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Quaternary International 211 (2010) 42–54

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A multivariate analysis of measurements recorded in early and more moderncrania from East Asia and Southeast Asia

Michael Pietrusewsky*

Department of Anthropology, University of Hawaii, 2424 Maile Way, Saunders 346, Honolulu, Hawaii 96822, USA

a r t i c l e i n f o

Article history:Available online 13 January 2009

* Tel.: þ1 808 956 6653; fax: þ1 808 956 9541.E-mail address: mikep@hawaii.edu

1040-6182/$ – see front matter � 2009 Elsevier Ltd adoi:10.1016/j.quaint.2008.12.011

a b s t r a c t

Stepwise discriminant function analysis and Mahalanobis’ generalized distance are applied to 28 land-mark measurements recorded in 38 prehistoric and modern cranial series from Eastern Asia for inter-preting biological relationships and population history. The cranial series are from Japan, China, NorthernAsia, mainland Southeast Asia, and island Southeast Asia. The results of this analysis indicate a markedseparation of East/North Asian and Southeast Asian cranial series, a finding that supports hypotheses oflong-term continuity in northern and southern regions of Eastern Asia rather than models that suggestintrusion and replacement. The results also support a common recent origin of the Chinese, Japanese,and Koreans in Northeast Asia. A major intrusion of people into the Japanese archipelago beginning inthe Yayoi Period is further supported by these results. The relationships of Ainu, Jomon, RyukyuIslanders, and Taiwan Aboriginals are discussed.

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1. Introduction

Comprehending human biological variation and human originshas been important in physical anthropology since the discipline’sinception. Earlier attempts to understand human variation throughthe construction of typological classifications of race failed largelydue to the limitations of the theoretical approach and the lack ofappropriate statistical methods. Breakthroughs in evolutionarybiology and improvements in statistical methods now providea much sounder approach for measuring and interpreting variationwithin and between human groups.

While no longer conveying the initial meaning of earlier typo-logical classifications, labels such as Caucasoid, Negroid,Mongoloid, and Australoid provide a starting point for describinghuman variation for major geographical regions of the world.Mongoloid is the label most commonly applied to the people ofEastern Asia and Southeast Asia and groups derived from these,including Pacific Islanders and Native Americans (Bowles, 1977;Howells, 1986, 1992). While considerable heterogeneity is foundamong this division of humanity (Bowles, 1977:344), a southern(e.g., Southeast Asia) and a northern (e.g., China, Korea, Japan,northeastern Siberia) subdivision are often further recognized(Howells, 1986; Bellwood, 1997). Although not substantiated inlater work (e.g., Steegmann, 1970), several of the identifying

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features of northern Mongoloids (e.g., non-projecting noses, facialflatness, round eye sockets, brachycranic vaults, etc.) were viewedas representing adaptations to the extreme dry cold conditions ofthe Late Pleistocene (Coon, 1965). These features appear before theexpansion of rice agriculturalists at the beginning of the Holocene.While the southern Mongoloids possess many of the same cranialfeatures found in northern Mongoloids, these features are lessintense and of a more generalized nature in the southern groups(Bowles, 1977; Howells, 1986, 1992).

Interpretations of when (and where) these physical character-istics first appeared in Eastern Asians have generated two con-trasting viewpoints commonly referred to as the continuity anddiscontinuity models. Examining the paleoanthropological recordfor the region, some researchers (e.g., Kamminga and Wright, 1988;Kamminga, 1992; Wright, 1995; Brown 1998, 2001; Cunninghamand Wescott, 2002) maintain the features of dental and facialmorphology seen in Eastern Asians since the early Neolithic are notfound in Late Pleistocene hominin specimens such as the Zhou-koudian Upper Cave 101, Liujiang, or Minatogawa. Others (e.g.,Wolpoff, 1995; Wu and Poirier, 1995), have interpreted the earlyhuman fossil record for the region differently. These investigatorscontend that there are strong morphological links between theanatomically modern Homo sapiens in China and earlier datedhominin specimens in this region right back to Homo erectus.Likewise, Etler (1996) maintains that East Asian specimens, espe-cially those from northern China, although displaying a mosaic oftransitional features, demonstrate morphological continuitybetween archaic and modern humans in Eastern Asia. In his

M. Pietrusewsky / Quaternary International 211 (2010) 42–54 43

assessment, the Late Pleistocene specimens from southern China,such as the Liujiang cranium from Guangxi dated at 67,000 yearsago (but see Shen et al., 2002), show typical Mongoloid featuresincluding shovel-shaped incisors. Further, the retention of diag-nostic facial and dental features in Eastern Asian skeletal materialsuggests continuity rather than replacement from Homo erectus topre-modern and modern East Asians (Etler, 1996).

