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,

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 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 , 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: , 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 -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. , 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 physical features (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 Asian mainland 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 , 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 prehistoric Southeast 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 CraniofacialFeatures of SoutheastAsians 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 correlation coefficients, biplot graphic display, group average cluster analysis, and multidimen sionalscaling. On the basis of the recent findings for the population history of the Japanese, the samples 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 analyses Table 3 shows three (length-breadth, length-height, and breadth-height) indices of neurocranial part. The Australian and Melanesian samples are character izedby the dolicho- to hyperdolichocranic and orthocranic skull shape. Regarding the breadth-height index, none of the Asian and Pacific samples are tapeinocranic, but the Australian sample is distinctly acrocranic. Another important character is the facial morphology (Table 4). Regarding Kallman's upper facial indices, the two Chinese samples show higher indices than any other samples. The orbital, nasal, and upper facial indices indicate that a geographical cline exists from Australia to northern China through Southeast Asia and the Pacific regions. The East Asian samples are leptene or mesene, mesoconch, and mesorrhine or leptorrhine. On the other hand, the Australian and Southeast Asian samples together with the Jomonese sample are characterized by mesene-euryene, mesoconch-chamaeconch, and chamaerrhine. In the Pacific samples, Melanesians show close similarity to Austral ians.Almost all the samples are orthognathic; only the Australian sample falls within the prognathic range. Although Melanesian, early Thailand, and Okinawa Island samples fall within the range of mesognathic, the first lies just within the upper limit and the latter two near the lower limit.

Multivariate analyses In 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 the inter-group relationship and variance and correlation of variables on the same dimensions. The variance is represented by the length of variable vectors, and correlation 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, and Jomonese 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

Fig. 2. Biplot graphic display based on seven cranial and facial indices.

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, and Taiwanese, 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 upper faces 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-island Japanese. All the Nansei Island samples together with the Taiwanese sample are included in the Jomonese cluster, which links to the cluster containing two Southeast Asian samples. The three Pacific samples, Guamanians and two Hawaiians, loosely attach to the Southeast Asian - Jomonese subdivision. The Australian and Melanesian 36 T. HANIHARA

Fig. 3. Cluster analysis applied to Euclidean distance between every pair of samples based on seven craniofacial indices. 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 Craniofacial Features of Southeast Asians 37

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.

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. 38 T. HANIHARA

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.

DISCUSSION Reevaluation of Southeast Asian population history The 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 evidence supporting the broadly accepted theory of the migration of East Asians into Southeast Asia having caused the biological gracilization of the region can be summarized as follows (reviewed by Turner, 1987); 1) People with morphological affinities with present Australians and Melanesians originally occupied Southeast Asia (Callenfels, 1936; Mijsberg, 1940; Jacob, 1967; Howells, 1973a; Bellwood, 1978, 1985, 1987; Kennedy, 1979); and 2) North China was the cultural hearth of CraniofacialFeatures of SoutheastAsians 39 eastern Asia: The content and wealth of the Chinese civilization, e.g. the Shang Dynasty at Anyang, casts a long shadow over all of eastern Asia when contrasted with the simple Hoabinhian stone tools of Southeast Asia (Barth, 1952; Van Heekeren and Knuth, 1976; Jacob, 1967; Li, 1977; Bellwood, 1985). Howells (1973a, 1976, 1977) further identified a separate region he referred to as "Old Melanesia," corresponding to island Southeast Asia, New Guinea, and Australia, as the ancestral homeland of the western Pacific peoples some 40,000-50,000 years ago. However, the available human fossil records do not necessarily support the hypothesis of a "Australoid" occupation in Southeast Asia in the late Pleistocene and early Holocene (Bulbeck,1982). In the Southeast Asian region, three important human 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 recent Tasmanians and Australians. Kennedy (1979) suggested, however, that the skull was actually towards the gracile end of the Australoid range of variation. Although the Wajak remains were identified as Australoid by Dubois (1921), Jacob (1975) found in the upper face not only Australomelanesian but also Southeast Asian features. Glinka (1981) further suggests that there may be relations not only to the Australomelanesians but also to the "Proto-Malays" who represent the oldest Indonesian populations with a certain Southeast Asian character. The human remains from Tabon consisted of fragments of at least 5 individuals; the mandible is considered close to the Australian range by Macintosh (1978). Glinka (1981) reviewed the Southeast Asian human remains discovered in 41 sites from the period between 12,000-4,000 years B.P.According to him, these remains have racial elements not only of Australomelanesians but also of Southeast Asians and Negritos (Glinka, 1981). The dental traits of the specimens from the Gua Cha rock-shelter on the Nenggiri River, Kelantan, Malay Peninsula, divided chronologically into a Hoabinhian (Mesolithic) and a later (Neolithic) series, are more similar to those of Southeast Asians than to those of Australians (Trevor and Brothwell, 1962). More recently, Cuong (1996) described two nearly complete skulls from Vietnam dated to the early Hoabinhian Culture, or approximately 10,000 years ago. He found that the two skulls exhibited morphological features which were intermediate between modern 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 hypothesis arguing 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 Australians and Melanesians. This may be a sign of evolutionary divergence in craniofacial shape among populations of different geographic areas. The uniqueness of Southeast Asian craniofacial morphology presented herein is an additional support of the local evolution hypothesis of present Southeast Asian physical features.

