Negritos Australian Aborigines and the Уproto-sundadontФ dental pattern The basic populations in East Asia V.код для вставкиСкачать
AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 88:183-196 (1992) Negritos, Australian Aborigines, and the “Proto-Sundadont” Dental Pattern: The Basic Populations in East Asia, V TSUNEHIKO HANIHARA Department of Anatomy, Sapporo Medical College, South I , West 17, Chuo-ku, Sapporo, 060, Japan KEY WORDS Tooth crown feature, Sundadont, Southeast Asia ABSTRACT Five evolutionarily significant dental traits were identified from a B-square distance analysis of nine crown characters recorded for several populations of East Asia and Oceania. Intergroup variation in these traits distinguishes three major divisions of the Mongoloid dental complex: sundadonty, sinodonty, and the dental pattern of Australian Aborigines. The Australian crown features may be characterized a s having high frequencies of evolutionarily conservative characters. Negritos, one of the probable representatives of indigenous inhabitants of Southeast Asia who may have shared a common ancestor with Australians, possess the more derived sundadont dental pattern. As far as the five crown traits treated here are concerned, Australian dental features may be described as conforming to a “proto-sundadont” dental pattern, applying Turner’s terminology. This pattern may represent a microevolutionary step prior to the emergence of the sundadont and sinodont patterns. o 1992 Wiley-Liss, Inc. The peoples of East Asia are distinguished by the so-called Mongoloid dental complex (K. Hanihara, 1968), which is characterized by high frequencies of shovel-shaped upper first incisors, the sixth cusp, the seventh cusp, the deflecting wrinkle, and the protostylid on lower first molars. Turner (1976, 1979, 1983, 1987, 1989, 1990) distinguished two types or subdivisions of this complex: sundadonty and sinodonty. The sinodont pattern differs from the sundadont in frequencies of secondary traits (e.g., number of cusps on the molars, number of roots, various tiny ridges and grooves in the enamel, and other small anatomical features). According to Turner, eight key crown and root traits distinguish sundadonty and sinodonty: shoveling and double-shoveling of the upper first incisors; root number of upper third premolars; enamel extension of upper first molars; peg, reduced, or congenital absence of upper third molars; deflecting wrinkle and root number of lower first molars; and cusp number of lower second molars. Addition and intensification of these traits is characteristic of sinodonty, whereas 0 1992 WILEY-LISS,INC reduction and simplification characterize sundadonty (Turner, 1985). Turner (1979, 1985, 1987, 1989, 1990) has long argued that the sundadont pattern was present among late Pleistocene peoples of Southeast Asia, and th a t the more specialized sinodont pattern that characterizes today’s Northeast Asians and all indigenous Americans arose from sundadont antecedents at least 12,000-20,000 years B.P. The antecedents of the sundadont pattern are, however, less clear. Negritos, one possible representative of the aboriginal population of Southeast Asia, have a sundadont dental pattern (T. Hanihara, 1989c, 1990a,b,c, 1991a,b, 1992). The dental pattern characteristic of modern Aus- Received October 30,1990; accepted January 1, 1992. Address reprint requests to Tsunehiko Hanihara, Department of Anatomy, Sapporo Medical College, South 1,West 17, Chuoku, Sapporo, 060, Japan. T.HANIHARA 184 TABLE 1. Materials used Population Japanese Ainu #I Ainu #2 Nansei Islands Jomonese Collection of Provenience University of Tokyo, Jichi Medical School University of Tokyo, Sapporo Medical College University of Tokyo, Sapporo Medical College University of Tokyo, Kyoto University, etc. Yayoi #I University of Tokyo, National Science Museum, Tokyo Kyushu University Yayoi #2 Kyushu University Hirota Kyushu University Chinese Negrito University of Tokyo, Kyoto