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Page 1: Anthropol. Sci. 101(1), 25-46, 1993 Craniofacial Features

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

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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

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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

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28 T. HANIHARA

Table 2. Basic statistics for 24 craniofacial measurements in each sample.

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Craniofacial Features of Southeast Asians 29

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30 T. HANIHARA

Table 2 (cont'd)

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Craniofacial Features of Southeast Asians 31

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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.

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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-

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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

Page 11: Anthropol. Sci. 101(1), 25-46, 1993 Craniofacial Features

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.

Page 12: Anthropol. Sci. 101(1), 25-46, 1993 Craniofacial Features

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.

Page 13: Anthropol. Sci. 101(1), 25-46, 1993 Craniofacial Features

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.

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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.

Page 15: Anthropol. Sci. 101(1), 25-46, 1993 Craniofacial Features

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

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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

Page 17: Anthropol. Sci. 101(1), 25-46, 1993 Craniofacial Features

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.

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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.

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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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