Further evidence for discontinuity, based mainly on archaeo-logical and historical linguistic data, has been provided by Bellwoodand others (e.g., Blust, 1996; Higham, 2001; Diamond and Bell-wood, 2003; Bellwood, 2005), who have proposed an agriculturallydriven language expansion model to explain population move-ments in Eastern Asia, particularly Southeast Asia, beginning in theNeolithic onward. According to this model rice domestication/farming and languages spread from a source in southern China. Intheir view the immigrant early farmers eventually replace theearlier in situ hunter–gatherers of Southeast Asia who shareaffinities with Australian Aboriginals, Melanesians, and Negritos.This traditional view has also been referred to as the admixture/replacement model.

Except for work by Matsumura and colleagues (e.g., Matsumuraand Hudson, 2005), there has not been wide support of theadmixture/replacement model from studies in physical anthro-pology. Rather, the majority of work in physical anthropology (e.g.,Bulbeck, 1982; Turner, 1987, 1992; Pope, 1992; Hanihara, 1993;Pietrusewsky, 2006) supports continuity between earlier and laterinhabitants of the region. In this view the spread of farming is seenmainly as the result of the adoption and/or diffusion of agriculturaltechnology by the descendents of in situ hunter–gatherers withoutnecessarily the spread of new languages or genes.

Most notably, the work by Turner (1987, 1990, 1992), usingdental non-metric traits, has identified two major dental complexesin this region. The first, Sundadonty, is expressed in the moderninhabitants of Southeast Asia, Polynesia, Micronesia, and southernChina. The distribution of the second dental complex, Sinodonty, ismore northern and includes the inhabitants of northeastern Asia(e.g., northern China, Japan, and Korea) and Northern Asian-derivedgroups such as those in the Americas. In Turner’s view the morespecialized Sinodont pattern found in Northeast Asia arose fromthe more generalized Sundadont pattern in Southeast Asia asa result of gene flow running from south to north along the EasternAsian corridor.

Molecular genetic work (e.g., Jin and Su, 2000; Shi et al., 2005)also supports an initial arrival of modern humans in southern Chinawith subsequent northward expansion. Other, more recent, work inphysical anthropology and archaeology has identified systematicdifferences between northern and southern groups within EasternAsia (Barnes, 1999; Zhang, 1999; Wu et al., 2007). However, recentgenetic studies (e.g., Ding et al., 2000) find no support for a majornorth/south division within China and Eastern Asia while an anal-ysis of the mitochondrial DNA molecule by Kivisild et al. (2002)arrives at an opposite conclusion. Likewise, molecular genetic workby Shi et al. (2005) supports a southern origin of modern humans inEastern Asia with subsequent northward migration while studies ofY chromosome (NRY) data (e.g., Karafet et al., 2001) suggest a morecomplex scenario.

Of equal importance for understanding the origins of the peopleof Eastern Asia is the relationship between the modern inhabitantsof the Korean peninsula, the Japanese archipelago, and theremaining regions of Eastern Asia and the origin of the prehistoricand modern people of the Japanese archipelago. The prehistoricJomon are assumed to be the descendents of Pleistocene hunter–gatherers who migrated to Japan approximately 30,000 years ago(Imamura, 1996; Kobayashi, 2004) from either Southeast Asia(Hanihara, 1991, Turner, 1992) or northern central Asia (Omoto and

Saitou, 1997; Pietrusewsky 1999). Much later, approximately 2500years ago, Yayoi agriculturalists began to migrate into the Japanesearchipelago bringing with them wet-rice agriculture from a regionthat today corresponds to Korea and northern China (Hudson, 1999;Imamura, 1996). This later population expansion, which wasaccompanied by varying degrees of interbreeding with the indig-enous Jomon foragers of Japan, is assumed to be largely responsiblefor the origin of the modern Japanese (Hanihara, 1991). Of furtherinterest is the relationship of modern Japanese to Jomon, Ainu, andRyukyu Islanders and the relationship of Taiwan’s Aboriginalpeoples to the present day peoples of Eastern Asia.

