Metrical and Nonmetrical Analyses of Modern Female Crania in the Northwestern Kyushu Area TOMOHIDE WATANABE1, KAZUNOBU SAIKI1, KEISHI OKAMOTO1, TETSUAKI WAKEBE1*

ANTHROPOLOGICAL SCIENCE Vol. 112, 147–159, 2004

Metrical and nonmetrical analyses of modern female crania in the northwestern Kyushu area TOMOHIDE WATANABE1, KAZUNOBU SAIKI1, KEISHI OKAMOTO1, TETSUAKI WAKEBE1*

1Department of Developmental and Reconstructive Medicine, Course of Medical and Dental Sciences, Graduate School of Biomedical Sciences, Nagasaki University, 1-12-4 Sakamoto, Nagasaki, 852-8523 Japan

Received 29 September 2003; accepted 18 December 2003

Abstract In the northwestern Kyushu area, at the southwestern region of the mainland of Japan, a native people of Jomon lineage lived during the Aeneolithic Yayoi period, and differed morphologically from the immigrant people of continental lineage of the northern Kyushu area. In this study, the metric and nonmetric characteristics of 204 modern female crania from northwestern Kyushu were examined in order to investigate their morphological characteristics and lineage relationships by comparing them with several modern Japanese, Ryukyuan, Hokkaido Ainu, Korean, Chinese, and northern and northwestern Kyushu Yayoi and Jomon cranial series. The craniometric characteristics of the northwestern Kyushu Japanese females were found to be similar to those of the mainland Japanese, as well as to those of Koreans, Chinese, and the northern Kyushu Yayoi, but differed somewhat from the Ryukyuan and Hokkaido Ainu conditions, and distinctly diverged from the northwestern Kyushu Yayoi and Jomon conditions. The results of multivariate analyses of nonmetric traits coincided generally with those of the craniometrical analyses; however, the northwestern Kyushu Japanese was found to be relatively close to the Ryukyuans, and markedly remote from the Hokkaido Ainu. These findings suggest that the northwestern Kyushu Japanese possess morphological traits in common with other mainland Japanese populations due to genetic influences of the continental lineage during or after the protohistoric Kofun period.

Key words: Japanese, northwestern Kyushu, female, craniometry, nonmetric cranial trait

Introduction area had a markedly higher face and taller stature than the Jomon (Kanaseki, 1956, 1959, 1966; Nakahashi et al., 1985; Numerous morphological studies—both metric and non- Nakahashi and Nagai, 1989; Nakahashi, 1993). Therefore metric—of crania from the Japanese archipelago have been the Yayoi people of the northern Kyushu area are considered reported (e.g. Koganei, 1893; Hasebe, 1917; Miyamoto, to have been strongly influenced genetically by immigrants 1924; Morita, 1950; Yamaguchi, 1973, 1978; Dodo, 1974; from the Asian continent or their descendants. It is widely Mouri, 1976; K. Hanihara, 1984). In recent years, detailed considered—based on many morphological and genetic comparative studies of Ryukyuan crania have also been pub- studies—that people of continental lineage were influential lished by Dodo and others (Dodo et al., 1998, 2000; Dodo, in the formation of the modern Japanese (e.g. Mouri, 1986, 2001). Because morphological regionality is closely related 1988; Ossenberg, 1986; Dodo, 1987; Dodo and Ishida, 1988, to the lineage history of populations, clarifying the morpho- 1990, 1992; T. Hanihara, 1989, 1991; Kozintsev, 1990; logical characteristics of the modern peoples in each area of Yamaguchi, 1990; K. Hanihara, 1991, 1993; Nakahashi, Japan is useful in the study of the origin and formation of the 1993; Omoto, 1995; Omoto and Saito, 1997; Ikeda, 1998). Japanese. Noda (1993) morphologically investigated male crania to In this study we examined modern female crania from the ascertain whether modern males in the northwestern Kyushu northwestern Kyushu area, i.e. the northwest region of area had inherited the characteristics of Yayoi males, and Kyushu Island, including many islets (Figure 1). The people concluded—because of similarities such as a lower cranial of the Yayoi period (ca. 300 BC to 300 AD) in this area vault, a lower face, and a lower orbit—that the modern inherited some morphological characteristics from the peo- males of this area show influence of Yayoi males from the ple of the Jomon period (ca. 10,000 BC to 300 BC), who had same area. However, the morphological characteristics of a low, wide face and short stature (Naito, 1971, 1981, 1984). modern female crania from northwestern Kyushu have not However, the Yayoi people in neighboring northern Kyushu been clarified. Cranial morphological studies have mainly been carried out using male or mixed samples, but little * Corresponding author. e-mail: [email protected] detailed work has been performed on female samples. How- phone: 81-095-849-7022; fax: 81-095-849-7024 ever, there have been some cases of excavation of exclu- Published online 29 June 2004 sively female human skeletal remains from ancient sites. in J-STAGE (www.jstage.jst.go.jp) DOI: 10.1537/ase.00100 Also, lineage analyses using mitochondrial DNA (which is

Figure 1. Map showing the location of the northwestern Kyushu area and samples from other areas used in the present study.