Affinities of Jomonese The 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 Southeast Asia with lesser admixture with East Asian invaders. However, the assessment of the possibility mentioned above will require further comparative study including both recent and prehistoric Southeast Asian and Australomelanesian cranial series. In a previous study, I pointed out the possibility that the early Southeast Asians without intensive admixture with Chinese can be regarded as members having recent Southeast Asian cranial features using the cranial data published by Pietrusewsky (1981, 1984, 1988) (Hanihara, T., 1992d). In the present study, the biological relationships among prehistoric and recent East and Southeast Asians, Australians, Melanesians, and Jomonese based on the 13 cranial measurements were re-analyzed using Q-mode correlation coefficients. The variables used are: maximum cranial length (M-1); nasio-occipital length (M-1d); maximum cranial breadth (M-8); minimum frontal breadth (M-9); maximum frontal breadth (M-10); biauricular breadth (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. 5 represents the dendrogram obtained by group average clustering, and Fig. 6 shows a two dimensional scattergram obtained by multidimensional scaling. In Fig. 6, 81.7% of the total variance is accounted for. Although the variables used are restricted within the neurocranial part, these findings suggest that certain modern traits of Southeast Asian cranial features such as brachycrany and gracilization may developed inside of Southeast Asia, not from East Asia. The two figures reveal that the Jomonese cranial features are all significantly different from Australomelanesians and East Asian Chinese. Jomonese exhibit strong similarities to the post-Pleistocene early Southeast Asians, or the "generalized Asian populations"; they share cranial traits indicative of a regional source in Southeast Asia. These results suggest the hypothesis that there was population continuity from at least late Pleistocene times to the Christian period in Southeast Craniofacial Features of Southeast Asians 41

Fig. 5. Cluster analysis applied to Q-mode correlation coefficients based on 13 cranial measurements.

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 of human mitochondrial DNA (mtDNA) from Asian and Pacific populations, including Jomonese specimens dated to about 3,000-6,000 years B.P.,and found a close affiliation of Jomonese with present Southeast Asians from Malaysia and Indonesia. Taking all of these together, the present findings support the previous dental suggestion that the sundadont ancestors of Jomonese likely arrived from Sundaland via the now-submerged East Asian continental shelf during and after the late Pleistocene. One of the main routes for the peopling of the Japanese Archipelago might 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 the Pacific populations (Hawaiians and Guamanians) and Jomonese, as emphasized by 42 T. HANIHARA

Fig. 6. Two dimensional graph of MDS applied to the same matrix used in Fig. 5, expressing 81.7% of total variance.

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 the distinctions 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. 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.

