Metric dental variation of major human populations.

AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 128:287–298 (2005)
Metric Dental Variation of Major Human Populations
Tsunehiko Hanihara1* and Hajime Ishida2
1
2
Department of Anatomy and Biological Anthropology, Saga Medical School, Saga 849-8501, Japan
Department of Anatomy, Faculty of Medicine, University of the Ryukyus, Nishihara, 903-0215, Japan
KEY WORDS
relationships
odontometrics; phenotypic diversity; geographic variation; interpopulation
ABSTRACT
Mesiodistal and buccolingual crown diameters of all teeth recorded in 72 major human population groups and seven geographic groups were analyzed.
The results obtained are fivefold. First, the largest teeth
are found among Australians, followed by Melanesians,
Micronesians, sub-Saharan Africans, and Native Americans. Philippine Negritos, Jomon/Ainu, and Western Eurasians have small teeth, while East/Southeast Asians and
Polynesians are intermediate in overall tooth size. Second,
in terms of odontometric shape factors, world extremes
are Europeans, aboriginal New World populations, and to
a lesser extent, Australians. Third, East/Southeast Asians
share similar dental features with sub-Saharan Africans,
and fall in the center of the phenetic space occupied by a
wide array of samples. Fourth, the patterning of dental
variation among major geographic populations is more or
less consistent with those obtained from genetic and
craniometric data. Fifth, once differences in population
size between sub-Saharan Africa, Europe, South/West
Asia, Australia, and Far East, and genetic drift are taken
into consideration, the pattern of sub-Saharan African
distinctiveness becomes more or less comparable to that
based on genetic and craniometric data. As such, worldwide patterning of odontometric variation provides an additional avenue in the ongoing investigation of the origin(s) of anatomically modern humans. Am J Phys
Anthropol 128:287–298, 2005. © 2005 Wiley-Liss, Inc.
During the last few decades, many dental anthropologists have focused on nonmetric crown and root traits for
defining the dental characteristics of major geographic
groups, and for reconstructing the population history,
structure, and origin of modern humans (Hanihara, 1966,
1968; Turner, 1983; Irish, 1997, 1998; Scott and Turner,
1997; Stringer et al., 1997; Tyrrell and Chamberlain,
1998; Irish and Guatelli-Steinberg, 2003). On the other
hand, odontometric variation has often been used in the
investigation of hominid evolution and/or diversity of local
and regional population groups (e.g., Brace, 1967, 1980;
Wolpoff, 1971; Hanihara, 1976; Frayer, 1977; Brace and
Hinton, 1981; Brace et al., 1984, 1991; Lukacs and Hemphill, 1991; Hillson, 1996; Lukacs et al., 1998; Bermudez de
Castro et al., 2004). During the past few decades, some
articles on metric dental variation covering a wide range
of regional populations were published (Lasker and Lee,
1957; Falk and Corruccini, 1982; Harris and Bailit, 1988;
Kieser, 1990; Harris and Rathbun, 1991; Schnutenhaus
and Rösing, 1998), and these studies classified contemporary and recent populations as microdontic, mesodontic,
and megadontic (Harris and Rathbun, 1991). However,
few researchers have attempted a systematic analysis of
geographic variation in metric dental traits on a worldwide scale.
Recent genetic studies demonstrated that the majority
of the diversity exists within regional populations, with
much less among large geographic population groups (Lewontin, 1972; Latter, 1980; Barbujani et al., 1997; Jorde et
al., 2000; Brown and Armelagos, 2001). Apart from issues
of among-regional variation, analyses of within-group
variation consistently reveal that highest levels occur
among sub-Saharan Africans (Cann et al., 1987; Stoneking and Cann, 1989; Stoneking, 1993; Bowcock et al.,
1994). It is generally accepted that morphological varia-
tion among recent human populations is greater than
genetic variation (Stringer and Andrews, 1988). Using the
craniometric data of Howells (1973, 1989, 1995), Relethford (1994, 2002) and Relethford and Harpending (1994)
showed that craniometric diversity among major geographic regions is very limited, at ca. 11–14% of the total
variance, and sub-Saharan Africans show the greatest
intraregional variation. Relethford and Harpending
(1994, 1995) showed, moreover, that sub-Saharan Africa
accounted for ca. 50% of the total population size during
recent human evolution using craniometric data. Until
now, however, assessments of odontometric diversity
among and within group variation in major geographical
regions have not been presented.
The present study has two objectives. The first is to
present the odontometric characteristics and interpopulation relationships among human populations of major geographic regions to elucidate how such data contribute to
©
2005 WILEY-LISS, INC.
Grant sponsor: Ministry of Education, Science and Culture in
Japan; Grant numbers: Grants-in-Aid for Scientific Research
1440521, 14540659; Grant sponsor: Japan Fellowship for Research in
United Kingdom, Japan Society for the Promotion of Science; Grant
sponsor: Smithsonian Opportunities for Research and Study, Smithsonian Institution Fellowship Program.
*Correspondence to: Tsunehiko Hanihara, Department of Anatomy
and Biological Anthropology, Saga Medical School, 5-1-1 Nabeshima,
Saga 849-8501, Japan. E-mail: [email protected]
Received 26 July 2003; accepted 1 April 2004.
DOI 10.1002/ajpa.20080
Published online 18 April 2005 in Wiley InterScience
(www.interscience.wiley.com).
288
T. HANIHARA AND H. ISHIDA
our understanding of the patterning of population affinities among modern humans. The second is to present the
estimates of the degree of population differentiation
within, and diversity among, world regions based on odontometric data.