matrilinearly inherited) in both ancient and modern Japanese Kyushu Yayoi, and Tsukumo Jomon samples were used. For have increased in recent years (Horai et al., 1989, 1991; direct comparisons of trait incidences in the nonmetrical Shinoda and Kunisada, 1990, 1994; Kurosaki et al., 1993; analyses, northwestern Kyushu, northern Kyushu, Kinki, Oota et al., 1995; Shinoda and Kanai, 1999). Therefore we and Kanto–Tohoku Japanese, Hokkaido Ainu, northwestern considered it necessary to clarify the lineage relations of not Kyushu Yayoi, northern Kyushu Yayoi, and Jomon samples only males but also females. were used. For multivariate analyses, all samples were used This study employed standard metrical and nonmetrical in both metrical and nonmetrical comparisons. methods for comparative analysis of modern and earlier female crania. We attempted to clarify metric and nonmetric Metrical analysis characteristics of the modern female crania from the north- The 41 cranial measurements and 16 indices shown in western Kyushu area and to estimate their lineage relation- Appendix 1 were measured according to Martin’s methods ships by analyzing and comparing modern Japanese, (Bräuer, 1988; Baba, 1991), and 17 measurements and 9 Ryukyuans, Ainu, Koreans, Chinese, Yayoi, and Jomon indices shown in Table 2 were used in the analyses. It has female crania, along with comparisons and consideration of been reported that the method of measuring upper facial previous findings based on male crania. height varies between researchers, and this affects the results of analysis (Dodo, 2001). In this study, using Martin’s meth- Materials and Methods ods, the distance between the ‘nasion’ and the ‘alveolare’ was measured. The upper facial height of the northwestern Materials Kyushu Yayoi (Naito, 1971) was corrected by adding The 204 female crania stored at the Nagasaki University 2.0 mm (Dodo, 2001) because it had been measured between Graduate School of Biomedical Sciences were examined. the ‘nasion’ and the ‘prosthion’. The upper facial indices of These were crania of people born between 1866 and 1937 in the northwestern Kyushu Yayoi were also corrected by a Nagasaki prefecture, and their birth and death dates and proportional amount. domiciles were documented. Comparative samples used in Student’s t-test was performed to test the significance of this study are shown in Table 1. For direct comparisons of differences in the measurements. For the indices, the nor- measurements and indices in metrical analyses, northwest- mality of distribution was examined in the large sample of ern Kyushu, northern Kyushu, Kinki, and Kanto Japanese, northwestern Kyushu Japanese. As a result, the null hypoth- Hokkaido Ainu, northwestern Kyushu Yayoi, northern esis of normal distribution could not be rejected at the 5% Vol. 112, 2004 MODERN FEMALE CRANIA IN NORTHWESTERN KYUSHU 149

Table 1. Female samples compared in this study Sample name Period1 No. of crania Source and brief information Metrical analyses Northwestern Kyushu Japanese Modern 202 Present study: Nagasaki prefecture Northern Kyushu Japanese Modern 57 Nakahashi and Nagai (1989): mainly Fukuoka prefecture Central Kyushu Japanese Modern 45 Matsumoto (1956): Kumamoto prefecture Kinki Japanese Modern 20 Miyamoto (1924): Kinki district Hokuriku Japanese Modern 20 Otsuki (1930): Hokuriku district Kanto Japanese Modern 84 Morita (1950): Kanto district Tohoku Japanese Modern 23 Yamasaki et al. (1967): Tohoku district Amami–Okinawa Islander Modern 34 Dodo et al. (2001): Amami and Okinawa islands (Ryukyuan) Miyako Islander (Ryukyuan) Modern 48 Ikeda (1974): Miyako island Hokkaido Ainu Modern 64 Koganei (1893), Yamaguchi (1978): Hokkaido Korean Modern 50 Shima (1934): Kyonggi-do, Korea Chinese Modern 17 Shima (1933): Fuschun, Liaoning Province, China Northern Kyushu Yayoi Aeneolithic Yayoi 86 Nakahashi and Nagai (1989): mainly from plain sites of Fukuoka prefecture Northwestern Kyushu Yayoi Aeneolithic Yayoi 15 Naito (1971): from seaside sites of Nagasaki prefecture Tsukumo Jomon Neolithic Jomon 28 Kiyono and Miyamoto (1926): from Tsukumo site of Okayama prefecture Nonmetrical analyses Northwestern Kyushu Japanese Modern 204 Present study: Nagasaki prefecture Northern Kyushu Japanese Modern 27 Dodo et al. (1992, unpublished data): mainly Fukuoka prefecture Kinki Japanese Modern 85 Mouri (1988): Kinki district Hokuriku Japanse Modern 70 Mouri (1988): Hokuriku district Kanto–Tohoku Japanese Modern 50 Dodo (1974), Dodo and Ishida (1987): Kanto and Tohoku districts Amami–Okinawa Islander Modern 39 Mouri (1988): Amami and Okinawa islands (Ryukyuan) Hokkaido Ainu Modern 81 Dodo (1974), Dodo and Ishida (1987): Hokkaido Korean Modern 26 Takenaka (1994): Kyongsangnam-do, Korea Chinese Modern 35 Wang and Sun (1988): Taiyuan, Shanxi Province, China Northern Kyushu Yayoi Aeneolithic Yayoi 67 Saiki et al. (2000): mainly from plane sites of Saga prefecture Northwestern Kyushu Yayoi Aeneolithic Yayoi 57 Saiki et al. (2000): from seaside sites of Nagasaki prefecture Jomon Neolithic Jomon 71 Dodo and Ishida (1990): from sites of Kanto and Tohoku districts 1 Modern: since the latter half of 19th century; Aeneolithic Yayoi: ca. 300 BC to 300 AD; Neolithic Jomon: ca. 2000 BC to 300 BC. level, except in the upper facial (K) and nasal indices. Stu- Analysis version 4.0 (Esumi Inc., 2002). dent’s t-test was used with the other indices. Furthermore, to compare visually the various measurements, Mollison’s Results deviation curves were prepared. For multivariate analyses, principal component analysis and cluster analysis using Cranial measurements Penrose’s shape distances were performed in an attempt to The 41 measurements and 16 indices for the modern assess biological lineage relations among the samples female crania of the northwestern Kyushu Japanese are (Constandse-Westermann, 1972). shown in Appendix 1.

