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Anthropol. Sci. 101(1), 25-46, 1993
Craniofacial Features of Southeast Asians and Jomonese:
A Reconsideration of Their
Microevolution Since the Late Pleistocene
TSUNEHIKO HANIHARA
Department of Anatomy, Sapporo Medical College,
S-1, W-17, Chuo-ku, Sapporo 060, Japan
Received June 8, 1992
•ôGH•ô Abstract•ôGS•ô Univariate and multivariate statistical procedures were applied to 24
measurements recorded in 944 crania from East and Southeast Asia and Oceania
to assess the possible origins and affinities of these populations with special
reference to the origin of Jomonese. The results show that Jomonese are much
more like the prehistoric mainland Southeast Asians and recent aboriginal people
in Borneo, the Dajaks, than like East Asians, Polynesians (Hawaiians), western
Micronesians (Guamanians), Melanesians, and Australians. The orthodox view of
Southeast Asian prehistory has held that the region was occupied by
Australomelanesians before the southern expansion of Chinese between 2,000 and
4,000 years ago. The contradictory findings presented here form the basis of an
alternative view that Southeast Asian craniofacial features resulted from local
evolution, not admixture. The present findings favor the prehistoric Southeast
Asians, with lesser admixture with East Asian invaders from the north, as the most
likely source for not only the present-day Southeast Asians, but also prehistoric
Jomonese, and the Pacific populations.
•ôGH•ô Key Words•ôGS•ô: Sundaland, Australians, Local evolution hypothesis, Generalized
Asian populations
INTRODUCTION
Population movements from the major initial centers may have served to absorb
and replace the non-agricultural peoples in broadly contiguous areas, for example,
in the China-Southeast Asian region ca. 2,000-4,000 years B.P., China-Japan
region ca. 2,300-1,300 years B.P., and the Tigris-Euphrates region ca. 7,000-
8,000 years B.P. (Bellwood, 1978, 1985, 1987; Glinka, 1981; Hanihara, K., 1985,
1987, 1991; Brace et al., 1989, 1990; Brace and Hunt, 1990; Brace, 1992).
According to Brace (1992), increased inter-regional contact in the Bronze and Iron
ages and subsequent periods accelerated the process of gene flow between what had
previously been relatively isolated regional populations.
The ongoing and accelerating reduction in regional human biological diversity
in Southeast Asia is said to have taken place by a series of migrations from the north
between 2,000 and 4,000 years ago (Coon, 1962; Jacob, 1967; Birdsell,1977; Brues,
1977; Kennedy, 1979; Bellwood, 1978, 1985). Although human skeletal materials
26 T. HANIHARA
are too sparse to give us a clear picture of regional diversity during the late
Pleistocene and early Holocene times in Southeast Asia, the archaeological findings
and somatological characteristics of the indigenous inhabitants of Southeast Asia,
e.g. Negritos, argue for the once wide distribution of people having physical
affinities with present Australians and Papuans (Von Koenigswald, 1952; Coon,1962; Jacob, 1967; Howells, 1976; Brues, 1977; Kennedy, 1979; Bellwood, 1978,
1985; Glinka, 1981).
On the other hand, the operation of similar selective forces, such as caused by
tropical rain-forests, on the widely dispersed but contiguous human populations for
a long period of time (20,000-30,000 years) may have been a more important factor
than the extent of gene flow on the formation of the Southeast Asian physicalfeatures (Cheboksarov, 1966; Bulbeck, 1982; Omoto, 1984, 1992; Turner, 1987,
1990; Pietrusewsky, 1988; Hanihara, T., 1992a, b, c, d).
Turner (1976, 1979, 1989, 1990) proposed that sundadonty evolved in Southeast
Asia among earlier late Pleistocene peoples, and that it subsequently evolved into
the more specialized sinodont pattern of present Chinese and Northeast Asians. The
keystone of his hypothesis has been the dental characteristics of prehistoric Jomon
populations in Japan, the Jomonese, who were considerably isolated from the Asianmainland for about 12,000 years (Turner, 1987, 1990). Considering the marked
morphological and biological differences between Jomonese and all late Pleistocene
and Holocene humans from north China and Siberia, sundadont Jomonese could
only have originated in Southeast Asia or now-flooded Sundaland (Turner, 1990).
According to Turner (1983, 1985, 1989, 1990), the sundadont dental pattern had
to have existed at least 12,000 years ago, long before the so-called southern
expansion of East Asian Chinese into Southeast Asia.If prehistoric Southeast Asians with lesser admixture with East Asian migrants,
or Chinese, have morphological features like those of Australomelanesians, then
Jomonese must have had characteristics similar to those of Australians or Melanesians.