University University of Tokyo Early Thailand University of Hawaii Micronesian Polvnesian Meianesian Australian Bishop Museum BishoD Museum Bishop Museum, University of Tokyo University of Tokyo, Kagoshima University tralian Aborigines resembles sundadonty in seven out of eight key traits (the exception is the low frequency of four-cusped lower second molars) (Turner, 1990). It is now widely held that the modern Australians and the indigenous inhabitants of Southeast Asia, or Negritos, diverged in the early late Pleistocene from an ancestral “Proto-Australoid” population inhabiting Sundaland (Bellwood, 1978, 1985; Omoto, 1984). If so, then a comparison between the dental features of Negritos and Australian Aborigines might shed light on the morphology of that Proto-Australoid ancestry and the process of its divergence and expansion into Australia and eastern Asia. This study undertakes such a comparison. MATERIALS In addition to the samples previously described by the present author (T. Hanihara, 1989a,b,c, 1990a,b,c, 1991a,b, 19921, this study includes data for some groups of Australian Aborigines reported by various authors (K. Hanihara, 1976; Richards and Tefler, 1979; Smith et al., 1981; Townsend et Recent main-island Japanese Central and eastern Hokkaido (recent) Southwest Hokkaido (recent) Recent Nansei Islanders: Tanegashima, Amami-Oshima, Kikai, Okinoerabu, Yoro, Yoron, Okinawa, Miyako, Ishigaki, Hateruma, and Yonaguni Islands Honshu, Japan (12,000-2,300 years B.P.) Doigahama site, Yamaguchi Prefecture (2,300-1,700 years B.P.) Kanenokuma site, Fukuoka Prefecture Mitsu site, Saga Prefecture (2,300-1,700 years B.P.) Hirota site, Tanegashima, Kagoshima Prefecture (2,300-1,700 years B.P.) Manchuria (19th century) Aeta tribe, Bataan Peninsula, Luzon, The Philippines (recent) Early metal age of Thailand (ca. 3,000-6,000 years B.P.) Ban Chiang site, Nong Han district of Udon Thani province in Northeast Thailand Guam. Manana (me-contact) Mokapu site, Oahu, Hawaii (pre-contact) Fiji, New Guinea, Bismarck Arch. (recent) Mainly from Western Australia (recent) al., 1990; Turner, 1990). A list of the samples recorded by the present author is given in Table 1.The number of Melanesian individuals (mainly from Fiji) is relatively small, so that the sampling reliability is low. However, some examinations reported by Dahlberg (19491, Bailit et al. (19681, Barksdale (1972), Doran (19771, and Turner and Swindler (1978) provide results similar to those presented here. Accordingly, my data on Melanesians are included in the comparisons. Initially, nine permanent tooth crown traits were recorded for each individual. The exact sample sizes for each trait are given in Table 2. For the upper first incisor, the presence of shoveling was recorded. If the lingual fossa was less than 0.5 mm depth, the tooth was classified as no shovel (-); if between 0.5 mm and 1.0 mm, as moderate (+); and if deeper than 1.0 mm, as strong (+ +). For the upper first and second molars, the presence or absence of Carabelli’s cusp and the hypocone was scored, respectively. For the lower first molar, the existence of sixth cusp, seventh cusp, deflecting wrinkle, dis- "PROTO-SUNDADONT"DENTAL PATTERN 185 tal trigonid crest, and protostylid were recorded. For the lower second molar, cusp number and development of hypoconulid were scored. Data were obtained from the right-side teeth. When a right tooth was missing or badly damaged, the corresponding left tooth was investigated. The criteria for classification of non-metric crown traits are outlined elsewhere (T. Hanihara, 1990b, 1991a, 1992). In this study, male and female samples were combined, since almost no sexual dimorphism in frequencies was found in most of the groups studied. Modern Japanese have two primary origins: aboriginal Jomonese and Yayoi people from Northeast Asia. The Jomonese may have arrived from Southeast Asia via the now-submerged East Asian continental shelf by 12,000 years ago. During the period