Nonmetrical analysis Neurocranium The 22 nonmetric cranial traits shown in Appendix 2 were The neurocrania of the northwestern Kyushu female assessed according to Dodo’s criteria (Dodo, 1974, 1975, Japanese were shorter in maximum cranial length, narrower 1986). However, only 17 traits were used in the analyses in maximum cranial breadth, and higher in basion–bregma because biasterionic suture vestige, foramen of Vesalius, cli- height than those of the northern Kyushu Yayoi, northwest- noid bridging, juglar foramen bridging, and sagittal sinus ern Kyushu Yayoi, and Tsukumo Jomon. The measurements groove left were not investigated in the Kinki and Hokuriku of the four modern Japanese series were comparable overall, Japanese, and in the Amami–Okinawa islanders. The inci- except that the Kinki Japanese had somewhat shorter maxi- dence of each trait was analyzed by a 2 test or Fisher’s exact mum cranial length and wider maximum cranial breadth. probability test. For multivariate analyses, Smith’s mean Among these samples, the northwestern Kyushu Japanese measure of divergence (MMD) was calculated in order to was closest to the northern Kyushu Japanese. The cranial- assess biological distances among the samples (Sjøvold, type classifications based on three cranial indices, the 1973), and principal coordinate analysis was then per- length–breadth, length–height, and breadth–height indices, formed. The statistical tests and multivariate analyses were were mesocranic, hypsicranic, and acrocranic in both Japa- processed by a computer software package from Kyoritu- nese groups. shuppan Inc. (Tanaka et al., 1984) and Excel Multivariate 150 T. WATANABE ET AL. ANTHROPOLOGICAL SCIENCE

Table 2. Comparison of measurements and indices of the modern, Yayoi and Jomon female crania in Japan (in mm except for angular measurements) Northwestern Northern Kinki Kanto Hokkaido Northern Northwestern Tukumo Metrical characters Kyushu Kyushu Japanese Japanese Ainu Kyushu Kyushu Jomon with Martin’s Japanese Japanese Yayoi Yayoi number n Mean S.D. n Mean n Mean n Mean n Mean n Mean n Mean n Mean 1 Maximum cranial 201 172.0 5.79 57 172.8 20 169.3* 82 170.8 63 177.2** 86 177.0** 15 178.1** 18 175.7* length 8 Maximum cranial 201 134.6 5.15 57 134.0 20 137.7** 82 135.9 64 136.9** 84 138.4** 15 139.3** 21 141.9** breadth 17 Basion–bregma 199 131.8 4.08 57 131.3 20 132.5 81 132.5 55 133.3* 66 130.7 7 128.3* 14 126.1** height 8/1 Length–breadth 201 78.3 4.00 57 77.6 20 81.5** 82 79.7** 63 77.3 72 78.1 15 78.2 18 80.8** index 17/1 Length–height index 199 76.7 2.85 57 76.0 20 77.9 81 77.7* 54 75.1** 62 74.1** 7 71.2** 14 71.5** 17/8 Breadth–height 199 98.1 4.07 57 98.0 20 95.8* 81 97.7 55 97.2 56 94.9** 7 92.5** 14 89.3** index 5 Basion–nasion 200 95.4 3.70 57 95.5 20 95.0 84 95.6 61 99.9** 65 96.5* — 12 96.3 length 23 Horizontal 197 491.4 12.29 57 483.0** 20 493.0 81 493.7 63 502.8** 60 512.5** 14 502.1** 14 506.0** circumference 24 Transverse arc 198 304.3 10.42 57 301.4 20 307.0 81 313.2** 62 317.9** 81 305.0 11 309.8 19 298.4* 25 Total sagittal arc 198 359.0 11.89 57 359.3 20 359.0 82 357.6 54 360.7 45 364.8** 13 363.1 12 358.7 40 Basion–prosthion 55 94.7 4.90 55 95.0 20 94.3 83 94.4 41 100.2** 45 96.2 — 12 95.8 length 45 Bizygomatic breadth 202 123.5 4.57 57 123.9 20 125.8* 84 124.9* 44 130.1** 61 131.3** 6 130.2** 9 132.8** 46 Bimaxillary breadth 200 93.6 4.65 57 93.4 20 95.9* 84 93.5 46 96.5** 67 99.8** 11 95.9 12 100.8** 47 Total facial height 31 116.6 6.29 14 112.9 20 115.4 84 115.0 21 112.1** 45 116.3 9 104.9** 9 103.3** 48 Upper facial height 57 69.2 3.58 55 68.2 20 68.3 83 67.1** 41 65.6** 66 70.1 12 62.9** 10 63.1** 47/45 Total facial index (K) 31 95.0 6.35 14 90.8* 20 91.6* 84 92.2* 18 87.8** 34 88.7** 6 81.7** 6 78.1** 48/45 Upper facial index (K)1 57 56.1 3.28 55 55.0 20 54.6 83 53.8 35 50.9 49 53.7 6 49.2 6 48.0 47/46 Total facial index (V) 31 125.2 9.00 14 119.0* — 84 123.3 — 39 116.7** 9 109.5** — 48/46 Upper facial index (V) 57 74.1 4.63 55 72.9 20 71.5* 83 72.0** 38 67.9** 57 70.2** 11 65.6** 8 62.8** 51 Orbital breadth (left) 202 41.1 1.60 57 40.5* 20 41.0 84 41.1 45 41.8* 66 41.6* 10 41.1 10 41.6 52 Orbital height (left) 202 34.8 1.86 57 34.0** 20 34.4 84 33.8** 45 33.5** 65 34.1* 10 31.2** 7 33.0* 52/51 Orbital index (left) 202 84.7 4.39 57 83.9 20 83.0 84 82.4** 45 80.3** 62 82.0** 10 75.9** 7 81.0* 54 Nasal breadth 197 24.9 1.80 57 25.0 20 25.1 84 24.5 46 24.8 72 26.6** 12 26.6** 11 25.4 55 Nasal height 199 49.0 2.47 57 48.6 20 48.6 84 49.0 48 47.4** 71 49.8* 12 46.3** 11 45.2** 54/55 Nasal index1 196 50.8 3.98 57 51.4 20 51.2 84 50.2 45 52.1 69 53.5 12 57.4 10 56.1 72 Facial profile angle 54 83.5 2.79 55 83.0 20 82.4 83 83.6 41 81.4** 48 83.5 10 81.5* 10 81.8 1 t-test not performed because of rejection of normal distribution at the 0.05 level. *, ** significantly different at the 0.05 and 0.01 levels, respectively, from the northwestern Kyushu Japanese series (t-test).

