howells WW. 1959. Mankind in the making. New York: Doubleday & Company.
Howells WW. 1973. Cranial variation in man: a study by multivariate analysis of patterns of difference among recent human
populations. Papers of the Peabody Museum of Archaeology
and Ethnology 67. Cambridge: Harvard University.
Howells WW. 1986. Physical anthropology of the prehistoric
Japanese. In: Pearson RJ, editor. Windows on Japanese
past: studies in archaeology and prehistory. Ann Arbor:
Center for Japanese Studies, The University of Michigan. p
85–99.
Howells WW. 1989. Skull shapes and the man: craniometric
analyses in the dispersion of modern Homo. Papers of the
Peabody Museum of Archaeology and Ethnology 79. Cambridge: Harvard University.
Howells WW. 1995. Who’s who in skulls: ethnic identification
of crania from measurements. Papers of the Peabody Museum
of archaeology and Ethnology 82. Cambridge: Harvard
University.
Hubbe M, Neves WA, Amaral HL, Guidon N. 2007. ‘‘Zuzu’’
strikes again—morphological affinities of the early Holocene
human skeleton from Toca dos Coqueiros, Piaui, Brazil. Am J
Phys Anthropol 134:285–291.
Hudson MJ. 2004. The perverse realities of change: world
system incorporation and the Okhotsk culture of Hokkaido.
J Anthrolpol Archaeol 23:290–308.
Ishida H. 1988. Morphological studies of Okhotsk crania from
Omisaki, Hokkaido. J Anthropol Soc Nippon 96:17–45.
Ishida H. 1994. Skeletal morphology of the Okhotsk people on
Sakhalin Island. Anthropol Sci 102:257–269.
Ishida H. 1995. Nonmetric cranial variation of Northeast Asian
populations and their population affinities. Anthropol Sci
103:385–401.
American Journal of Physical Anthropology
Ishida H. 1996. Metric and nonmetric cranial variation of
the prehistoric Okhotsk people. Anthropol Sci 104:233–
258.
Ishida H, Kida M. 1991. An anthropological investigation of the
Sakhalin Ainu with special reference to nonmetric cranial
traits. J Anthropol Soc Nippon 99:23–32.
Ishida H, Kondo O. 1999. Nonmetric cranial variation of the
Ainu and neighbouring human populations Perspective Hum
Biol 4:127–138.
Jantz RL, Owsley DW. 2001. Variation among early North
American crania. Am J Phys Anthropol 114:146–155.
Jantz RL, Owsley DW. 2003. Reply to Van Vark et al.: is European upper Paleolithic cranial morphology a useful analogy
for early Americans? Am J Phys Anthropol 121:185–188.
Kodama S. 1970. Ainu: historical and anthropological studies.
Sapporo: Hokkaido University School of Medicine.
Koganei Y. 1893. Beiträge zur physischen Anthropologie der
Ainu. I. Untersuchungen am Skelet. Mittheilungen aus der
medicinischen Facultät der Kaiserlich-Japanischen Universität 2:1–249.
Komesu A, Hanihara T, Amano T, Ono H, Yoneda M, Dodo Y,
Fukumine T, Ishida H. 2008. Nonmetric cranial variation in
human skeletal remains associated with Okhotsk culture.
Anthropol Sci 116:33–47.
Kondo O. 1995. An analysis of Ainu population structure, based
on cranial morphology. Anthropol Sci 103:369–384.
Kondo O. 2005. Regional diversity of the Ainu cranial morphology was caused by influences from the Okhotsk cultural people. In: Archaeological issue publication team on the maritime
cultural exchange, editors. Sea and archaeology. Tokyo: Rokuichi-Shobo. p 233–242 (in Japanese).
Kozintsev A. 1990. Ainu, Japanese, their ancestors and neighbours: cranioscopic data. J Anthropol Soc Nippon 247–267.
Kozintsev A. 1992. Prehistoric and recent populations of Japan:
multivariate analysis of cranioscopic data. Arctic Anthropol
29:104–111.