from Yayoi age to the early historic age, or from 2,300 years B.P. to 1,300 years B.P., admixture between the Jomonese and the incoming Yayoi people from Northeast Asia blurred the distinction between the two populations. The impact of the post-Yayoi migrants was unexpectedly large, so that a majority of modern main-island Japanese carry a large amount of Northeast Asian characteristics (Turner, 1976,1979, 1987,1990; K. Hanihara, 1985, 1987, 1991). Some Aeneolithic Yayoi populations, as represented by the skeletal remains excavated from the Doigahama site in the westernmost part of Honshu and Kanenokuma and Mitsu-Nagata sites in the northern part of Kyushu, are now believed to be typical representatives of immigrants from Northeast Asia (K. Hanihara, 1985, 1987, 1991; Dodo and Ishida, 1990). On the other hand, the more geographically isolated Ainu and residents of the Nansei Islands (the island chain stretching from the south end of main-island Japan to the east of Taiwan for about 1,200 kilometers) are clearly descended from Jomonese (Turner, 1979, 1987, 1990; T. Hanihara, 1989a,b,c, 1990a,b,c, 1991a,b, 1992). On the basis of such findings, some Japanese groups with physical affinities with Jomonese, or the Jomon lineagesnamely, Ainu and the inhabitants of the Nansei Island chain-are distinguished from a majority of main-island Japanese. In the present study, prehistoric and recent 186 T. HANIHARA groups from the Nansei island chain are separately analyzed. The prehistoric material consists of skeletal remains excavated from the Hirota site in Tanegashima Island, the northern end of the island chain (ca. 2,3001,700 years B.P.); the recent material comprises a pool of small local collections containing individuals from almost all islands of the chain. The populations properly referred to as the Neolithic Jomon population, Aeneolithic Yayoi populations, residents of the Nansei Island chain, the skeletal remains excavated from Hirota site in Tanegashima, those from Ban Chiang site in north Thailand, etc., will be simply referred to as Jomonese, Yayoi, Nansei Islands, Hirota, early Thailand, etc., in the tables and figures. METHODS Balakrishnan and Sanghvi (1968) proposed a B-squared distance coefficient. The distance between two populations P , and P , is defined as tained according to the formula for the elements: 9 9 i= 1 i= 1 This method also offers an advantage in providing contribution rates of each trait to the distance obtained (Constandse-Westermann, 1972). Although no formulae for this procedure could be found in the paper of Balakrishnan and Sanghvi (19681,the contribution of the difference in each trait between is defined by every pair of populations (Bmnj) the formula Si si RESULTS B-squared distance coefficients applied to the original nine discrete dental traits recorded in 14 samples are given in Table 3. In the scattergram resulting from multidimen*: S, sional scaling of the B-squared values (Fig. B,L = 11, three major clusters are evident. The j = l k=l 1=1 first includes modern Japanese, Chinese from the northern part of China (Manchuriwhere ans), and the immigrant Yayoi populations, corresponding to the sinodont dental group. The second is comprised of sundadont Jomonese and their lineages including samP,k is the frequency of the kth class, k = 1,2, ples from the Hirota site, Negritos, early 3, . , . , sj + 1, of the j t h character Sj, J = 1, Thailand people, Polynesians, and Microne2 , . , . , r, in the ith population Pi, z = 1, sians. Australian Aborigines are separated 2, . . . , q. The detailed procedure for comput- from the former two groups and comprise a ing this coefficient is as follows: the matrix third, separate entity. [A,,], being the per-trait dispersion matriFigure 2 shows the contribution of each ces for each population separately, has the trait to the B-squared distance coefficients elements between the Australian sample and the others calculated from the last formula in the previous