Facial skeleton Kyushu Japanese. On the other hand, the curves of the The facial skeletons of the northwestern Kyushu and other Hokkaido Ainu, northern Kyushu Yayoi, northwestern modern Japanese were narrower in breadth measurements Kyushu Yayoi, and Tsukumo Jomon deviated far from the (Martin’s no. 45, 46, 51, and 54), greater in height measure- baseline. In particular, the amplitude of curves for the north- ments (Martin’s no. 47, 48, 52, and 55), larger in facial, western Kyushu Yayoi and Tsukumo Jomon was large, upper facial, and orbital indices, and smaller in the nasal indicating that there were large morphological differences index than those of the Hokkaido Ainu, northern Kyushu with the northwestern Kyushu Japanese. Yayoi, northwestern Kyushu Yayoi, and Tsukumo Jomon, except for nasal breadth of the Hokkaido Ainu, and facial, Multivariate analyses upper facial, orbital, and nasal heights of the northern Figure 3 shows the results of principal component analy- Kyushu Yayoi. Moreover, the northwestern Kyushu Japa- sis for 15 samples using the mean values of ten cranial mea- nese were taller than other modern Japanese women. surements (Martin’s no. 1, 8, 17, 40, 45, 48, 51, 52, 54, and 55), which are generally used, with no duplication of length, Mollison’s deviation curves height, and breadth measurements in the neurocranial, As shown in Figure 2, Mollison’s deviation curves, with facial, orbital, and nasal parts. The northwestern Kyushu the northwestern Kyushu Japanese as the baseline, depict the Japanese had a different first principal component score morphological relationships of the various groups well. The from the Tsukumo Jomon, northwestern Kyushu Yayoi, curves of the northern Kyushu, Kinki, and Kanto Japanese northern Kyushu Yayoi, Hokkaido Ainu, Miyako Islanders, did not deviate much from the baseline, indicating that these and Amami–Okinawa Islanders; and they had a different groups are morphologically close to the northwestern second principal component score from the Chinese and Vol. 112, 2004 MODERN FEMALE CRANIA IN NORTHWESTERN KYUSHU 151

Figure 2. Mollison’s deviation curves of the female cranial measurements and indices, using the northwestern Kyushu Japanese as the baseline of comparison.

western Kyushu Yayoi, and were also distant from the Hokkaido Ainu, Miyako Islanders, and Amami–Okinawa Islanders. Figure 4 shows the results of cluster analysis using the group average method based on the Penrose’s shape distances. The 15 groups were divided into two large clusters: one cluster consisted of the Tsukumo Jomon and northwestern Kyushu Yayoi, while the other cluster contained the remaining 13 groups. The latter cluster was further divided into a cluster consisting of the Hokkaido Ainu and the Amami–Okinawa and Miyako Islanders, and a cluster con- taining the other ten groups. Of those ten groups, the northwestern Kyushu, northern Kyushu, central Kyushu and Tohoku Japanese formed a cluster, and this cluster merged with a cluster comprising the Kinki Japanese, Kanto Japa- nese, and Koreans, and then with the Hokuriku Japanese, the northern Kyushu Yayoi, and Chinese, in that order.

Figure 3. Scatter diagram of first and second principal component Incidences of nonmetric cranial traits scores calculated from 10 cranial measurements of 15 female cranial The incidences of 22 nonmetric traits for the modern series. With the first principal component, strong positive factors are female crania of the northwestern Kyushu Japanese are bizygomatic breadth, maximum cranial breadth and maximum cranial shown in Appendix 2. length, while strong negative factors are upper facial height and basion-bregma height. In the second principal component, the positive factor is the whole size of the cranium. Abbreviations are the same as Comparison of incidence patterns those given in the footnote of Table 3. The incidences of 17 nonmetric traits are given in Table 4. When compared to the other modern Japanese, the incidence pattern of the northwestern Kyushu Japanese female was very similar to that of the northern Kyushu Japanese, and no Hokuriku Japanese. The northwestern Kyushu Japanese had significant difference existed in any of the traits. Although a score similar to the other modern Japanese and Koreans, the incidence pattern of the northwestern Kyushu Japanese and had the closest score to the Kanto and northern Kyushu was similar to that of the Kanto–Tohoku Japanese, signifi- Japanese. Table 3 shows Penrose’s shape distances calcu- cant differences were seen in three traits. Compared with the lated from ten mean values used in the principal component Hokkaido Ainu, there were significant differences in five analysis. The distances show that the northwestern Kyushu traits and the differences in these traits were marked. Japanese were closest to the northern Kyushu Japanese, fol- Although there were no significant differences in any trait lowed by the Kanto Japanese and Central Kyushu Japanese, compared with the northern Kyushu Yayoi, compared to the in that order. Conversely, the northwestern Kyushu Japanese northwestern Kyushu Yayoi and Jomon, there were signifi- were markedly distant from the Tsukumo Jomon and north- cant differences in five and seven traits, respectively. The 152 T. WATANABE ET AL. ANTHROPOLOGICAL SCIENCE