MATERIALS AND METHODS
The principal analyses performed in the present study
are limited to males because of larger sample sizes than in
females, and are based mainly on samples drawn from
countries, tribes, and cultural background. Table 1 provides the sample name, sample size, and brief information
for those samples included in the dataset. Mesiodistal and
buccolingual crown diameters of all teeth (a total of 32
metric variables) were recorded by T.H. to avoid possible
interobserver differences. Using a Japanese male sample,
intraobserver error was tested by t-statistics and found to
be insignificant. Dental measurements were taken according to the procedures of Moorrees (1957) and Hillson
(1996).
Datasets were analyzed in the classic way, with the use
of principal component analysis (PCA) based on the pooled
within-group variance-covariance matrix weighted by
sample sizes. Two separate analyses were performed using raw measurements and C-scores to investigate the
possible effects of size and shape (Howells, 1989; Brace
and Hunt, 1990).
Fst values were calculated for both male and female
samples drawn from broad geographical areas, following
the methods outlined by Relethford and Blangero (1990)
and Relethford (1994) to estimate the relative degree of
dental variation among regional groups. The detailed procedures for calculation were given elsewhere (Relethford,
1991, 1994; Relethford and Blangero, 1990; Relethford
and Harpending, 1994, 1995). Briefly, FST is defined as
F ST ⫽ (⌺w i C ii )/(2t ⫹ ⌺w i C ii ),
where t is the number of traits and wi is the weighting
factors, the relative size of population i, defined as
w i ⫽ N i /⌺N j .
Nj is the effective size of population j. Cii is the diagonal
element of codivergence matrix (C), computed as
C ⫽ ⌬G ⫺ 1⌬ⴕ,
where G⫺1 is the inverse of the pooled within-group additive genetic variance-covariance matrix, and the elements
of ⌬ are the deviations of group means from the total
means averaged over all populations weighted by population size. Calculation of the pooled within-group additive
genetic variance-covariance matrix (G) from the phenotypic variance-covariance matrix (V) requires an estimate
of average heritability for phenotypic traits. According to
Relethford (1994), for any given trait, G ⫽ h2V, where h2
is the heritability of the trait. The lowest possible value of
FST, the minimum FST, is given when h2 ⫽ 1 (G ⫽ V). The
actual Fst, or estimated Fst, requires the estimates of
average heritability of odontometric traits. Estimates of
standard errors for FST are approximated by the jackknife
method (Miller, 1974; Relethford et al., 1997).
Many investigators reported relatively high heritability
rates for dental dimensions based on family and twin
studies (Garn et al., 1968; Goose 1971; Alvesalo and Tigerstedt, 1974; Potter et al., 1976; Townsend and Brown,
1978; summarized by Hillson, 1996). Mizoguchi (1977)
reported the heritabilities of mesiodistal crown diameters
in permanent dentitions of modern Japanese as ranging
from 0.07–1.00 with an average of 0.81 in males, and from
0.41–1.00 with an average of 0.74 in females. Townsend
and Brown (1978) estimated heritability valued to average
0.54 ⫾ 0.23 for mesiodistal dimensions and 0.68 ⫾ 0.27 for
buccolingual dimensions in Australians. In these studies,
however, the upper and lower third molars were not included. As heritability rate (h2) increases, C decreases, so
does Fst. In this study, therefore, we tentatively choose an
estimate of average heritability for odontometric dimension of 0.55 based on Townsend and Brown’s (1978) mesiodistal estimation to avoid the underestimate of actual
Fst values (intraregional variation).
We employed the R matrix approach developed by
Relethford and Blangero (1990) to estimate interpopulation relationships. The R matrix is given as
R ⫽ C(1 ⫺ F ST )/2t.
The elements of an R matrix are converted into distances:
d ij2 ⫽ r ii ⫹ r jj ⫺ 2r ij .
The distance matrix controlled for effective population
size and the potential effects of genetic drift on the genetic
distance
is
obtained
from
scaled
R
matrix. That is,
S ⫽ W 1/2RW 1/2,
where W is a diagonal matrix with relative size wi for the
ith population. The effect of this transformation is to
change the relationship rij between populations i and j to
rij(wiwj)1/2 (Relethford and Harpending, 1994, 1995).
RESULTS
The results of PCA applied to raw measurements are
given in Table 2, which summarizes the eigenvectors and
proportions of total variation accounted for by each of the
first three principal components (PCs) with eigenvalues
greater than 0.50. The first PC, which is only one factor
with eigenvalues greater than 1.0, represents overall
tooth size, and this factor plays a major role in odontometric variation measured by raw data. Variation in the overall dental size assessed by the first PC scores is shown in
Figure 1. The Philippine Negrito, Jomon/Ainu, and Western Eurasian samples are characterized by small dentition, and the Australian samples prove to have the largest
teeth, followed by the Melanesian, Micronesian, sub-Saharan African, and New World samples, with East/Southeast Asians and Polynesians in between.
Table 3 shows the results of PCA based on the C-score.
The first three PCs which account for just over half
(56.7%) of the total variance can be interpreted as representing the relative size of mesiodistal crown diameters
against buccolingual diameters, that of molar teeth
against other teeth, and that of anterior teeth against
premolars, respectively.