section. It is easily recognized that Ay.k. l = p .y k. (1- p .y.k )In.. y7 k = 1 = -pUk . pyllnu, k f 1 k, 1 = 1,2, . . . . , the contributions of the differences in the Sj+1, frequencies of shoveling, the hypocone, the sixth cusp, the deflecting wrinkle, and the and n y is the sample size for Sj from Pi. The four-cusped lower second molars are relasame dispersion matrices for the samples tively high. The five characters are almost can be symbolized by n,[Aijkll,from which perfectly consistent with the key traits disthe pooled dispersion matrices [ c j k , ] , per tinguishing sinodonty from sundadonty trait, for all the samples combined, are ob- (Turner, 1987,1989,1990). c “PROTO-SUNDADONTDENTAL PAlTERN 1 c I$ c 3r- I $ ? oc 187 Examinations of Figures 3-7 show the frequencies of five crown traits for each sample (modified from Turner, 1990). The left side of these figures is based on my own data. The right side gives the frequencies of the traits for Australian Aborigines reported by other workers (K. Hanihara, 1976; Richards and Tefler, 1979; Smith et al., 1981; Townsend et al., 1990; Turner, 1990). Although data from several mainland and island Southeast Asians are needed to assess the population history within East Asia and the Pacific, the frequencies of the five traits nevertheless accord fully with an underlying dichotomy between sinodonty and sundadonty (Turner, 1987,1989,1990). The pattern of frequencies of five traits of Philippine Negritos from Luzon can be characterized as one of overall crown simplification. These populations fall within the range of sundadonty. On the other hand, these data suggest a rather unique position of Australian Aborigines. The Australian dental pattern includes low frequencies of marked incisor shoveling, more examples of the sixth cusp and the deflecting wrinkle expression on mandibular first molar, and high frequencies of the hypocone and the hypoconulid on maxillary and mandibular second molars, respectively. DISCUSSION Phylogenetic significance of crown traits Population variation of shovel-shaped incisors was first studied by Hrdlicka (1911, 1920). Concerning the phylogenetic significance of shoveling, HrdliEka (1920),Weidenreich (19371, Robinson (19561, Remane (19601, and others regarded this character as being primitive, since it was observable not only in modern human populations but also in Australopithecus, Homo erectus pekinensis (Sinanthropus),Homo sapiens neanderthalensis, and Hylobates. On the other hand, Adloff (1938) insisted that an incisor with a well-developed lingual tubercle, as seen in Sinanthropus, Homo sapiens neanderthalensis, and other fossil hominids, was not shovel-shaped but rather “tubercleshaped.” He argued that shoveling was a special character derived at a late evolutionary stage. Gorjanovic-Kramberger (1906) re- T.HANIHARA 188 2 Australian Polynesian ~ A Early ThailandA HirotaA AindIA 0 i i ~ rYwoi#2~ ~ ~ 1 Jamnesei Negritor L Nansei Islands AJapanese A Y w oiX 1 9 0 AChinese Fig. 1. Two-dimensional expression of multidimensional scaling applied to B-square distances based on nine discrete crown traits, expressing 85.7% of the total variance. ported that the Krapina incisors definitely exhibited shoveling and well-developed lingual tubercles. According to Adloff (1927), the basal lingual tubercle had been generated from the cingulum and was a feature common to mammals, thus a primitive character. McCown and Keith (1939) investigated the Neanderthaloid teeth from Tabun and pointed out that the well-developed lingual tubercle seen in the maxillary incisors was a feature distinguishing Homo sapiens neanderthalensis from modern human populations. In his study of American Indians and American Whites, Dahlberg (1949) provided the following discussion (p. 148)of the evolution of incisor shoveling: “The ancestral pattern for the incisors is a large tooth with prominent lateral borders on the lingual surface. Sinanthropus and Neanderthal man have these