Table 3. Penrose’s shape distances among 15 female samples based on 10 cranial measurements NW- N- C- Kinki Hokuriku Kanto Tohoku Amami– Miyako Hokkaido Korean Chinese NK- NWK- Tsukumo Kyushu Kyushu Kyushu Okinawa Ainu Yayoi Yayoi Jomon NW-Kyushu1 — N-Kyushu2 0.0342 — C-Kyushu3 0.0943 0.0719 — Kinki4 0.1036 0.1187 0.1013 — Hokuriku5 0.1923 0.1778 0.2679 0.2793 — Kanto6 0.0796 0.0718 0.0775 0.0472 0.1845 — Tohoku7 0.2127 0.1409 0.0805 0.2519 0.2263 0.2088 — Amami– 0.4657 0.2900 0.3293 0.3225 0.3802 0.2618 0.3130 — Okinawa8 Miyako9 0.4726 0.3043 0.3188 0.3001 0.4032 0.2405 0.3274 0.0107 — Hokkaido 0.5550 0.4014 0.4578 0.4773 0.3456 0.3491 0.5014 0.2132 0.2298 — Ainu Korean 0.1912 0.2204 0.1810 0.0394 0.3903 0.0786 0.3626 0.4086 0.3535 0.6286 — Chinese 0.2524 0.2976 0.2503 0.3161 0.4797 0.2422 0.3873 0.8078 0.7481 0.8927 0.2657 — NK-Yayoi10 0.3169 0.2068 0.3138 0.2584 0.2823 0.2508 0.2876 0.2538 0.2599 0.3815 0.3320 0.4457 — NWK-Yayoi11 1.2925 1.0058 1.1828 1.0436 0.8828 0.9752 1.0426 0.3517 0.3898 0.4860 1.1578 1.6649 0.5032 — Tsukumo 1.3811 1.1909 1.2832 1.0418 0.8665 1.0783 1.1549 0.6488 0.6912 0.5572 1.1985 1.7639 0.6199 0.2650 — Jomon 1 Northwestern Kyushu Japanese, 2 Northern Kyushu Japanese, 3 Central Kyushu Japanese, 4 Kinki Japanese, 5 Hokuriku Japanese, 6 Kanto Japanese, 7 Tohoku Japanese, 8 Amami–Okinawa Islander, 9 Miyako Islander, 10 Northern Kyushu Yayoi, 11 Northwestern Kyushu Yayoi.

ern Kyushu Japanese and northern Kyushu Yayoi. They were also close to the Koreans, Hokuriku, and Kanto– Tohoku Japanese, and somewhat close to the Amami– Okinawa Islanders, Kinki Japanese, and Chinese. On the other hand, the northwestern Kyushu Japanese were markedly distant from the Hokkaido Ainu, northwestern Kyushu Yayoi and Jomon. A principal coordinate analysis using the MMDs was conducted to prepare a two-dimensional diagram (Figure 5). The 12 samples were divided into two major clusters: one cluster consists of the Jomon, northwestern Kyushu Yayoi, and Hokkaido Ainu on the positive side of axis I, the other cluster consists of the remaining nine samples placed on the negative side. In the latter cluster, the Chinese, Amami–Okinawa Islanders, and Kinki Japanese were slightly distant from the six samples comprising the northwestern Kyushu, northern Kyushu, Hokuriku and Kanto–Tohoku Japanese, Koreans, and northern Kyushu Yayoi. The northwestern Kyushu Japanese was located near the center.

Figure 4. Group average clustering analysis based on the Pen- rose’s shape distances matrix shown in the Table 3. Abbreviations are Discussion the same as those given in the footnote of Table 3. Regionality of modern female populations in the Japa- nese archipelago The regionality and formation of modern people in the degree of difference in traits that are considered important Japanese archipelago is well explained by the ‘dual structure for ascertaining inter-group lineage relationships, such as model’ (K. Hanihara, 1991). According to this model, the supraorbital foramen, hypoglossal canal bridging, and modern Japanese are a mixture of two fundamentally dis- transverse zygomatic suture vestige (Dodo, 1987; Dodo and tinct groups, an earlier group of the Jomon, and a later group Ishida, 1990; Kozintsev, 1990; Saiki et al., 2000), was par- of immigrants who appeared during the Yayoi period. More- ticularly large. over, the Ainu and Ryukyuans are regarded as representing the relatively unmixed descendants of the Jomon. This Multivariate analyses model is supported by many morphological and genetic Smith’s MMDs were calculated among 12 samples based findings (T. Hanihara, 1991, 1992; Matsumura, 1995; on the incidences of the 17 traits (Table 5). Northwestern Omoto, 1995; Omoto and Saito, 1997; Tokunaga et al., Kyushu Japanese females were markedly close to the north- 2001), and also is consistent with the results of virological Vol. 112, 2004 MODERN FEMALE CRANIA IN NORTHWESTERN KYUSHU 153