Kozintsev A. 1999. Collateral relatives of American Indians
among the Bronze age populations of Siberia? Am J Phys
Anthropol 108:193–204.
Lahr MM. 1995. Pattern of modern human diversification:
implications for Amerindian origins. Yrbk Phys Anthropol
38:163–198.
Lahr MM. 1996. The evolution of modern human diversity: a study
of cranial variation. Cambridge: Cambridge University Press.
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.
Nei M. 1995. The origins of human populations: genetic, linguistic, and archeological data. In: Brenner S, Hanihara K,
editors. The origin and past of modern human as viewed from
DNA. Singapore: World Scientific. p 71–91.
Neves WA, Hubbe M. 2005. Cranial morphology of early
Americans from Lagoa Santa, Brazil: implications for the
settlement of the New World. Proc Natl Acad Sci USA 102:
18309–18314.
Neves WA, Pucciarelli HM. 1991. The origin of the first Americans: an analysis based on the cranial morphology of early
South American human remains. J Hum Evol 21:261–273.
Neves WA, Hubbe M, Correal G. 2007a. Human skeletal
remains from Sabana de Bogotá, Colombia: a case of Paleoamerican morphology late survival in South America? Am J
Phys Anthropol 133:1080–1098.
Neves WA, Hubbe M, Piló LB. 2007b. Early Holocene human
skeletal remains from Sumidouro Cave, Lagoa Santa, Brazil:
history of discoveries, geological and chronological context,
and comparative cranial morphology. J Hum Evol 52:16–30.
Ohyi H. 1985. On the process of crystallization of Sakhalin
Ainu. Bull Inst Study North Eurasian Culture Hokkaido Univ
17:165–192.
Omoto K. 1995. Genetic diversity and the origins of the ‘‘Mongoloids.’’ In: Brenner S, Hanihara K, editors. The origin and
past of modern human as viewed from DNA. Singapore:
World Scientific. p 92–109.
CRANIOMETRIC VARIATION OF THE AINU
Omoto K. Saitou N. 1997. Genetic origins of the Japanese: a
partial support for the dual structure hypothesis. Am J Phys
Anthropol 102:437–446.
Oota H, Settheetham-Ishida W, Tiwawech D, Ishida T, Stoneking M. 2001. Nat Genet 29:20–21.
Ossenberg NS, Dodo Y, Maeda T, Kawakubo Y. 2006. Ethnogenesis and craniofacial change in Japan from the perspective of
nonmetric traits. Anthropol Sci 114:99–115.
Pietrusewsky M. 2000. Metric analysis of skeletal remains:
methods and applications. In: Katzenberg MA, Saunders SR,
editors. Biological anthropology of the human skeleton. New
York: Wiley-Liss. p 375–415.
Powell JF, Neves WA. 1999. Craniofacial morphology of the first
Americans: pattern and process in the peopling of the New
World. Yrbk Phys Anthropol 42:153–188.
Relethford JH. 1991. Genetic drift and anthropometric variation
in Ireland. Hum Biol 63:155–165.
Relethford JH. 1994. Craniometric variation among modern
human populations. Am J Phys Anthropol 95:53–62.
Relethford JH. 1996. Genetic drift can obscure population history: problem and solution. Hum Biol 68:29–44.
Relethford JH. 2001. Genetics and the search for modern
human origins. New York: Wiley.
Relethford JH. 2002. Apportionment of global human genetic
diversity based on craniometrics and skin color. Am J Phys
Anthropol 118:393–398.
Relethford JH. 2004. Boas and beyond: migration and craniometric variation. Am J Hum Biol 16:379–386.
Relethford JH, Blangero J. 1990. Detection of differential gene
flow from patterns of quantitative variation. Hum Biol 62:5–25.
Relethford JH, Harpending HC. 1994. Craniometric variation,
genetic theory, and modern human origins. Am J Phys
Anthropol 95:249–270.
Roseman CC, Weaver TD. 2004. Multivariate apportionment of
global human craniometric diversity. Am J Phys Anthropol
125:257–263.