characters, but specialization has taken place in degree and in minor characteristics associated with the cingulum in modern man. Some populations, notably Whites, have proceeded in sirnplification to the point of having high frequency of no shovel-producing borders at all, whereas in the Indian the borders have specialized to even greater degrees of prominence than is seen in the specimens of early man.” Hominid maxillary molars are primitively four-cusped, but the hypocone tends to be reduced or lost in recent human populations as part of a general trend toward reduction and simplification of the molar dentition. In the lower molars, this trend is expressed in the reduction or loss of the hypoconulid on M, and M,, converting the primitive Y-5 pattern inherited from Dryopithecus into a symmetrical 4-cusped arrangement (“+4” pattern). The sixth cusp or “tuberculum accessorium” (Selenka, 1898) is seen on the lower molars of many fossil and extant hominoid (Dryopithecus, Australopithecus, “Sinanthropus,” Hylobates, and Neanderthals), and is generally thought to be a primitive character in modern human populations (Gregory, 1916, 1922; Gregory and Hellman, 1926;Hellman, 1928;Weidenreich, 1937; Jorgensen, 1956; Dahlberg, 1949; K. Hanihara, 1956, 1957; Robinson, 1956; Remane, 1960; Frisch, 1965; M. Suzuki and T. ~ ~ Hirota ..~ Nansei Islands Early Thailand Negrito Micronesian Polynesian Aina Japanese Chinese Ywoi#l Y w o i#2 Jixmnese Ain&l Aina I aM1) , I. 0 CI aM1) 1. 0 Japanese Chinese Y w o i#l 1.0 Protosty1id CMD 1.0 I. 0 4-cusP aim I. 0 1.0 Distal Trigonid Crest QM1) Fig. 2. Contribution of each trait to the B-square distances between Australians and other populations. Jamnese Yay0i#Z AinHirota Nansei Islands Early Thailand Negrito Mi cronesian Polynesian Hypocone US9 2. 0 Hirota Nansei Islands Early Thai land Negrito Mi cronesiau Polynesian 1. 0 1.0 Deflecting Wrinkle QMD Hirota Nansei Islands Early Thailand Negrito Micronesian Polynesian Ain&E Aindl Japanese Chinese Ywo i#l Y w o i#2 Jixmnese A i m 1 Carabelli Wl) Shovel UID Japanese Chinese Ywoi#l Y w o i#Z Jixmnese AindFl Japanese Chinese Y w o i#l Yay0i#2 Jananese Ainu#] AinHirota Nansei Islands Early Thai land Negrito Micronesian Polynesian T.HANIHARA 190 30 1 1 - P A i n u S l Negrito 20 Jamnese Micronesian Polynesian Ai n&2 Australian Early Thai land Hirota hklanesian in Shovelling W11) 50 ( X ) Broadbeach*): - 1 4 0 Pan+ustral ia* P a n A s t r a l ia* Ai nuU2 Hirota Anson B w Jamnese Swanport= %%% AinuSl Negrito Austral iao Micronesian Early Thai land 20 i i Polynesian Nansei-Islands Japanese YayoiXt Chinese Yayoi#2 f Yuenchmu*rr Yuenrhmu*#* ~a I mim rum Fig. 3. Percentages of shovel-shaped upper first incisors *Turner, 1990; **Smith et al., 1981; ***Richards and Tefler, 1979; ***“K. Hanihara, 1976; *****Townsendet al., 1990. Shading indicates the sinodont populations; the others are the sundadont populations. (Modified from Turner, 1990). Sakai [unpublished]). The deflecting wrinkle, a variant of the metaconid median ridge, is also regarded as primitive, for similar reasons (Weidenreich, 1937, 1945; K. Hanihara, 1956; M. Suzuki and T. Sakai unpublished; Swindler and Ward, 1988). Australians are generally conservative in all these features, and also retain a high frequency of lingual tubercles on I’ (Smith et al., 1981). Population history in Southeast Asia and Australia Human occupation in Australia is now believed to date back at least 40,000-50,000 years (Bowler, 1976; Howells, 1976; Thorne, 1976; Birdsell, 1977; Brace and Hinton, 1981; Groube et al., 1986). Because of the substantial morphological variation in Australians, diverse schemes of their origin(s) have been advanced, including monophyletic (Abbie, 19681, diphyletic (Thorne, 1976), and triphyletic models (Birdsell, 1967).Although early Australian fossils and recent Australians present a morphological continuum ranging from gracile (e.g., Keilor and Lake Mungo) to quite robust crania (e.g., Kow Swamp and Broadbeach), the gracile and robust groups are more similar to each other than they are to other anatomically modern Homo sapiens crania (Macintosh, 1963; Larnach, 1974; Macintosh and “PROTO-SUNDADONT DENTAL PATTERN 191 Hypocone Kh€2 I00 (X)+ Japanese Chinese Yayoi#;! Australian Me 1 anes i an Polynesian Hirota Negrito Early-Thailand Ywoitll Nansei-I s I ands Broadbeach= Micronesian Ai nuff2 Jamnese 70 I Ainu#I 1 60 5 50 1 Fig. 4. Frequency distribution of hypocone on upper second molar (modified from Turner, 1990). Symbols as in Fig. 3. 