Figure 2 provides scattergrams of the first and second
(Fig. 2a) as well as first and third (Fig. 2b) PC scores in
each sample. In Figure 2a, the shape factors of the dental
measurements identify separate phenotypic relationships,
with the Native American, Arctic, and Northeast Asian
samples having relatively large mesiodistal crown diameters, in contrast to Polynesian and Western Eurasian samples with narrow mesiodistal crown diameters. Polynesian
and Western Eurasian samples are separated by the second PC representing the relative size of molars. With the
exception of Jomon and Ainu, East and Southeast Asian
DENTAL VARIATION OF MAJOR HUMAN POPULATIONS
289
TABLE 1. Materials used and brief information1
Sample name
N
Information
East Asia
1. Japanese
2. Ainu
47–50
25–77
3. Jomon
20–77
4. Northern Chinese
19–53
5. Southern Chinese
12–58
6. Tibetans/Nepalese
Northeast Asia
7. Mongol/Buryats
11–77
Tohoku and Kanto regions, Northern part of Honshu (Univ. of Tokyo)
Mainly from Hokkaido, including a few specimens from Sakhalin Island
(Univ. of Tokyo)
Prehistoric (Neolithic) Japan, Middle to the latest Jomon periods (5,300–2,300
years B.P.) (Univ. of Tokyo, National Sci. Museum, Tokyo)
Chinese from north of Cheng Kiang River (Univ. of Tokyo, American Museum
of Natural History)
Chinese from south of Cheng Kiang River (Natural History Museum, Musée
de l’Homme, American Museum of Natural History)
Tibet and Nepal (lowland) (Natural History Museum, Cambridge Univ.)
27–95
8. Chukchis/NE Asia
6–18
Southeast Asia
9. Vietnam/Thailand
21–130
10. Myanmar
14–90
11. Malay
11–55
12. Early SE Asia
27–60
13. Andaman/Nicobar
14–67
14. Java/Sumatra
27–88
15. Borneo
11–76
16. Philippines
26–89
17. Lesser Sunda
18–51
18. Negritos
13–37
Arctic
19. Aleutian Islands
20. Eskimos
North America
21. Northwest Coast
22. Plateau/Great Basin
21–51
17–125
13–86
9–27
23. California
14–53
24. Southwest
20–40
25. Plains/North
31–52
26. Plains/South
11–35
27. Northeastern Woodland/West
21–38
28. Northeastern Woodland/East
18–40
29. Southeastern Woodland
38–55
Ulan Bator, Mongol/Troiskosavsk, Buryats (Musée de l’Homme, Natural
History Museum, American Museum of Natural History, National Museum
of Natural History)
Mainly Chukchis, including a few samples of Evenki Yukagirs and Yakuts
(Musée de l’Homme, American Museum of Natural History, National
Museum of Natural History)
Saigon, Bangkok; Vietnam, and Thailand (Musée de l’Homme, Natural
History Museum, American Museum of Natural History)
Arakan Hill, Rangoon, Karen; Myanmar (Natural History Museum,
Cambridge Univ.)
Mainly from Singapore (Natural History Museum, American Museum of
Natural History)
Mesolithic Malay, Gua Cha; Neolithic Vietnam and Laos; Bang Chang site,
early Iron age, Thailand (Cambridge Univ., Musée de l’Homme, Univ. of
Hawaii)
Recent Andamanese (Onge, Jarawa, etc.) and Nicobarese (Natural History
Museum, Cambridge Univ.)
Including Nias, Billiton, and Mentawei Islands, Batavia, Madura, Fagal,
Pekalongan (Natural History Museum, Cambridge Univ., American
Museum of Natural History, Musée de l’Homme)
Mainly Land Dayaks, including Iban, Sea Dayaks (Natural History Museum,
Cambridge Univ., Musée de l’Homme)
Non-Negrito Filipinos from Luzon, Palawan, Panay, Negroes, and Mindanao
(Natural History Museum, Cambridge Univ., Musée de l’Homme)
Bali, Sumbawa, Flores, and Timor Islands and Celebes and Molucca Island
chain (Natural History Museum, Cambridge Univ., Musée de l’Homme,
American Museum of Natural History)
Mainly Aeta, Luzon, Philippines (Univ. of Tokyo, Natural History Museum,
Musée de l’Homme, National Museum of Natural History)
Unalaska, Unga, Umnak, Shiplock, Amaknak, Kagamil, Atka, Kanaga, Amlia,
Adak, Amchitka, Kiska, Attu, Agatu Islands (American Museum of Natural
History, National Museum of Natural History)
Lower Yukon River basin, Seward Peninsula, Point Barrow around Baffin
Bay, Somerset Island, Prince of Wales, Victoria Island, Greenland (Natural
History Museum, American Museum of Natural History, National Museum
of Natural History)
Tlingit and Bancouver Island (American Museum of Natural History,
National Museum of Natural History, Natural History Museum)
Washington, Oregon, Utah, Colorado, Wyoming, and Nevada (National
Museum of Natural History, American Museum of Natural History)
Sacramento Angel Island, San Nicolas Island, Angeles Bay, Santa Barbara,
Santa Rosa Island, and Santa Cruz Island (National Museum of Natural
History, Musée de l’Homme)
Arizona and New Mexico (American Museum of Natural History, National
Museum of Natural History)
North and South Dakota, Montana, Iowa, and Nebraska (National Museum of
Natural History)
Kansas, Missouri, Arkansas, Texas, and Oklahoma (National Museum of
Natural History)
Illinois, Michigan, Wisconsin, Indiana, and Ohio (National Museum of
Natural History)
New York, Massachusetts, Connecticut, New Jersey, Pennsylvania, Delaware,
Maryland, and Kentucky (American Museum of Natural History, National
Museum of Natural History)
Virginia, Tennessee, Georgia, Alabama, North and South Carolina, Louisiana,
Mississippi, and Florida (National Museum of Natural History)
(continued)
290
T. HANIHARA AND H. ISHIDA
TABLE 1. (Continued)
Sample name
Central/South America
30. Mexico
31. Peru
32. Fuegians/Patagonians
Australia
33. Australia/East
34. Australia/South
35. Australia/North and West
Melanesia
36. Papua New Guinea
N
Information
12–78
Mainly recent Mexico Indians, including Central Cultural region (American
Museum of Natural History, National Museum of Natural History)
Recent Peruvians (Natural History Museum, National Museum of Natural
History)
Terra del Fuego and Patagonia region (Natural History Museum, Cambridge
Univ., American Museum of Natural History, National Museum of Natural
History)
112–115
10–38
19–68
54–155
12–36
11–156
Queensland, New South Wales, and Victoria (Australian Museum, American
Museum of Natural History, Natural History Museum, National Museum
of Natural History, Cambridge Univ., Musée de l’Homme)
Near Adelaide and Murray River Basin (South Australian Museum)
Northern Territory and Western Australia (American Museum of Natural
History, Natural History Museum, National Museum of Natural History,
Cambridge Univ., Musée de l’Homme)
Purari River, Fly River, Sepik River, including Torres Strait (Australian
Museum, Univ. of Sydney, South Australian Museum, Natural History
Museum)
Bismarck Archipelago, New Ireland, New Britain (Australian Museum, Univ.