REFERENCES Ballinger, S.W., Schurr, T.G., Torroni, A., Gan, Y.Y., Hodge, J.A., Hassan, K., Chen, K.H., and Wallce, D.C. (1992) Southeast Asian mitochondrial DNA analysis reveals genetic continuity of ancient migrations. Genetics 130, 139-152. Barth, F. (1952) The southern Mongoloid migration. Man 52, 5-8. Bellwood, P. (1978) Man's Conquest of the Pacific: The Prehistory of Southeast Asia and Oceania, Oxford Univ. Press, New York. Bellwood, P. (1985) Prehistory of the Indo-Malaysian Archipelago, Academic Press, Sydney. Bellwood, P. (1987) The prehistory of island Southeast Asia; A multidisciplinary review of recent research. J. World Prehist. 1, 171-224. Birdsell, J.B. (1977) The recalibration of a paradigm for the first peopling of greater Australia. In Sunda and Sahul: Prehistoric Studies in Southeast Asia, Melanesia, and Australia (Allen, J., Golson, J., and Jones, R., eds.), Academic Press, London, pp. 113-167. Brace, C.L. (1992) Maximum human biological diversity in the Mesolithic, and the peopling of the Americas. Am. J. Phys. Anthropol. (Suppl.) 14, 52. Brace, C.L., Brace, M.L., and Leonard, W.R. (1989) Reflections on the face of Japan: A multivariate craniofacial and odontometric perspective. Am. J. Phys. Anthropol. 78, 93-113. Brace, C.L., Brace; M.L., Dodo, Y., Hunt, K.D., Leonard, W.R., Li, Y., Sangvichien, S., Xiang-Oing, S., and Zhenbiao, Z. (1990) Micronesians, Asians, Thais and relations: A craniofacial and odontometric perspective. Micronesica (Suppl.) 2, 323-348. Brace, C.L., and Hunt, K.D. (1990) A nonracial craniofacial perspective on human variation: A(ustralia) to Z(uni). Am. J. Phys. Anthropol. 82, 341-360. Brothwell, D.R. (1960) Upper Pleistocene human skull from Niah Caves, Sarawak. Sarawak Mus. J. 9, 323-349. 44 T. HANIHARA