6o 50 (%) 5 0 1 - YwortR Polynesian Japanese Yayortl Jawnese Melanesian Australian Micronesian Chinese - - Hi rota Nansei Islands 20 Ainuff2 Early-Thailand Ainu#l Negrito 2o _] Fig. 5. Percentages of sixth cusp on lower first molars (modified from Turner, 1990). Symbols as in Fig. 3. Larnach, 1976; Pietrusewsky, 1984, 1990; both gracile and robust types (Jacob, 1976), Habgood, 1986; Brown, 1987). Smith et al. represent a t least part of the morphological (1981) concluded from the available dental background of the Australians. More immedata that metric and non-metric dental vari- diate antecedents may be represented by the ation within the various groups of Austra- Wajak, Niah, and Tabon individuals lians is associated with differences in envi- (Thorne, 1976). ronmental conditions, rather than gene The geographic source of Australian Abflow. Thorne (1976) suggested that the origines, or the original homeland of “ProtoHomo erectus remains from Java, including Australoids” from which the early Austra- 192 T.HANIHARA Deflecting wrinkle a 0 50 50 (%) YaYOI%2 Chinese Micronesian Ywi#l Melanesian Early-Thailand Australian Polynesian 4 0 Japanese Jamnese Nansei-Islands AinuSI AinuffL Negrito 10 Hi rota 0 Fig. 6. Percentages of deflecting wrinkle on lower first molars (modified from Turner, 1990). Symbols as in Fig. 3. 4 cusp pattern W Negrito Early-Thailand Melanesian Nansei-Islands Jamnese 30 Japanese Micronesian 20 Yayoill2 Australian Chinese 10 Kalduru*rr Pan++ustral la* Broadbeach*l: Fig. 7. Frequency distributions of four-cusped pattern on lower second molars (modified from Turner, 1990) Symbols as in Fig 3 lians may have diverged, can be traced to late Pleistocene mainland Southeast Asia or the landmass of the continental shelf called Sundaland. In the tropical rain forest, open inland areas, and the shores of Sundaland, a Proto-Australoid population may have evolved into modern “Proto-Malays” beginning some 20,00040,000 years ago (Omoto, 1984; T. Hanihara, 1990a,b, 1991a, 1992).A number of investigators have emphasized the degree of Australoid inheritance in the physical characteristics of the Proto-Ma- lays-for example, Negritos (Birdsell, 1949, 1977; Garn, 1961; Coon, 1962; Howells, 1976; Jacob, 1976; Brues, 1977; Kennedy, 1979; Glinka, 1981; Bellwood, 1985). Genetic and metric dental studies suggest, however, that the Philippine Negritos do not show close similarity to Australians (Omoto, 1984; T. Hanihara, 1989c, 1990a,b, 1991a, 1992).The sundadont dental pattern of Negritos documented here provides more direct biological evidence favoring a local-evolution hypothesis for the origin of the reduced “PROTO-SUNDADONTDENTAL PATTERN Years B P. Japan Asia 0 10.000 20.000 1 Negr i to \t Jmnese Sinodont Minatogawa tr t 40. 0 0 0 50.000 60. 000 70. 0[10 0 90.000 100.000 Sahul land tr Proto-fvhlw Sundadont 30. 