of Sydney, South Australian Museum, American Museum of Natural
History, National Museum of Natural History, Natural History Museum)
Solomon, New Caledonia, Vanuatu, and Fiji (Australian Museum, Univ. of
Sydney, South Australian Museum, American Museum of Natural History,
National Museum of Natural History, Natural History Museum, Bishop
Museum)
37. West Melanesia
24–105
38. East Melanesia
21–148
Micronesia
39. Mariana Islands
40. Caroline Islands
35–75
17–33
Guam, Saipan, and Tinian Islands (Bishop Museum, Musée de l’Homme)
Yap, Palau, and Ponape Islands (American Museum of Natural History, Univ.
of Tokyo)
Polynesia
41. Society
42. Cook
15–48
20–54
Mainly from Tahiti (Musée de l’Homme, Natural History Museum)
Manuai, Atiu, Titiaroa, and Mangaia Islands (Natural History Museum,
Kyoto Univ., Musée de l’Homme)
New Zealand and Chatham Islands (Australian Museum, Univ. of Sydney,
South Australian Museum, Natural History Museum, Cambridge Univ.,
American Museum of Natural History, National Museum of Natural
History)
Hatua Tua, Nuku Hiva, Ua Huka, and Fatu Hiva (Bishop Museum, Natural
History Museum, Musée de l’Homme, American Museum of Natural
History)
Mainly from Mokapu site, Oahu (Bishop Museum, Cambridge Univ.)
Including a few samples from Lanai and Kauai (Bishop Museum)
Kona site, H1 site; Hawaii Island (Bishop Museum)
43. Maori/Moriori
15–119
44. Marquesas
15–71
45. Oahu/Hawaii
46. Maui-Molokai/Hawaii
47. Hawaii/Hawaii
South Asia (India)
48. India/North
49. India/South
West Asia
50. Afghanistan/Pakistan
51. Turkey/Cyprus
52. Early Iran/Israel
48–123
21–50
16–40
26–169
13–57
9–44
7–42
21–89
Bengal, Behar, Orissa, Bombay, Punjab, and Kashmir (Natural History
Museum, American Museum of Natural History)
Madras, Malabar, and Mysore (Natural History Museum, American Museum
of Natural History)
Helmand Valley; Mogul, Kandahar; Kotal; Priest; Pecheen Valley; Kalati;
Pamir Plat., including a few specimens from West Asia (Natural History
Museum, American Museum of Natural History)
Adevis, Marash; Aintab; Adalea; Ephesos: Kurd, Mohammedan, Armenian;
Hellenismic period of Cyprus (Natural History Museum, American Museum
of Natural History)
Tell Duweir (Lachish), Bronze and Iron Age; Israel/Bronze to Achaemenian
period (6–4 centuries B.C.) of Iran (Natural History Museum, Univ. of
Tokyo)
North Africa
53. Egypt/Predynasty
54. Egypt/12–29th Dynasty
17–63
12–176
Badari and Naqada, ca. 5,000–4,000 B.P. (Cambridge Univ.)
Lisht, Cairo, Omdurman, and Gizeh (Cambridge Univ., National Museum of
Natural History)
Europe
55. Greek
11–28
Ancient and recent Greece; Cyrene, Rhodes, Corfu, Athens, Attica (Natural
History Museum)
Crimea, Courland, Latvia, Pskoff, Uzbekistan, Odessa, Tambow (Natural
History Museum, National Museum of Natural History)
Charvaty, near Olomouc, Central Moravia; including a few samples from East
Europe (Poland, Yugoslavia, Herzegovina) (South Australian Museum,
Natural History Museum)
(continued)
56. Russia
12–38
57. Czecho
12–54
DENTAL VARIATION OF MAJOR HUMAN POPULATIONS
291
TABLE 1. (Continued)
Sample name
N
58. Hungary
12–40
59. Germany
24–46
60. Austria/Swiss
10–40
61. Italy
62. France
26–93
12–56
63. Finland/Ural
10–72
64. UK/Ensay
65. UK/Poundbury
66. UK/Spitalfields1
67. UK/Spitalfields2
Sub-Saharan Africa
68. Nigeria/Ghana
69. Somalia
70. Kenya
36–59
47–95
10–64
33–54
34–259
13–59
19–77
71. Tanzania
13–94
72. South Africa
26–86
Information
Demko-Hegy; Czakbereny, Roman Period; Demko Hegy, 11–12th centuries;
Nagy-sap, Csakover, 11th century; Stuhlweisenberg; Pressburg (Natural
History Museum, American Museum of Natural History)
Berlin, Mainz, Holstein, Westphalia (Natural History Museum, National
Museum of Natural History)
Tirol; St. Peter above Tweng, Carinthia, 16–17th centuries; Malta, Millstatt,
Carinthia; Raach, near Gloggnitz; Modling, near Vienna; Vienna/Austria;
Lenz Canton, Grisons/Switzerland (American Museum of Natural History
National Museum of Natural History)
Ancient and recent Italians (Natural History Museum)
Merovingian, Gallo-Roman, Brittany times; Paris, etc. (Natural History
Museum)
Recent Finns, including a few specimens of Ural language people (Natural
History Museum)
Late Medieval to Postmedieval periods, Scotland (Natural History Museum)
Late Roman period, Southwest England (Natural History Museum)
Mid-Victorian period, London (Natural History Museum)
Pre-17th century, London (Cambridge Univ.)