However, I have seen no references to anyone finding such close affinities. On the
other hand, if the physical features of Southeast Asians are local evolutionary
products under the environmental conditions and sufficient time depth, say 20,000
years or so, then sundadont ancestor of Jomonese can be traced back to prehistoricSoutheast Asians.
In my previous studies, I have presented two lines of evidence that favor the local
evolution hypothesis for Southeast Asians; 1) The dental characters of the Philippine
Negritos, who share somatological similarities with Australians and are therefore
regarded as "Australoid" in Southeast Asia, fall within the range of sundadonty. On
the other hand, Australian dental features are characterized by high frequencies of
evolutionarily conservative characteristics, called "proto-sundadont" patter. 2) The
prehistoric Southeast Asians and recent Negritos show closer affinities to present
Craniofacial Features of Southeast Asians 27
Southeast Asians and Jomonese in their dental features than to Australians and
Melanesians (Hanihara, T., 1992a, b, c, d). Based on such findings, the purpose of
this paper is to present a detailed comparison of East and Southeast Asian and
Oceanian craniofacial variation and to investigate the possible origin and affinities
of prehistoric Jomonese.
MATERIAL AND METHODS
The materials of this study were prehistoric and near-recent adult male samples
from East and Southeast Asia and Oceania (Table 1). Twenty-four craniofacial
measurements used for the analyses were given in Table 2. These measurements
were taken according to Martin and Saller (1957).
Table 1. Sample names and provenience
28 T. HANIHARA
Table 2. Basic statistics for 24 craniofacial measurements in each sample.
Craniofacial Features of Southeast Asians 29
30 T. HANIHARA
Table 2 (cont'd)
Craniofacial Features of Southeast Asians 31
32 T. HANIHARA
The sample size of the Australians is small, so the sampling reliability is low.
However, the broad examination reported by Pietrusewsky (1984) provides the basic
statistics similar to those presented here. The Australian sample used here falls
within a cluster of nine Australian samples in comparison with 38 Asian and the
Pacific samples based on the 16 of the author's 24 craniofacial measurements that
are in common with those reported by Pietrusewsky (1984) (Fig. 1). Accordingly,
the Australian data are included in the comparisons.
Fig. 1. Group average cluster analysis based on Q-mode correlation coefficients between every pair
of samples reported by Pietrusewsky (1984) and presented in this study (Australians).
Craniofacial measurements used are: M-1, M-5, M-8, M-9, M-11, M-17, M-29, M-30, M-40,
M-46, M-48, M-51a, M-52, M-54, M-55, and M-61. The same symbols represent the same
population groups.
Craniofacial Features of Southeast Asians 33
The statistical procedures used are Euclidean distance, Q-mode correlationcoefficients, biplot graphic display, group average cluster analysis, and multidimensionalscaling.
On the basis of the recent findings for the population history of the Japanese, thesamples from the Nansei Island chain are referred to as Jomonese lineages in the
present study (Hanihara, K., 1985, 1987, 1991; Kozintsev, 1990; Hanihara, T., 1989,1990a, b, 1991a, b, 1992a, b, c, d).
RESULTS
Univariate analysesTable 3 shows three (length-breadth, length-height, and breadth-height)
indices of neurocranial part. The Australian and Melanesian samples are characterizedby the dolicho- to hyperdolichocranic and orthocranic skull shape. Regardingthe breadth-height index, none of the Asian and Pacific samples are tapeinocranic,but the Australian sample is distinctly acrocranic. Another important character isthe facial morphology (Table 4). Regarding Kallman's upper facial indices, the twoChinese samples show higher indices than any other samples. The orbital, nasal, andupper facial indices indicate that a geographical cline exists from Australia tonorthern China through Southeast Asia and the Pacific regions. The East Asiansamples are leptene or mesene, mesoconch, and mesorrhine or leptorrhine. On theother hand, the Australian and Southeast Asian samples together with the Jomonesesample are characterized by mesene-euryene, mesoconch-chamaeconch, andchamaerrhine. In the Pacific samples, Melanesians show close similarity to Australians.Almost all the samples are orthognathic; only the Australian sample falls withinthe prognathic range. Although Melanesian, early Thailand, and Okinawa Islandsamples fall within the range of mesognathic, the first lies just within the upper limitand the latter two near the lower limit.