Although outside the scope of the present analysis, readers whowish to view the evolution of East Asians within the broadercontext of modern human origins and connections to the earlyinhabitants of the Americas should consult Wright (1992), Lahr(1995), Brace et al. (2001), and Cunningham and Jantz (2003)among others.

Physical anthropology, especially studies of human skeletalremains that span the transition from hunting and gathering toagriculture in Eastern and Southeast Asia, should help demonstratewhether the indigenous peoples of Eastern Asia and Southeast Asiawere displaced by later colonists or if population continuity, char-acterized by a common genetic heritage of people, bridged thetechnological and social transition in this region. Alternatively, theevidence from physical anthropology may necessitate an entirelydifferent scenario.

The primary focus of this paper is to investigate, usingmeasurements recorded in prehistoric and modern crania, thehistorical–biological relationships of the peoples of Eastern Asiaand surrounding regions. Comparisons of these results with thosebased on genetic, archaeological, and historical linguistic data willbe used to evaluate some of the competing hypotheses that relateto the origins and peopling of Eastern Asia.

While morphological variation, including craniometric varia-tion, is subject to non-genetic or environmental influences, thiscategory of variation is generally viewed as reflecting geneticsimilarity and provides the basis for the use of this category ofvariation in biodistance studies (Van Vark and Howells, 1984;Buikstra et al., 1990; Kohn, 1991). Concordance in results based onanthropometric and quantitative genetic analyses strengthensthese assertions, which allow distances based on metric data to beinterpreted within a population genetic framework (Konigsbergand Ousley, 1995; Relethford, 2002; Roseman, 2004; Roseman andWeaver, 2004).

2. Material and methods

2.1. Cranial series

Because multivariate statistical procedures do not toleratemissing variables, only complete, or nearly complete, adult malecrania are used in the present study. Comparable data for femaleadult specimens were not available for this study. Information,including the number of crania and their origin, for cranial seriesfrom Eastern Asia and Southeast Asia are given in Table 1. A mapshowing the location of these cranial series is provided in Fig. 1. Themodern Chinese series include crania from eastern and westernprovinces, Hong Kong, Hainan Island, and Taiwan. Two cranialseries are from northeastern Asia, including Manchuria (nowcommonly referred to as northeastern China) and Korea, and one isfrom Mongolia.

While the majority of the cranial series used in this analysisrepresent modern and near modern specimens, a few of the seriesrepresent earlier time periods. Included in these earlier dated series

Table 1Thirty-eight male cranial series used in the present analysis.

Series name (abbreviation)a Samplesize

Collection locationand number of specimensb

Remarks

Eastern and Northern AsiaShanghai (Sha) 50 SHA-50 The specimens are mostly from post-Qing cemeteries in Shanghai.Hangzhou (Han) 50 SHA-50 Ancient skeletal remains exhumed in the modern city of Hangzhou,

Zhejiang Province in eastern China.Nanjing (Nan) 49 SHA-49 Ancient remains exhumed from the modern city of Nanjing, Jiangsu

Province in eastern China.Chengdu (Che) 53 SHA-10; CHE-43 A majority of these specimens date to the Qing Dynasty (1644 A.D.–1911 A.D.)

and are from Chengdu, Sichuan Province in western China. Ten crania arefrom Leshan, Lizhong County, Sichuan Province.

Hong Kong (Hk) 50 HKU-50 Specimens represent individuals who died in Hong Kong between 1978 and 1979.Taiwan Chinese (Tai) 47 TPE-47 Modern Chinese living in Taiwan who trace their immediate origins to Fujian

and Guangdong Provinces on the mainland of China.Hainan Island (Hai) 47 TPE-47 Chinese immigrants originally from the Guangdong region of China who began

arriving around 200 B.C. This material was excavated by T. Kawasaki in HaikouCity on Hainan Island.

Anyang (Any) 56 TPE-56 Bronze-Age (11th century B.C.) Shang Dynasty sacrificial victims excavated atAnyang in northern Henan Province in northern China (Howells, 1983; Li, 1977).