Table 4. Comparison of side incidences of 17 nonmetric traits in modern, Yayoi and Jomon female crania Northwestern Northern Kinki Japanese Kanto–Tohoku Hokkaido Ainu Northern Northwestern Jomon Kyushu Kyushu Japanese Kyushu Yayoi Kyushu Yayoi Traits Japanese Japanese n % n % n % n % n % n % n % n % Metopism1 204 4.4 27 11.1 85 14.1** 50 6.0 79 2.5 57 7.0 46 0.0 66 16.7** Supraorbital 408 12.5 54 14.8 170 17.6 98 27.6** 100 15.0 79 17.7 50 24.0* 100 17.0 nerve groove Supraorbital 407 43.2 54 33.3 170 38.8 100 42.0 148 14.2** 98 40.8 63 12.7** 108 8.3** foramen Ossicle at the 158 5.1 27 3.7 79 3.8 46 2.2 80 1.3 35 11.4 40 10.0 61 4.9 lambda1 Ossicle at asterion 393 5.3 52 3.8 165 6.1 96 6.3 158 7.0 58 3.4 71 7.0 88 8.0 Occipitomastoid 384 8.9 54 13.0 162 17.9** 98 8.2 158 13.9 45 6.7 38 7.9 52 15.4 ossicle Parietal notch bone 402 20.6 50 16.0 167 34.1** 92 17.4 156 20.5 55 20.0 69 7.2** 68 11.8 Condylar canal 405 74.3 54 64.8 170 75.3 98 73.5 156 80.2 37 59.5 22 68.2 42 88.1* Precondylar 370 4.6 54 3.7 164 15.9** 100 3.0 136 4.4 53 7.5 44 0.0 68 4.4 tubercle Paracondylar 327 0.6 50 0.0 164 5.5** 88 0.0 108 7.4** 33 3.0 16 6.3 12 8.3 process Hypoglossal 407 11.1 54 7.4 170 10.0 100 6.0 156 23.7** 69 11.6 60 33.3** 64 21.9* canal bridging Tympanic 408 42.9 54 53.7 170 47.6 100 26.0** 154 19.5** 79 38.0 89 40.4 102 35.3 dehiscence Foramen ovale 407 2.7 54 1.9 170 1.2 100 2.0 152 3.9 71 2.8 53 0.0 36 5.6 incomplete Pterygospinous 408 2.0 54 0.0 170 0.0 100 0.0 126 1.6 68 1.5 35 2.9 48 0.0 foramen Medial palatine 391 3.1 52 3.8 170 9.4** 96 4.2 122 22.1** 69 5.8 23 4.3 56 8.9* canal Transverse zygomatic 407 7.4 52 3.8 165 11.5 94 16.0** 76 13.2 47 8.5 50 36.0** 54 24.1** suture vestige Mylohyoid 398 3.0 20 0.0 168 4.8 96 3.1 108 6.5 49 6.1 66 6.1 100 11.0** bridging 1 Median traits. *, ** significantly different at the 0.05 and 0.01 levels, respectively, from the northwestern Kyushu Japanese series (chi-square or exact probability test).

Table 5. Smith’s mean measure of divergences (MMDs) among 12 female samples based on 17 nonmetric traits NW- N- Kinki Hokuriku Kanto– Amami– Hokkaido Korean Chinese NK- NWK- Jomon Kyushu Kyushu Tohoku Okinawa Ainu Yayoi Yayoi NW-Kyushu1 — N-Kyushu2 0.0064 — Kinki3 0.0338 0.0180 — Hokuriku4 0.0090 0.0062 0.0272 — Kanto–Tohoku5 0.0133 0.0120 0.0430 0.0196 — Amami–Okinawa6 0.0334 0.0240 0.0082 0.0018 0.0274 — Hokkaido Ainu 0.0800 0.0878 0.0765 0.0945 0.0680 0.0738 — Korean 0.0089 0.0160 0.0361 0.0311 0.0097 0.0215 0.0523 — Chinese 0.0384 0.0415 0.0597 0.0621 0.0206 0.0479 0.1010 0.0116 — NK-Yayoi7 0.0030 0.0095 0.0161 0.0126 0.0075 0.0271 0.0636 0.0043 0.0057 — NWK-Yayoi8 0.0893 0.0958 0.1302 0.0869 0.0792 0.1066 0.0578 0.0959 0.0886 0.0637 — Jomon 0.0882 0.0767 0.0668 0.1016 0.0767 0.0656 0.0225 0.0625 0.0936 0.0699 0.0345 — 1 Northwestern Kyushu Japanese, 2 Northern Kyushu Japanese, 3 Kinki Japanese, 4 Hokuriku Japanese, 5 Kanto–Tohoku Japanese, 6 Amami– Okinawa Islander, 7 Northern Kyushu Yayoi, 8 Northwestern Kyushu Yayoi. studies (Hinuma, 1985, 1986). ing to the metrical and nonmetrical analyses (Figure 3, Fig- The regional features of modern females of the Japanese ure 4, Figure 5); (2) the Hokkaido Ainu and Ryukyuans are archipelago identified in this study may be summarized as comparatively close and form one cluster in the metrical follows: (1) the modern peoples of mainland Japan, includ- analyses but are unlike the mainland Japanese (Figure 3, ing the northwestern Kyushu Japanese, are relatively close Figure 4), while the Ryukyuans are distant from the to the Koreans, Chinese, and northern Kyushu Yayoi accord- Hokkaido Ainu and close to the mainland Japanese 154 T. WATANABE ET AL. ANTHROPOLOGICAL SCIENCE

al., 1998, 2000; Manabe et al., 1999; Pietrusewsky, 1999; Dodo, 2001; Higa et al., 2003). The nonmetrical findings of this study support these reports.