Multivariate analysesIn the first step, the method of biplot graphic display developed by Gabriel (1971)
was applied to the seven indices described above. This method can display both theinter-group relationship and variance and correlation of variables on the samedimensions. The variance is represented by the length of variable vectors, andcorrelation between variables is given by the cosine between variable vectors
(Fig. 2).Four major clusters are evident in Fig. 2. The Australian and Melanesian samples
form a relatively tight cluster and are characterized by being prognathic, acrocranic,and chamaerrhine. The samples from the Nansei Island chain, Southeast Asia, andJomonese form a second major cluster. They show more similarities with Australian-
34 T. HANIHARA
Table 3. Three indices of neurocranial part (%)
*Mesocrany , **Dolichocrany: +Hypsicrany, ++Orthocrany: -Metriocrany, --Acrocrany
Table 4. Four indices of facial part (%)
*Euryene , **Mesene, ***Leptene: +Chamaeconch, ++Mesoconch: -Leptorrhine, --Mesorrhine,---Chamaerrhine: /Orthognath , //Mesognath, ///Prognath
Craniofacial Features of Southeast Asians 35
Melanesian samples than with East Asian samples. However, the features of the
samples of the second cluster are not so prominent as those of the Australian and
Melanesian samples. The East Asian samples, two Chinese, Japanese, Korean, andTaiwanese, form a third cluster bordering on the Hawaiian - Guamanian grouping,
the fourth cluster. These two are characterized by being mesene - leptene,
mesoconch, hypsicranic, and mesocranic. The East Asian samples have higher upperfaces and higher orbits, and the Pacific samples have particularly round and high
cranial vaults.
In the next step, cluster analysis was applied to the Euclidean distance based on
the seven craniofacial indices (Fig. 3). In the first main cluster, which contains East
and Southeast Asian and the Pacific samples, there are two main groups. The first
contains four East Asian samples, namely Chinese #1 & #2, Korean, and main-islandJapanese. All the Nansei Island samples together with the Taiwanese sample are
included in the Jomonese cluster, which links to the cluster containing two SoutheastAsian samples. The three Pacific samples, Guamanians and two Hawaiians, loosely
attach to the Southeast Asian - Jomonese subdivision. The Australian and Melanesian
Fig. 2. Biplot graphic display based on seven cranial and facial indices.
36 T. HANIHARA
samples are well differentiated from the other samples.
Multidimensional scaling was applied to Q-mode correlation coefficients based
on the 24 craniofacial measurements taken from the 16 populations. Fig. 4a is a plot
of the samples for the first two dimensions, expressing 71.4% of the total variance.
A scattergram of the first and third dimensions resulting from the same method is
shown in Fig. 4b; this accounts for 68.6% of the total variance. These two
representations express 90.0% of the information contained in the Q-mode cone
lationmatrix.
These two graphs indicate a clear regional separation into East Asian, Polynesian,
western Micronesian, Australomelanesian, and Southeast Asian groups. The posi
tionof the Australian and Melanesian samples reaffirms the distinctiveness of the
two population complexes. The Jomonese and Nansei Island samples are included
in the Southeast Asian constellation. The Pacific samples (Guamanians and
Fig. 3. Cluster analysis applied to Euclidean distance between every pair of samples based on seven
craniofacial indices.
Craniofacial Features of Southeast Asians 37
Hawaiians) are plotted at a relatively isolated position. This group forms the closest
association with the East Asian group. With the possible exception of Taiwanese,
the clustering of the East and Southeast Asian samples corresponds to the
association presented in Figs. 2 and 3. The unstable position of the Taiwanese
sample may be due to the strong mainland Chinese influence on the aboriginal
Taiwanese, who otherwise had physical affinities with Southeast Asians (Bellwood,
1985, 1987; Turner and Lien, 1984; Turner, 1987).
The marked separation between recent main-island Japanese and the Jomon-
Nansei Island group agrees with the dual structure model for the population history
of Japanese proposed by K. Hanihara (1991), which assumes the separate primary
origins of the modern Japanese.
Fig. 4a. Two dimensional expression of multidimensional scaling (MDS) applied to Q-mode corre
lationcoefficients between every pair of samples based on 24 craniofacial measurements.
Using first two axes, 71.4% of total variance is expressed.
38 T. HANIHARA
DISCUSSION
Reevaluation of Southeast Asian population historyThe earlier interpretation of the physical and cultural variability exhibited by the
recent inhabitants of Southeast Asia has stressed the importance of historic or even
prehistoric invasions of Chinese from the north (Barth, 1952; Von Koenigswald,1952; Coon, 1962; Bruce, 1977; Bellwood, 1978, 1985, 1987). The evidencesupporting the broadly accepted theory of the migration of East Asians intoSoutheast Asia having caused the biological gracilization of the region can besummarized as follows (reviewed by Turner, 1987); 1) People with morphologicalaffinities with present Australians and Melanesians originally occupied SoutheastAsia (Callenfels, 1936; Mijsberg, 1940; Jacob, 1967; Howells, 1973a; Bellwood,1978, 1985, 1987; Kennedy, 1979); and 2) North China was the cultural hearth of
Fig. 4b. Two dimensional expression of MDS using first and third axes in the same analysis in Fig.