Ibo and Ashanti tribes (Natural History Museum, Cambridge Univ.)
Erigavo District, Ogaden Somali (Cambridge Univ.)
Ngorongoro, Turkana, Kikuyu, Kaurite to Fort Hall (Cambridge Univ.,
Natural History Museum, Cambridge Univ., American Museum of Natural
History)
Tanganika Territory, Loe, Haya Tribe (Natural History Museum, Cambridge
Univ., American Museum of Natural History)
Zulu and once called Kaffirs (Natural History Museum, Cambridge Univ.,
American Museum of Natural History, National Museum of Natural
History)
1
N, range of N in samples (since each of traits observed has a different N); Univ., University; Sci., Science; Univ. of Tokyo, Tokyo,
Japan; Kyoto Univ., Kyoto, Japan; National Sci. Museum, Tokyo, Japan; American Museum of Natural History, New York; National
Museum of Natural History, Washington, DC; Bishop Museum, Honolulu; Univ. of Hawaii, Honolulu; Natural History Museum,
London, UK; Cambridge Univ., Cambridge, UK; Musée de l’Homme, Paris, France; Australian Museum, Sydney, Australia; Univ. of
Sydney, Sydney, Australia; South Australian Museum, Adelaide, Australia.
samples together with the Melanesian and Micronesian
samples share similar characteristics with each other and
with sub-Saharan African samples. Australian, Jomon,
and Indian Subcontinent samples occupy a position intermediate between New World and Western Eurasian samples. The Ainu sample is plotted at the near position of
Northeast Asian samples. The Australian samples show
distinctive dental features in the third PC representing
relatively large anterior teeth.
Among-group
dental
variation
estimated
by
Fst was computed for five Old World samples (Far
East, Australia, South/West Asia, Europe, and
sub-Saharan Africa) and seven worldwide samples including Pacific and New World in both male and female datasets. The geographic groups used and the number of individuals for each sex are provided in Table 4. Table 5 gives
the minimum Fst values and estimated Fst values using
an average heritability of 0.55. The Fst values obtained
reveal that extensive odontometric diversity exists within
regional populations, with much less diversity between
regional populations.
Table 6 gives the results of the method of Relethford and
Blangero (1990) for estimating intraregional variations
under the assumption of equal effective population size.
Intraregional variation is largest in the sub-Saharan African group, followed closely by the Australian group. The
New World and European groups, on the other hand, show
small intraregional variation.
Using the estimated Fst values, the R matrix for the five
Old World samples is calculated. The New World and
Pacific samples are excluded, since they represent newly
diverged populations (Relethford and Harpending, 1994).
Figure 3a illustrates the interpopulation relationships derived from the first two eigenvectors of the R matrix
scaled by the square root of the corresponding eigenvalues, assuming equal effective population sizes in the five
geographical samples. Among males, the Australian sample is distinctive, while sub-Saharan African and Far East
samples appear close. On the other hand, Australian and
sub-Saharan African samples are distinctive for females.
Relethford and Harpending (1994, 1995) pointed out that
it is necessary to control for variation in population size
for reconstructing population relationships without the
effect of different degrees of genetic drift. Following the
suggestion of Relethford and Harpending (1994) that the
population of sub-Saharan Africa was three times that of
any of the other geographical regions, a scaled R matrix
was calculated whereby Far East, Australia, South/West
Asia, and Europe samples were weighted as one each and
sub-Saharan Africa as three, to account for relative effective population size. Figure 3b was drawn using the scaled
eigenvectors of the scaled R matrix. In both sexes, interpopulation relationships are similar to those obtained by
unscaled R matrix analysis, in which the Australian sample stands out as distinctive. However, the sub-Saharan
African sample moves more or less away from the Far
East sample in males.