Morphological characteristics of female crania in the northwestern Kyushu Japanese As briefly mentioned in the introduction, modern male crania (n 143) of the northwestern Kyushu Japanese are metrically characterized by a lower cranial vault, lower face, and lower orbit compared with those of the other modern Japanese and Koreans (Figure 6, reproduced from Noda, 1993). Moreover these characteristics are generally similar to the condition seen in the Jomon and northwestern Kyushu Yayoi series. For this reason, Noda (1993) suggested that the modern male Japanese of the northwestern Kyushu area was genetically influenced by the northwestern Kyushu Yayoi. We calculated Penrose’s shape distances of the modern samples compared by Noda (1993) to the northern Kyushu Yayoi, northwestern Kyushu Yayoi, and Tsukumo Jomon series. The findings shown in Table 6 indicate that the males of the northwestern Kyushu Japanese (as well as the other modern Japanese) were closer to the northern Kyushu Yayoi Figure 5. Two-dimensional display of the principal coordinate than to the northwestern Kyushu Yayoi or Tsukumo Jomon, analysis based on the MMD matrix shown in Table 5. The cumulative but that among the six modern samples compared, they were proportion is 67.6%. Abbreviations are the same as those given in the footnote of Table 5. the closest to the northwestern Kyushu Yayoi and Tsukumo Jomon. The following conclusion may be drawn from the metrical analyses of female crania of the northwestern Kyushu Japa- according to the nonmetrical analysis (Figure 5). These find- nese: (1) the height of the neurocranium is relatively great in ings are in agreement with the dual structure model in the relation to length and breadth; and (2) facial breadth is rela- metrical analyses, but somewhat in disagreement in the non- tively narrow and facial height is relatively great (Table 2, metrical analysis. In recent years, many studies have found Figure 2). These characteristics were particularly marked in that the Ryukyuans share more morphological similarities the northwestern Kyushu Japanese, but are also common in with modern mainland Japanese than with the Ainu (Dodo et modern Japanese. The results of principal component analy-

Figure 6. Mollison’s deviation curves of the male cranial measurements and indices, using the northwestern Kyushu Japanese as the baseline of comparison (reproduced from Noda, 1993). Length–height index (17:1), breadth–height index (17:8), upper facial height (48), upper facial indices (48:45, 48:46), and orbital index (52:51) of the five cranial series are all situated in the positive side of the baseline. In other words, the male of the northwestern Kyushu Japanese has a lower cranial vault, face, and orbit compared to the five populations. Vol. 112, 2004 MODERN FEMALE CRANIA IN NORTHWESTERN KYUSHU 155

Table 6. Penrose’s shape distances from six modern Japanese and not the direct descendants of the northwestern Kyushu Korean male series (used in Noda, 1993) to two Yayoi and Yayoi, and are genetically closer to the northern Kyushu Jomon male series, based on 10 cranial measurements Yayoi. Dodo and Ishida (1992) investigated the incidence of Northern Northwestern Tsukumo nonmetric cranial traits at specific periods spanning the Kyushu Yayoi Kyushu Yayoi Jomon Jomon to the modern, and stated, “The northern Kyushu Northwestern 0.2118 0.5251 0.8388 Yayoi of continental lineage must be one of the forerunners Kyushu Japanese of the modern Japanese of the central islands of Japan.” Northern Kyushu 0.1390 0.5792 0.9517 Saiki et al. (2000) offered the same conclusion. The results Japanese of this study also support the opinions of Dodo and Ishida Central Kyushu 0.1584 0.6912 1.1920 Japanese (1992). Kinki Japanese 0.1936 0.5947 1.1084 In the northwestern Kyushu area, hardly any human skel- Kanto Japanese 0.2011 0.6169 0.9968 etal remains have been excavated from the protohistoric Korean 0.3368 0.9805 1.5097 Kofun period (ca. 300 AD to 700 AD) to recent. It is therefore difficult to ascertain temporally based morphological changes of the skeleton. According to the results of metrical and nonmetrical analyses of the female crania, it can at least sis in Figure 3 show these findings clearly. With the first be suggested that people in the northwestern Kyushu area principal component, high scores represent broader and inherited the characteristics of the Jomon people up to the lower cranial vault and facial skeleton, while low scores rep- Yayoi period, and subsequently—during and after the Kofun resent narrower and higher vault and face. As a result, all period—were influenced by people migrating from the con- modern Japanese are on the negative side, and the north- tinent or their descendants. The resulting changes produced western Kyushu Japanese have the lowest score. In contrast, the characteristics seen in the modern Japanese people. the Tsukumo Jomon and northwestern Kyushu Yayoi, with a lower neurocranium and a lower, wider face, have the high- Acknowledgments est scores. The craniometric characteristics of the northwestern The authors wish to express our sincere thanks to Profes- Kyushu Japanese differ to some extent between males and sor Seiji Nagashima, Graduate School of Biomedical Sci- females. Consequently, we consider that it is necessary to ences, Nagasaki University for his guidance. We also wish to examine not only males but also females when discussing thank Professor Yukio Dodo, Graduate School of Medicine, and understanding regionality of the modern Japanese popu- Tohoku University for providing us with the raw data of the lation. nonmetric traits of the modern Japanese in northern Kyushu The following may be concluded from the nonmetrical area, Professor Takahiro Nakahashi, Graduate School of analyses: (1) the incidence pattern seen in the northwestern Social Science and Cultural Studies, Kyushu University, Kyushu female Japanese is quite similar to that of the other with unpublished measurements of the same modern Japa- modern Japanese; (2) compared to the Hokkaido Ainu, nese, and Professor Hajime Ishida, Faculty of Medicine, northwestern Kyushu Yayoi, and Jomon, incidences of the University of the Ryukyus for observational advice on the supraorbital foramen, hypoglossal canal bridging, and nonmetric cranial traits. transverse zygomatic suture vestige were particularly large (Table 4). 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Appendix 1. Measurements and indices of modern female crania of the northwestern Kyushu area (in mm except for angular measurements) Metrical characters with n Mean S.D. Min. Max. Martin’s number 1. Maximum cranial length 201 172.0 5.79 159 186 5. Basion-nasion length 200 95.4 3.70 82 106 7. Foramen magnum length 198 33.7 2.13 28 42 8. Maximum cranial breadth 201 134.6 5.15 121 148 9. Least frontal breadth 201 89.6 3.95 81 103 9/8 Transverse frontparietal index 200 66.7 3.34 58.1 76.0 10. Maximum frontal breadth 193 109.9 4.94 95 125 11. Biauricular breadth 202 120.3 4.30 110 135 12. Biasterionic breadth 200 104.8 3.79 94 116 13. Bimastoid breadth 201 97.5 4.35 87 109 16. Foramen magnum breadth 200 28.6 1.84 24 34 17. Basion–bregma height 199 131.8 4.08 123 141 8/1 Length–breadth index 201 78.3 4.00 68.8 91.9 17/1 Length–height index 199 76.7 2.85 70.7 85.6 17/8 Breadth–height index 199 98.1 4.07 87.0 108.8 23. Horizontal circumference 197 491.4 12.29 461 521 24. Transverse arc 198 304.3 10.42 278 330 25. Total sagittal arc 198 359.0 11.89 327 392 26. Frontal sagittal arc 200 121.8 6.01 104 139 27. Parietal sagittal arc 200 120.3 7.64 102 137 28. Occipital sagittal arc 198 116.8 7.82 97 141 29. Frontal sagittal chord 200 107.2 4.31 96 121 30. Parietal sagittal chord 200 108.0 5.88 94 124 31. Occipital sagittal chord 199 98.0 5.40 88 114 26/25 Frontosagittal index 198 33.9 1.29 29.5 38.2 27/25 Parietosagittal index 198 33.5 1.85 28.1 38.3 28/25 Occipitosagittal index 198 32.5 1.82 28.1 38.7 40. Basion-prosthion length 55 94.7 4.90 81 103 42. Basion-gnathion length 72 103.3 5.63 87 119 43. Outer biorbital breadth 201 98.5 4.28 71 107 44. Biorbital breadth 202 94.1 3.58 82 103 45. Bizygomatic breadth 202 123.5 4.57 110 134 46. Bimaxillary breadth 200 93.6 4.65 80 107 47. Total facial height 31 116.6 6.29 101 130 48. Upper facial height 57 69.2 3.58 60 77 47/45 Total facial index (Kollmann) 31 95.0 6.35 84.8 107.4 47/46 Total facial index (Virchow) 31 125.2 9.00 108.6 147.7 48/45 Upper facial index (Kollmann) 57 56.1 3.28 49.2 62.8 48/46 Uupper facial index (Virchow) 57 74.1 4.63 64.5 84.9 50. Anterior interorbital breadth 202 16.6 1.87 12.0 21.3 50/44 Interorbital index 202 17.7 1.70 12.2 22.7 51. Orbital breadth (right) 202 41.3 1.65 36 46 (left) 202 41.1 1.60 36 46 52. Orbital height (right) 202 34.8 1.80 30 40 (left) 202 34.8 1.86 30 39 52/51 Orbital index (right) 202 84.4 4.13 69.8 93.0 (left) 202 84.7 4.39 72.1 95.1 54. Nasal breadth 197 24.9 1.80 21 30 55. Nasal height 199 49.0 2.47 40 57 54/55 Nasal index 196 50.8 3.98 42.0 62.8 57. Least nasal breadth 201 7.2 1.74 2.4 12.2 60. Maxilloalveolar length 52 51.6 2.90 42 58 61. Maxilloalveolar breadth 27 61.9 2.97 56 67 61/60 Maxilloalveolar index 26 119.6 6.82 103.4 132.0 72. Facial profile angle 54 83.5 2.79 77 91 73. Middle facial profile angle 185 87.3 3.01 79 95 74. Alveolar profile angle 54 71.2 5.79 60 86 Vol. 112, 2004 MODERN FEMALE CRANIA IN NORTHWESTERN KYUSHU 159