0 0 0 80.000 Sunda land b 193 t Australoid t r Proto-Sundadont Dental Pattern Fig, 8. A hypothetical schema showing modification of dental morphology and racial diversification in East Asia and West Oceania during late Pleistocene times (modified from Omoto, 1984). size and simple crown morphology that characterize sundadonty. On morphological and historical grounds it is quite likely that in Australian dental groups the formation of a Negrito-like sundadont dental pattern may have accompanied a phenotypic specialization to small size and gracilization under the environmental condition of late Pleistocene Sundaland over a period of some 20,000 years. Turner (1990) pointed out the strong similarity between the dentitions of Southeast Asians and Australians. From Turner’s review of frequency variation of eight key crown and root traits, the Australian dentition is closely associated with the sundadont dental pattern (Turner, 1990). The five characters treated here are minor traits which distinguish Australians from Southeast Asian sundadont populations. Australian Aborigines are distinctive in exhibiting high frequencies of weak shoveling (I1), hypocone (M2), sixth cusp and deflecting wrinand hypoconulid (MJ, traits that kle (MI), also appear frequently in individuals of the Upper Pleistocene or earlier ages. It is probable, therefore, that high frequencies of these traits characterized a microevolutionary phase prior to the emergence of the Southeast Asian sundadont pattern. If so, it may be tentatively termed the “proto-sundadont” dental pattern, applying Turner’s terminology. Figure 8 displays a hypothetical reconstruction of microevolution and modification of dental characters in East Asian and western Oceanic populations (modified from Omoto, 1984). This hypothesis is in full agreement with that proposed by Turner (1987,1989,1990). The frequencies of the five traits discussed above (Figs. 3-7) link Australians to the sinodont groups. However, it is still far from obvious whether the sinodont dental pattern was derived from a “proto-sundadont” pattern or from the sundadont pattern. As Turner (1987) points out, we cannot rule out genetic drift or founder effect i n explaining the origin of sinodonty from sundadonty, particularly in the system of small, relatively isolated populations that must have prevailed in the late Pleistocene of Northeast Asia. A great many more data on early occupants of Southeast and Northeast Asia are needed to elucidate the relationship among sinodonty, sundadonty, and protosundadonty. ACKNOWLEDGMENTS I am deeply indebted to Professor C.G. Turner I1 of the Department of Anthropology, Arizona State University, for his great encouragement, invaluable support, and critical advice at Kyoto in September, 1990. I wish to express my sincere gratitude to Professor G. Ito of the Department of Orth- 194 T. HANIHARA odontics, Kagoshima University Dental School; Professor M. Pietrusewsky of the Department of Anthropology, University of Hawaii; Professor Y.H. Sinoto of the Department of Anthropology, Bernice P. Bishop Museum in Honolulu; Professor K. Omoto and Professor B. Endo of the Department of Anthropology, Faculty of Science, the University of Tokyo; Professor T. Yohro of the Department of Anatomy, Faculty of Medicine, the University of Tokyo; Professor Y. Dodo and Professor H. Ishida of the Department of Anatomy, Sapporo Medical College; Professor H. Ishida and Professor K. Katayama of the Department of Zoology, Faculty of Science, Kyoto University; and Dr. T. Nakahashi and Dr. N. Doi of the Department of Anatomy, Faculty of Medicine, Kyushu University, for permission to examine materials in their care. This study was supported in part by a grant-in-aid for overseas scientific surveys, “Anthropological Studies on the Origin of the Pacific Populations,” and “The Population Genetic Survey of Negritos” from the Ministry of Education, Science, and Culture in Japan. The principal investigators are Professor K. Hanihara of the International Research Center for Japanese Studies and Professor K. Omoto, respectively. This research was also financially supported by grants-in-aid for scientific research 01740483, 02740412, 02225213, 03209210, and 03740424 from the Japanese Ministry of Education, Science, and Culture. LITERATURE CITED Abbie AA (1968) The homogeneity of Australian Aborigines. Archaeol. Phys. Anthropol. Oceania 3t223-231. Adloff P (1927) Das Gebiss des Menschen und der Anthropoiden und das Abstammungsproblem. Z. Morph. 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