DISCUSSION
The results of this study confirm the trifurcation of
contemporary and recent populations into those that are
microdontic, mesodontic, and megadontic (Harris and
Rathbun, 1991). It was asserted for several decades that
292
T. HANIHARA AND H. ISHIDA
TABLE 2. Eigenvalues and eigenvectors of variance-covariance
matrix by PCA, based on raw measurement data
Tooth
UI1-MD
UI2-MD
UC-MD
UP1-MD
UP2-MD
UM1-MD
UM2-MD
UM3-MD
LI1-MD
LI2-MD
LC-MD
LP1-MD
LP2-MD
LM1-MD
LM2-MD
LM3-MD
UI1-BL
UI2-BL
UC-BL
UP1-BL
UP2-BL
UM1-BL
UM2-BL
UM3-BL
LI1-BL
LI2-BL
LC-BL
LP1-BL
LP2-BL
LM1-BL
LM2-BL
LM3-BL
Eigenvalues
Proportion
Cumulative proportion
PC-1
PC-2
PC-3
0.16934
0.16405
0.18002
0.18397
0.18992
0.18779
0.19168
0.17043
0.17068
0.17968
0.18390
0.18213
0.18600
0.18651
0.19132
0.18675
0.16763
0.16277
0.17656
0.17897
0.18043
0.19254
0.18785
0.16091
0.12138
0.13628
0.15836
0.16610
0.16733
0.18717
0.19356
0.19159
4.40350
0.43823
0.43823
⫺0.16099
⫺0.29241
⫺0.18091
⫺0.09076
⫺0.10830
⫺0.14915
⫺0.09757
⫺0.08201
⫺0.20374
⫺0.21945
⫺0.17263
0.02432
0.03953
⫺0.16409
⫺0.07491
0.01090
⫺0.01470
⫺0.00779
0.14221
0.11318
0.21718
⫺0.01858
0.13670
0.23574
0.35629
0.36690
0.17372
0.28941
0.30720
⫺0.12776
⫺0.01166
0.04129
0.90329
0.08989
0.52812
⫺0.19786
0.06159
0.08594
0.33084
0.16924
⫺0.10561
⫺0.07744
⫺0.27485
⫺0.01214
⫺0.06007
0.07390
0.34568
0.22341
0.09128
⫺0.11603
⫺0.15608
⫺0.20763
⫺0.05580
⫺0.20498
0.35693
0.17422
⫺0.14932
⫺0.14911
⫺0.13629
⫺0.24713
⫺0.08257
⫺0.12772
0.26393
0.15686
⫺0.03258
⫺0.07386
0.00403
0.56595
0.05632
0.58445
tooth size has been exposed to natural selective forces
beginning in the Upper Paleolithic (Frayer, 1978; Calcagno and Gibson, 1988, 1991). Macchiarelli and Bondioli
(1986) suggested the increasing population density effect
on tooth size reduction. Brace (1967) and Brace et al.
(1984, 1991) argued that tooth size reduction is a consequence of what they called the probable mutation effect,
i.e., large tooth size is maintained by strong natural selection, but reduction in tooth size occurs in the absence of
such pressures in relation to the development of sophisticated food preparation techniques and pottery use.
As expected, modern Australian Aboriginals possess the
least reduced teeth, followed by Melanesians, sub-Saharan Africans, and Native Americans. Together with the
Philippine Negritos, the most extensive dental size reduction occurs in populations occupying the western part and
eastern edge of the Eurasian continent, i.e., Europeans
and the Jomon/Ainu of Japan. Such results are in agreement with the relationship between tooth size reduction
and cultural background among recent populations of the
world found by Brace et al. (1991). It is well-known that
modern Europeans and the Ainu are living descendents of
the first people to use cooking in the preparation of food
and to use pottery (Brace et al., 1991). The small tooth size
observed in Negritos is likely a byproduct of an overall
diminutization of body size (Hillson, 1996).
When the allocation of relative tooth size across all 32
variables forms the basis of comparison, New World populations, including occupants of the Arctic, Polynesians, Europeans, and to a lesser extent, Australians, form the extremes
of contemporary worldwide variation. East/Southeast
Asians, Melanesians, and the Indian Subcontinent, together
with sub-Saharan Africans, form an average group. Such
findings are more or less consistent with those obtained by
Scott and Turner (1997) from nonmetric dental traits. In
Figure 2, the geographical groups with relatively large intraregional variation (Table 6) are plotted at a central position, and clinal variation of the traits is recognized. Such
results may be consistent with the center and edge model
proposed by Thorne and Wolpoff (1981), according to which
phenotypic variation is expected to be greatest at the geographical center of a species range, and lower in peripheral
populations.
The present findings (Fig. 2) do not support the division
of East Asian populations into sinodonts and sundadonts
proposed (Turner, 1987, 1990; Scott and Turner, 1997)
from analyses of nonmetric dental traits. Turner (1987,
1989, 1990, 1992) designated the major populations of
China and Japan as sinodonts, and suggested that these
populations were more similar in dental trait frequencies
to Northeast Asians and New World populations than to
the sundadont populations of Southeast Asia and the Pacific. Turner (1990) identified the prehistoric Jomon and
recent Ainu of the Japanese Archipelago as sundadont
populations residing as isolated pockets within the range
of sinodonts.
The patterning of phenetic affinities of East Asians
based on relative dental size is profoundly different from
those obtained from comparisons of morphological trait
frequencies. The results of this study indicate that Chinese and Japanese populations are more like Southeast
Asian populations, while the prehistoric Jomon show affinities with Australian and Papuan populations. In direct
opposition to results from dental morphological studies,
patterning of relative tooth size among the Ainu shows
similarities to that seen among Northeast Asian and New
World populations.
Recently, genetic affinities between Ainu and East/
Northeast Asians and morphological similarities between
Ainu and Paleoindians were suggested (Cavalli-Sforza et
al., 1988; Nei, 1995; Omoto, 1995; Omoto and Saitou,
1997; Brace et al., 2001). Unfortunately, we cannot confirm this Ainu-Paleoindian connection from results obtained in the current study. This is because an ancestordescendent relationship between Early and Late Holocene
American populations remains, as yet, unconfirmed
(Steele and Powell, 1992; Lahr, 1995; Jantz and Owsley,
2001, 2003; Van Vark et al., 2003). However, should it be
that Native Americans are derived from prehistoric
Northeast Asians with more generalized craniofacial features (Howells, 1959; Lahr, 1995, 1996), the dental morphological shift of Ainu to the Northeast Asians and Native Americans throws light on the relationships between
prehistoric Northeast Asia and Hokkaido.