4a, accounting for 68.6% of total variance.
Craniofacial Features of Southeast Asians 39
eastern Asia: The content and wealth of the Chinese civilization, e.g. the ShangDynasty at Anyang, casts a long shadow over all of eastern Asia when contrastedwith the simple Hoabinhian stone tools of Southeast Asia (Barth, 1952; VanHeekeren and Knuth, 1976; Jacob, 1967; Li, 1977; Bellwood, 1985). Howells
(1973a, 1976, 1977) further identified a separate region he referred to as "OldMelanesia," corresponding to island Southeast Asia, New Guinea, and Australia, asthe ancestral homeland of the western Pacific peoples some 40,000-50,000 yearsago.
However, the available human fossil records do not necessarily support thehypothesis of a "Australoid" occupation in Southeast Asia in the late Pleistoceneand early Holocene (Bulbeck,1982). In the Southeast Asian region, three importanthuman remains in the late Pleistocene have been discovered: Niah cave (Borneo,ca. 40,000 years B.P.); wajak (Java, ca. 25,000-40,000 years B.P.); and Tabon
(Palawan, ca. 22,000-24,000 years B.P.). The analysis by Brothwell (1960)addressed the closest morphological affinities of the Niah skull with recentTasmanians and Australians. Kennedy (1979) suggested, however, that the skull wasactually towards the gracile end of the Australoid range of variation. Although theWajak remains were identified as Australoid by Dubois (1921), Jacob (1975) foundin the upper face not only Australomelanesian but also Southeast Asian features.Glinka (1981) further suggests that there may be relations not only to theAustralomelanesians but also to the "Proto-Malays" who represent the oldestIndonesian populations with a certain Southeast Asian character. The humanremains from Tabon consisted of fragments of at least 5 individuals; the mandibleis considered close to the Australian range by Macintosh (1978). Glinka (1981)reviewed the Southeast Asian human remains discovered in 41 sites from the periodbetween 12,000-4,000 years B.P. According to him, these remains have racialelements not only of Australomelanesians but also of Southeast Asians and Negritos
(Glinka, 1981). The dental traits of the specimens from the Gua Cha rock-shelteron the Nenggiri River, Kelantan, Malay Peninsula, divided chronologically into aHoabinhian (Mesolithic) and a later (Neolithic) series, are more similar to those ofSoutheast Asians than to those of Australians (Trevor and Brothwell, 1962). Morerecently, Cuong (1996) described two nearly complete skulls from Vietnam datedto the early Hoabinhian Culture, or approximately 10,000 years ago. He found thatthe two skulls exhibited morphological features which were intermediate betweenmodern Australians and Southeast Asians.
The recent contradictory findings by Bulbeck (1982), Omoto (1984, 1986, 1987,1992), Turner (1987, 1990), Pietrusewsky (1988), and T. Hanihara (1989, 1990a,b, c, 1991a, b, 1992a, b, c, d), and to a lesser extent by Cavalli-Sforza et al. (1988)and Ballinger et al. (1992), have formed the basis of an alternative hypothesisarguing that local selection in Southeast Asia is responsible for the formation of the
40 T. HANIHARA
physical features of today's Southeast Asians.The present findings stress the considerable difference between East Asian and
Southeast Asian groups. At the same time, two Southeast Asians, the early Thailand
people and the aboriginal Borneans (Dajaks), are generally unrelated to Australiansand Melanesians. This may be a sign of evolutionary divergence in craniofacialshape among populations of different geographic areas. The uniqueness of SoutheastAsian craniofacial morphology presented herein is an additional support of the localevolution hypothesis of present Southeast Asian physical features.