Sardi ML, Rozzi FR, González-José R, Pucciarelli HM. 2005.
South Amerindian craniofacial morphology: diversity and
implications for Amerindian evolution. Am J Phys Anthropol
2005:747–456.
Sato T, Amano T, Ono H, Ishida H, Kodera H, Matsumura H,
Yoneda M, Masuda R. 2007. Origins and genetic features of
the Okhotsk people, revealed by ancient mitochondrial DNA
analysis. J Hum Genet 52:18–627.
Schurr TG. 2004. The peopling of the New World: perspectives
from molecular anthropology. Ann Rev Anthropol 33:551–583.
Schurr TG, Sukernik RI, Starikovskaya YB, Wallace DC. 1999.
Mitochondrial DNA variation in Koryaks and Itel’man: population replacement in the Okhotsk Sea-Bering Sea region during the Neolithic. Am J Phys Anthropol 108:1–39.
293
Seielstad MT, Minch E, Cavalli-Sforza LL. 1998. Genetic evidence for a higher female migration rate in humans. Nat
Genet 20:278–280.
Shigematsu M, Ishida H, Goto M, Hanihara T. 2004. Morphological affinities between Jomon and Ainu: reassessment based
on nonmetric cranial traits. Anthropol Sci 112:161–172.
Shriver MD. 2005. Female migration rate might not be greater
than male rate. Eur J Hum Genet 13:131–132.
Stojanowski CM. 2004. Population history of native groups in
pre- and postcontact Spanish Florida: aggregation, gene flow,
and genetic drift on the southeastern U.S. Atlantic coast. Am
J Phys Anthropol 123:316–332.
Stoneking M. 1998. Women on the move. Nat Genet 20:219–
220.
Tajima A, Hayami M, Tokunaga K, Juji T, Matsuo M, Marzuki
S, Omoto K, Horai S. 2004. Genetic origins of the Ainu
inferred from combined DNA analyses of maternal and paternal lineages. J Hum Genet 49:187–193.
Torgerson WS. 1952. Multidimensional scaling I. Theory and
method. Psychometrika 17:401–419.
Turner CG II. 1987. Late Pleistocene and Holocene population
history of East Asia based on dental variation. Am J Phys
Anthropol 73:305–321.
Turner CG II. 1990. Major features of sundadonty and
sinodonty, including suggestions about East Asian microevolution, population history, and late Pleistocene relationships
with Australian Aboriginals. Am J Phys Anthropol 82:295–
317.
Underhill PA, Passarino G, Lin AA, Shen P, Lahr MM,
Foley RA, Oefner PJ, Cavalli-Sforza LL. 2001. The phylogeography of Y chromosome binary haplotypes and the
origins of modern human populations. Ann Hum Genet
65:43–62.
Wilder JA, Kingan SB, Mobasher Z, Pilkington MM, Hammer
MF. 2004. Global patterns of human mitochondrial DNA
and Y-chromosome structure are not influenced by higher
migration rates of females versus males. Nat Genet 36:
1122–1125.
Yamaguchi B. 1974. Palaeoanthropology in Hokkaido. Quat Res
12:257–264 (in Japanese).
Yamaguchi B. 1981. Human skeletal remains in Hokkaido. In:
Ogata T, editor. Japanese I. Anthropology 5. Tokyo: Yuzankaku. p 137–156 (in Japanese).
Yamaguchi B. 1982. A review of the osteological characteristics
of the Jomon population in prehistoric Japan. J Anthropol Soc
Nippon 90 (suppl):77–90.
Yoshida K. 2006. Population of the Ainu and the Wajin in Hokkaido before the Meiji restoration. Anthropol Sci 114:255.
Yoshida K. 2007. Estimates of the Ainu population size before
the Meiji restoration. Anthropol Sci 115:266.
American Journal of Physical Anthropology
Документ
Категория
Без категории
Просмотров
2
Размер файла
343 Кб
Теги
flow, craniometric, northern, variation, hokkaido, japan, differential, genes, ainu, assessment, northeast, asia
1/--страниц
Пожаловаться на содержимое документа