Relethford (1994, 2002) emphasized that the patterning
of modern human variation yields important insights into
the origin and evolution of modern humans. However, the
technique employed to estimate such variation (interregional diversity (Fst), intraregional variance, and interregional relationships based on R matrix method) requires
neutrality with respect to natural selection (Relethford
and Blangero, 1990; Relethford, 1994, 2002; Relethford
and Harpending, 1994). It was asserted that tooth size has
been exposed to strong natural selective force (Brace et al.,
1991). It is true that overall tooth size plays a major role
in odontometric variation, but when the effects of overall
tooth size are eliminated with standardized data, several
independent vectors with eigenvalues greater than 1.0
Fig. 1. First PC scores based on raw measurement data, representing overall tooth size.
294
T. HANIHARA AND H. ISHIDA
TABLE 3. Eigenvalues and eigenvectors by PCA based on C-score data
Tooth
UI1-MD
UI2-MD
UC-MD
UP1-MD
UP2-MD
UM1-MD
UM2-MD
UM3-MD
LI1-MD
LI2-MD
LC-MD
LP1-MD
LP2-MD
LM1-MD
LM2-MD
LM3-MD
UI1-BL
UI2-BL
UC-BL
UP1-BL
UP2-BL
UM1-BL
UM2-BL
UM3-BL
LI1-BL
LI2-BL
LC-BL
LP1-BL
LP2-BL
LM1-BL
LM2-BL
LM3-BL
Eigenvalues
Proportion
Cumulative proportion
PC-1
PC-2
PC-3
PC-4
PC-5
PC-6
PC-7
⫺0.22335
⫺0.18840
⫺0.08187
⫺0.16201
⫺0.16839
⫺0.20793
⫺0.06484
⫺0.11277
⫺0.25291
⫺0.22331
⫺0.10153
⫺0.16187
⫺0.17386
⫺0.21998
⫺0.12699
⫺0.11090
0.18270
0.17877
0.26706
0.16003
0.15295
0.13461
0.19935
0.06116
0.21111
0.23865
0.30723
0.21658
0.18150
0.03302
0.10675
0.01628
9.38943
0.29342
0.29342
0.10680
0.18218
0.18718
0.26421
0.16645
⫺0.13457
⫺0.14163
⫺0.18236
0.12184
0.16635
0.18296
0.19182
0.09315
⫺0.18262
⫺0.22224
⫺0.28359
0.08271
0.10280
0.14224
0.24739
0.19008
⫺0.13838
⫺0.12569
⫺0.19313
0.07382
0.06074
0.05414
0.09345
0.06025
⫺0.23382
⫺0.27117
⫺0.32702
5.44359
0.17011
0.46353
0.22973
0.17820
0.14032
⫺0.23799
⫺0.24958
0.04751
⫺0.14380
0.05841
0.25650
0.24531
0.13036
⫺0.23278
⫺0.25798
0.04635
⫺0.13627
0.01663
0.20935
0.11141
0.16187
⫺0.23196
⫺0.28895
0.03864
⫺0.12785
⫺0.01663
0.25434
0.22533
0.14797
⫺0.14752
⫺0.23137
0.04094
⫺0.10987
0.03937
3.29825
0.10307
0.56660
⫺0.00541
⫺0.03247
0.03843
⫺0.04328
⫺0.04970
0.26044
0.13207
⫺0.40860
⫺0.03578
⫺0.00497
0.08958
0.03189
0.05289
0.25125
0.20492
⫺0.37093
0.05462
⫺0.08483
0.01212
⫺0.05023
⫺0.10188
0.27499
0.12994
⫺0.39146
0.05205
0.00336
0.02426
⫺0.01182
⫺0.02688
0.30512
0.18694
⫺0.28955
2.32693
0.07272
0.63932
0.10623
0.06819
⫺0.00379
0.03523
0.03963
0.21346
0.15127
0.06736
0.04473
0.02993
⫺0.15519
⫺0.22565
⫺0.25353
⫺0.08639
⫺0.22937
⫺0.14442
0.03569
⫺0.00087
⫺0.01250
0.16821
0.21729
0.39554
0.37601
0.25307
⫺0.12802
⫺0.19316
⫺0.20117
⫺0.17695
⫺0.21049
⫺0.08005
⫺0.20762
⫺0.16442
1.83195
0.05725
0.69657
⫺0.06150
0.03326
0.02725
⫺0.04944
0.19367
⫺0.00186
0.48994
0.08997
⫺0.09189
⫺0.14679
⫺0.12158
⫺0.05344
0.13985
⫺0.13264
0.30339
⫺0.02909
0.15807
0.24452
0.13667
⫺0.25620
⫺0.08535
⫺0.22107
0.12569
⫺0.06073
0.08032
0.10494
0.08264
⫺0.22781
⫺0.16988
⫺0.37151
⫺0.03774
⫺0.19843
1.58110
0.04941
0.74598
⫺0.18813
⫺0.07503
0.52194
⫺0.00693
⫺0.04832
0.01058
0.02062
0.06151
⫺0.24680
⫺0.10201
0.48720
⫺0.02439
⫺0.06685
0.00070
⫺0.02905
0.09786
⫺0.20974
⫺0.07970
0.26124
⫺0.01678
⫺0.00503
0.06649
0.02953
⫺0.00269
⫺0.30195
⫺0.18596
0.25746
⫺0.14516
⫺0.11698
⫺0.03660
⫺0.02409
0.05264
1.05119
0.03285
0.77883
TABLE 6. Intraregional variance
TABLE 4. Number of individuals used in R-matrix method
Number of individuals
Geographical regions
Male
Female
Far East
Australia
South/West Asia
Europe
Sub-Saharan Africa
Pacific
New World
1,162
308
923
1,006
772
844
1,052
278
193
78
171
90
258
431
TABLE 5. Fst values reflecting relative degree of interregional
variation for each sex
Sex
Male
Female
No. of
regions
Minimum
Fst
S.E.