Affinities of JomoneseThe results obtained in the present analyses support the dentally oriented idea that
the origin of Jomonese can be traced back to the indigenous inhabitants of SoutheastAsia with lesser admixture with East Asian invaders. However, the assessment ofthe possibility mentioned above will require further comparative study includingboth recent and prehistoric Southeast Asian and Australomelanesian cranial series.In a previous study, I pointed out the possibility that the early Southeast Asianswithout intensive admixture with Chinese can be regarded as members having recentSoutheast Asian cranial features using the cranial data published by Pietrusewsky
(1981, 1984, 1988) (Hanihara, T., 1992d). In the present study, the biologicalrelationships among prehistoric and recent East and Southeast Asians, Australians,Melanesians, and Jomonese based on the 13 cranial measurements were re-analyzedusing Q-mode correlation coefficients. The variables used are: maximum craniallength (M-1); nasio-occipital length (M-1d); maximum cranial breadth (M-8);minimum frontal breadth (M-9); maximum frontal breadth (M-10); biauricularbreadth (M-11b); biasterionic breadth (M-12); nasion-bregma chord (M-29); bregma-lambda chord (M-30); bifrontal breadth (M-43); interorbital breadth (M-49a);mastoid height (H-MDH); and mastoid width (H-MDW) (M: Martin's abbreviation,Martin and Saller, 1957; H: Howells' abbreviation, Howells, 1973b). Fig. 5represents the dendrogram obtained by group average clustering, and Fig. 6 showsa two dimensional scattergram obtained by multidimensional scaling. In Fig. 6,81.7% of the total variance is accounted for. Although the variables used arerestricted within the neurocranial part, these findings suggest that certain moderntraits of Southeast Asian cranial features such as brachycrany and gracilization maydeveloped inside of Southeast Asia, not from East Asia.
The two figures reveal that the Jomonese cranial features are all significantlydifferent from Australomelanesians and East Asian Chinese. Jomonese exhibitstrong similarities to the post-Pleistocene early Southeast Asians, or the "generalizedAsian populations"; they share cranial traits indicative of a regional source inSoutheast Asia. These results suggest the hypothesis that there was populationcontinuity from at least late Pleistocene times to the Christian period in Southeast
Craniofacial Features of Southeast Asians 41
Asia and the Japanese Archipelago. It is worth noting here that Horai et al. (1989,1991) performed nucleotide sequence analysis of the major non-coding region ofhuman mitochondrial DNA (mtDNA) from Asian and Pacific populations, includingJomonese specimens dated to about 3,000-6,000 years B.P., and found a closeaffiliation of Jomonese with present Southeast Asians from Malaysia and Indonesia.
Taking all of these together, the present findings support the previous dentalsuggestion that the sundadont ancestors of Jomonese likely arrived from Sundalandvia the now-submerged East Asian continental shelf during and after the latePleistocene. One of the main routes for the peopling of the Japanese Archipelagomight have been through the Nansei Island chain (Turner, 1979, 1987, 1989, 1990;Hanihara, T., 1991a, b, 1992b).
The results obtained do not specifically support a direct relationship between thePacific populations (Hawaiians and Guamanians) and Jomonese, as emphasized by
Fig. 5. Cluster analysis applied to Q-mode correlation coefficients based on 13 cranial measurements.
42 T. HANIHARA
Brace and his colleagues (Brace et al., 1989, 1990; Brace and Hunt 1990; Li et al.,
1991) and Katayama (1990). On the contrary, what does come as something of a
surprise is the nearly complete separation of the Hawaiian - Guamanian cluster from
the Jomonese - Southeast Asian cluster in Figs. 4a and 4b. Evidently the 3,000-
3,600 year time span since the presumed initial movement into the Pacific Rim and
genetic drift as well as the founder effect were sufficient for the production of thedistinctions observed in the cranial series of Micronesians and Polynesians. Both
diachronic comparison and analysis of geographical variation of Polynesians and
Micronesians will be required for evidence bearing on their population affinities
with Jomonese.
Fig. 6. Two dimensional graph of MDS applied to the same matrix used in Fig. 5, expressing 81.7%
of total variance.
Craniofacial Features of Southeast Asians 43
ACKNOWLEDGEMENTS
I wish to express my sincere gratitude to Professor M. Pietrusewsky of the
Department of Anthropology, University of Hawaii-Manoa; Professor K. Omoto
and Professor B. Endo of the Department of Anthropology, Faculty of Science, the
University of Tokyo; Professor H. Ishida and Professor K. Katayama of the
Department of Zoology, Faculty of Science, Kyoto University; Dr. T. Nakahashi,
Dr. N. Doi, and Professor Y. Shibata of the Department of Anatomy, Faculty of
Medicine, Kyushu University; and Professor Y.H. Sinoto of the Department of
Anthropology, B.P. Bishop Museum in Honolulu for their kind permission to study
the materials under their care.
This study was supported in part by the following grants from the Ministry of
Education, Science, and Culture in Japan: Grant-in-Aid for International Scientific
Research "Anthropological Studies on the Origin of the ;Pacific Populations"organized by Professor K. Hanihara of International Research Center for Japanese
Studies; Grants-in-Aid for Scientific Research Nos. 01740483, 02740412, 03740424,
02225213, 03209210, 04208210, and 04212118.
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