Brues, A.M. (1977) People and Race, The Macmillan Series in Physical Anthropology, Macmillan Publishing Co., New York. Bulbeck, D. (1982) A re-evaluation of possible evolutionary processes in Southeast Asia since the late Pleistocene. Bulletin of the Indo-Pacific Prehistory Association 3, 1-21. Cheboksarov, N.N. (1966) The Ethnic Anthropology of Eastern Asia, Nauka, Moscow. Callenfels, P.V. van S. (1936) The Melanesoid civilizations of eastern Asia. Raffles Mus. Bull. 1, 41- 51. Cavalli-Sforza, L.L., Piazza, A., Menozzi, P., and Mountain, J. (1988) Reconstruction of human evolution: Bringing together genetic archaeological and linguistic data. Proc. Natl. Acad. Sci. USA 85, 6002-6006. Coon, C.S. (1962) The Origin of Races, Alfred A. Knopf, New York. Cuong, N.L. (1986) Two early Hoabinhian crania from Thanh Hoa Province, Vietnam. Z. Morph. Anthrop. 77, 11-17. Dubois, E. (1921) Der proto-australische Mensch von Wajak (Java). Verhdl. K. Akad. Wetensch. Amsterdam 23, 10-13. Gabriel, K.R. (1971) The biplot graphic display of matrices with application to principal component analysis. Biometrika 58, 453-467. Glinka, J. (1981) Racial history of Indonesia. In Rassengeschichte der Menschheit (Schwidetzky, I., ed.) R. Lodenbourg, Munchen/Wien, pp. 97-113. Hanihara, K. (1985) Geographic variation of modern Japanese crania and its relationship to the origin of Japanese. Homo 36, 1-10. Hanihara, K. (1987) Estimation of the number of early migrants to Japan: A simulative study, J. Anthrop. Soc. Nippon 95, 391-403. Hanihara, K. (1991) Dual structure model for the population history of Japanese. Japan Review 2, 1- 33. Hanihara, T. (1989) Comparative studies of geographically isolated populations in Japan based on dental measurements. J. Anthrop. Soc. Nippon 97, 95-107. Hanihara, T. (1990a) Affinities of the Philippine Negritos with Japanese and the Pacific populations based on dental measurements: The basic populations in East Asia, I. J. Anthrop. Soc. Nippon 98, 13-27. Hanihara, T. (1990b) Dental anthropological evidence of affinities among the Oceania and the pan - Pacific populations: The basic populations in East Asia, II. J. Anthrop. Soc. Nippon 98, 233-246. Hanihara, T. (1990c) Studies on the affinities of Sakhalin Ainu based on dental characters: The basic populations in East Asia, III. J. Anthrop. Soc. Nippon 98, 425-437. Hanihara, T. (1991a) The origin and microevolution of Ainu as viewed from : The basic populations in East Asia, VIII. J. Anthrop. Soc. Nippon 99, 345-361. Hanihara, T. (1991b) Dentition of Nansei islanders and peopling of the Japanese Archipelago: The basic populations in East Asia, IX. J. Anthrop. Soc. Nippon 99, 399-409. Hanihara, T. (1992a) Dental and cranial evidence on affinities of the East Asian and Pacific populations. In Japan in the World IV: Japanese as a Member of the Asian and Pacific Populations, International Research SymposiumNo. 4 (Hanihara, K., ed.), IRCJS Press, Kyoto, pp. 119-136. Hanihara, T. (1992b) Dental and cranial affinities among the populations in East Asia and the Pacific: The basic populations in East Asia, IV. Am. J. Phys. Anthropol. 88, 163-182. Hanihara, T. (1992c) Negritos, Australian Aborigines and the "proto-sundadont" dental pattern: The basic populations in East Asia, V. Am. J. Phys. Anthropol. 88, 183-196. Hanihara., T. (1992d) Biological relationships among Southeast Asians, Jomonese, and the Pacific Craniofacial Features of Southeast Asians 45

populations as viewed from dental characters: The basic populations in East Asia, X.J. Anthrop. Soc. Nippon 100, 53-67. Horai, S., Hayasaka, K., Murayama, K., Wate, N., Koike, H., and Nakai, N. (1989) DNA amplification from ancient human skeletal remains and their sequence analysis. Proc. Jap. Acad. 65 (Ser. B), 229-233. Horai, S., Kondo, R., Murayama, K., Hayashi, S., Koike, H., and Nakai, N. (1991) Phylogenetic affiliation of ancient and contemporary humans inferred from mitochondrial DNA, Phil. Trans. R. Soc. Lond. B 333, 409-417. Howells, W.W. (1973a) The Pacific Islanders, Scribners, New York. Howells, W.W. (1973b) 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 Vol. 67, Harvard Univ. Press, Cambridge. Howells, W.W. (1976) Physical variation and history in Melanesia and Australia. Am. J. Phys. Anthropol: 45, 641-650. Howells, W.W. (1977) The sources of human variation in Melanesia and Australia. In Sunda and Sahul: Prehistoric Studies in Southeast Asia, Melanesia, and Australia (Allen, J., Golson, J., and Jones, R., eds.), Academic press, London, pp. 169-186. Jacob, T. (1967) Racial identification of Bronze age human from Bali, Indonesia. J. Dent. Res. 46, 930-910. Jacob, T. (1975) Morphology and palaeoecology of early man in Java. In Paleoanthropology: Morphology and Paleoecology (Tuttle, R.H., ed.), The Hague, Paris, pp. 311-324. Katayama, K. (1990) A scenario on prehistoric Mongoloid dispersals into the south Pacific, with special reference to hypothetic proto-Oceanic connection. Man and Culture in Oceania 6, 151- 159. Kennedy, K.A.R. (1979) The deep skull of Niah: An assessment of twenty years of speculation concerning its evolutionary significance. Asian Perspectives 20, 32-50. Kozintsev, A. (1990) Ainu, Japanese, their ancestors and neighbours: Cranioscopic data. J. Anthrop. Soc. Nippon 98, 247-267. Li, C. (1977) Anyang, Univ. Washington Press, Seattle. Li, Y., Brace, C.L., Qiang, G., and Tracer, D.P. (1991) Dimensions of face in Asia in the perspective of geography and prehistory. Am. J. Phys. Anthropol. 85, 269-279. Macintosh, N.W.G. (1978) The Tabon cave mandible. Archaeol. Phys. Anthropol. Oceania 13, 143- 159. Martin, R., and Saller, K. (1957) Lehrbuch der Anthropologic, Dritte Aufl, Bl., I, Gustav Fischer Verg, Stuttgart. Mijsberg, W.A. (1940) On a Neolithic Palaeo-Melanesian lower jaw found in a kitchen midden at Gua Kepah. Province Wellesley. Straits Settlements. Proc. III Cong. Prehist. Far East, pp. 100-118. Omoto, K. (1984) The Negritos: Genetic origins and microevolution. Acta Anthropogenetica 8, 137- 147. Omoto, K. (1986) Racial formation in East Asia and the Pacific. In The Origin of Japanese (Hanihara, K., ed.), Shogakukan, Tokyo, pp. 139-160. (In Japanese) Omoto. K. (1987) Discovery of Man: Molecular Anthropology, Yomiuri Kagaku Sensho 14, Tokyo, pp. 169-204. (In Japanese) Omoto, K. (1992) Mongoloid populations in north and south: The origins and variation of . Science J. Kagaku 62, 217-226. (In Japanese) Pietrusewsky, M. (1981) Cranial variation in early metal age Thailand and Southeast Asia studied by 46 T. HANIHARA