Estimated
Fst
S.E.
5
7
5
7
0.1275
0.1127
0.1328
0.1214
0.0013
0.0011
0.0036
0.0028
0.1958
0.1747
0.2033
0.1872
0.0012
0.0010
0.0032
0.0025
remain. The intention of these significant eigenvectors
confirms that the dataset used as the basis for R matrix
analyses provides information beyond overall size. Indeed,
as shown in the PCA results based on raw measurement
data, factors other than overall size account for more than
half (56.7%) of the total variance.
This study revealed that less than 20% of the overall
patterning of odontometric diversity occurs between geographic regions (Table 5). As such, the apportionment of
diversity estimated from metric dental data is in general
Variance observed
Regions
5 Old World regions
Far East
Australia
South/West Asia
Europe
Sub-Saharan Africa
7 Worldwide regions
Far East
New World
Australia
Pacific
South/West Asia
Europe
Sub-Saharan Africa
Male
Female
1.0318
1.0885
0.9828
0.8893
1.0972
1.0034
1.0216
1.0038
0.9976
1.0279
1.0499
0.9561
1.1080
0.9816
0.9999
0.9057
1.1151
1.0353
0.9453
1.0528
1.0520
1.0424
1.0248
1.0579
concordance with results obtained from genetic markers,
DNA polymorphism (6 –25%), and craniometrics (11–14%)
(Ryman et al., 1983; Barbujani et al., 1997; Jorde et al.,
2000; Relethford, 1994, 2002), all of which stand in direct
opposition to between-group allocation estimates obtained
from variation in skin color (ca. 90%) (Relethford, 2002).
Many genetic studies document a higher level of withinregional diversity in sub-Saharan Africa than in other
geographic regions (Cann et al., 1987; Stoneking and
Cann, 1989; Rogers and Jorde, 1995). Relethford and
Harpending (1994) found the same pattern of elevated
within-regional diversity of sub-Saharan Africans in their
worldwide craniometric study. The present results add
another system of biological variation, odontometry,
Fig. 2. Scattergrams of first and second PC scores (a) and first and third PC scores (b), based on C-score data.
296
T. HANIHARA AND H. ISHIDA
Fig. 3. Scattergrams based on eigenvectors of R matrix under assumption of equal effective population sizes (a) and of scaled R
matrix using relative effective population weights of sub-Saharan Africa ⫽ 3, and Far East ⫽ Australasia ⫽ South/West Asia ⫽
Europe ⫽ 1 (b).
which indicates that the largest amount of intraregional
diversity occurs among the inhabitants of sub-Saharan
Africa.
Examinations of the unscaled and scaled R matrix results reveal fundamentally different patterns by sex.
Among males, Australians are identified as the most distinctive regional group. Among females, this distinctiveness occurs for sub-Saharan Africans rather than Austra-
lians. Although potential problems remain with regard to
estimation of hereditability of metric dental traits, sample
selection, methodological concerns, and so forth, the patterns of variation presented here are in essential agreement with those obtained from genetic and craniometric
analyses. While some investigators suggested that the low
within- and between-group variance, relatively high interobserver error, and strong method dependency render
DENTAL VARIATION OF MAJOR HUMAN POPULATIONS
odontometric variation of little value for distinguishing
between recent human populations on a microevolutionary level (Lasker and Lee, 1957; Harris and Bailit, 1988;
Kieser, 1990; Falk and Corruccini, 1982; Schunutenhaus
and Rösing, 1998), the results of the present study suggests that odontometric variation is efficacious for assessment of patterning among modern human populations on
a larger, worldwide scale.
ACKNOWLEDGMENTS
For their kind permission to study the materials under
their care, we express our sincere thanks to T. Molleson, R.
Kruszynski, L.T. Humphrey, and C. Stringer of the Natural
History Museum, London; R. Foley, M.M. Lahr, and M.
Bellatti of the Department of Biological Anthropology, University of Cambridge; A. Langaney and M.A. Pereira da
Silva of the Laboratoire d’Anthropologie Biologique, Musée
de l’Homme, Paris; D. Hunt, D. Owsley, S. Ousley, R. Potts,
M. London, and D.H. Ubelaker of the Department of Anthropology, National Museum of Natural History, Smithsonian
Institution, Washington, DC; I. Tattersall, K. Mowbray, and
G. Sawyer of the Department of Anthropology, American
Museum of Natural History, New York; J. Spechit, P. Gordon, L. Bonshek, and N. Goodsell of the Department of
Anthropology, Australian Museum, Sydney; J. Stone and D.
Donlon of the Department of Anatomy and Histology, University of Sydney; D. Henley of the New South Wales Aboriginal Land Council, Sydney; M. Chow, a dentist in Sydney; M. Hanihara of the School of Languages, Macquarie
University, Sydney; C. Pardoe and G.L. Pretty of the
Department of Anthropology, South Australian Museum, Adelaide; Y.H. Sinoto of the Department of Anthropology, the Bernice P. Bishop Museum, Honolulu;
M. Pietrusewsky of the Department of Anthropology,
University of Hawaii at Manoa, Honolulu; and T. Akazawa of the University Museum, University of Tokyo.
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