Appendix 2. Incidences of 22 nonmetric traits of the modern female crania in the northwestern Kyushu area Side incidence Cranial incidence Traits Right Left Total n % n % n % n % 1 Metopism1 — — — 204 4.4 2 Supraorbital nerve groove 204 10.8 204 14.2 408 12.5 204 19.1 3 Supraorbital foramen 203 45.8 204 40.7 407 43.2 203 54.7 4 Ossicle at the lambda1 — — — 158 5.1 5 Biasterionic suture vestige 200 5.0 198 1.5 398 3.3 198 5.6 6 Ossicle at asterion 196 5.1 197 5.6 393 5.3 195 9.2 7 Occipitomastoid ossicle 192 7.8 192 9.9 384 8.9 189 14.8 8 Parietal notch bone 200 24.0 202 17.3 402 20.6 199 32.2 9 Condylar canal patent 204 80.4 201 68.2 405 74.3 201 88.6 10 Precondylar tubercle 191 5.2 179 3.9 370 4.6 177 6.8 11 Paracondylar process 164 1.2 163 0.0 327 0.6 150 0.7 12 Hypoglossal canal bridging 203 10.8 204 11.3 407 11.1 203 18.7 13 Tympanic dehiscence 204 43.6 204 42.2 408 42.9 204 53.9 14 Foramen ovale incomplete 203 3.0 204 2.5 407 2.7 203 5.4 15 Foramen of Vesalius 199 41.7 202 51.5 401 46.6 199 61.3 16 Pterygospinous foramen 204 1.5 204 2.5 408 2.0 204 3.4 17 Medial palatine canal 194 4.1 197 2.0 391 3.1 193 5.2 18 Transverse zygomatic suture vestige 204 6.9 203 7.9 407 7.4 203 9.9 19 Clinoid bridging 187 3.2 186 3.8 373 3.5 182 6.0 20 Mylohyoid bridging 199 3.0 199 3.0 398 3.0 199 5.5 21 Juglar foramen bridging 204 11.8 204 5.9 408 8.8 204 14.7 22 Sagittal sinus groove left1 ———19617.3 1 Median traits.