multivariate procedures. Homo 32, 1-26. Pietrusewsky, M. (1984) Metric and Non-metric Cranial Variaion in Australian Aboriginal Populations Compared with Populations from the Pacific and Asia, Occasional Papers in Human Biology 3, Australian Institute of Aboriginal Studies, Canberra. Pietrusewsky, M. (1988) Multivariate comparisons of recently excavated Neolithic human crania from the Socialist Republic of Vietnam. Int. J. Anthropol. 3, 267-283. Trevor, J.C., and Brothwell, D.R. (1962) The human remains of Mesolithic and Neolithic date from Gua Cha, Kelantan. Federation Museums J. 7, 6-22. Turner, C.G., II (1976) Dental evidence on the origins of the Ainu and Japanese. Science 193, 911- 913. Turner, C.G., II (1979) Dental anthropological indications of agriculture among the Jomon of central Japan. Am. J. Phys. Anthropol. 51, 619-636. Turner, C.G., II (1983) Sinodonty and sundadonty; A dental anthropological view of Mongoloid microevolution, origin and dispersal into the Pacific Basin, Siberia, and the Americas. In Late Pleistocene and Early Holocene Cultural Connections of Asia and America (Vasilievsky, R.S. ed.), USSR Academy of Science, Siberian Branch, pp. 72-76. Turner, C.G., II (1985) The dental search for native American origins. In Out of Asia; Peopling the Americas and the Pacific (Kirk, R., and Szathmary, E., eds.), Australian National University Printery, Canberra, pp. 31-78. Turner, C.G., II(1987) Late Pleistocene and Holocene population history of East Asia based on dental variation. Am. J. Phys. Anthropol. 73, 305-321. Turner, C.G., II (1989) Teeth and prehistory in Asia. Sci. Am. 263, 70-77. Turner, C.G., 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. Turner, C.G., II, and Lien, C.M. (1984) Diachronic differences in Taiwan dental morphology. Bull. Indo-Pacific Prehist. Assoc. 5, 74-82. Van Heekeren, H.R., and Knuth, E. (1967) Archaeological Excavations in Thailand, Vol. 1, Sai-Yok, Munksgaard, Copenhagen. Von Koenigswald, G.H.R. (1952) Evidence of a prehistoric Austral-melanesoid population in Malaya and Indonesia. Southwest J. Anthropol. 8, 92-96.