Manchuria (Man) 50 TKO-50 Many of the specimens are from northeastern China or the region formerly referredto as ‘‘Manchuria,’’ which today includes Heilongjiang and Jilin Provinces andadjacent northern Korea. A great many of these specimens are identified assoldiers or cavalrymen who died in battle in the late 19th century.

Korea (Kor) 32 KYO-7; SEN-3; TKM-2; TKO-20 Specific locations in Korea are known for most of these specimens.Mongolia (Mog) 50 SIM-50 The skulls are identified as coming from Ulaanbaatar (Urga), Mongolia and were

purchased by A. Hrdlika in 1912.Atayal (Aty) 36 TPE-28; TKM-7; TKO-1 The specimens in Taipei represent slain victims of Atayal, the second largest

surviving Aboriginal tribe in Taiwan. The incident took place in 1932 and thespecimens were collected by T. Kawasaki in the same year (Howells, 1989).

JapanKanto Japanese (Kan) 50 CHB-50 A dissecting room sample of modern Japanese from the Kanto District of eastern

Honshu Island. The majority of the individuals were born during the Meiji Period(1868–1911) and most died well before 1940.

Tohoku Japanese (Toh) 53 SEN-53 Dissecting room specimens of modern Japanese from the Tohoku District innorthern Honshu Island.

Kyushu Japanese (Kyu) 51 KYU-51 Modern Japanese which derive mostly from Fukuoka Prefecture in Kyushu Island.Other specimens are from Yamaguchi, Saga, Nagasaki, and adjoining Prefectures.

Edo (Edo) 55 NSM-55 The specimens are from the Joshinji (Tokyo) site and date to the Edo Period orapproximately the 17th to mid-19th centuries.

Kamakura (Kam) 52 NSM-9; TKO-43 Specimens are from the Medieval mass burial sites of Zaimokuza and Gokurakuji inthe city of Kamakura, victims of a war which occurred in 1333.

Marunouchi(Mar) 27 NSM-26; TKO-1 Specimens are from the Medieval Muromachi Period (1338–1603 A.D.) that wereexcavated from 16 temples in Marunouchi, Chiyoda-ku, Tokyo, by Hisashi Suzuki.

Kofun (Kof) 62 KYO-5; KYU-53; NSM-4 The Kofun Period (4th to 6th century A.D.) follows the Yayoi Period.Yayoi (Yay) 62 KYU-62 A combined sample of Yayoi specimens from Doigahama (39), Yoshimohama (14), and

Nakanohama (2) sites in Yamaguchi Prefecture and from the Koura (7) site in ShimanePrefecture. The dates for the Yayoi Period are approximately 300 B.C.–300 A.D.

Ainu (Ain) 50 SAP-18; TKM-5; TKO-27 Skeletons collected by Yoshikiyo Koganei in 1888–1989 from abandoned Ainucemeteries in Hokkaido (Koganei, 1883–1884).

Jomon (Jom) 51 TKO-16; NSM-19; KYO-15; SAP-1 All specimens represent Late to Latest [ca 3500 years B.P. to 2000 years B.P.] Jomonsites on Honshu Island. The largest series are Ebishima (11) in Iwate Prefecture inTohoku District and Tsukumo (12), Okayama Prefecture in the Chugoku District.

Amami Islands (Ama) 31 KYO-19; KYU-12 Specimens are from Tokunoshima (19) and Yoro Islands (12) of the Amami Islands,which are located north of the Okinawa Group in the central Ryukyu (or Nansei)Islands. Yoro Island, a small island off the southern tip of Amami-Oshima Island, of the AmamiGroup.

Okinawa Island (Oki) 53 KYO-11; KYU-1; TKO-1; GYO-19;KAN-21

Specimens are from Okinawa Island and neighboring Kume Island in the centralRyukyu Islands; 19 are from Gyokusendo Cave, located in southeastern Okinawa(18th A.D. century) and 21 are two different locations on Kume Island, an islandlocated west of Okinawa Island: Yattchi (17) and Hiyajo (4); the remainder are fromOkinawa Island.

Sakishima Group (Sak) 24 RYU-11 KYU-5; TKO-8 The specimens are from five separate islands in the Sakishima Group of the southernRyukyu Islands: Hateruma (2); Miyako (6); Iriomote (3); Ishigaki (1), andYonaguni Islands (12).

Mainland and Island Southeast AsiaVietnam (Vtn) 49 HCM-49 Specimens are from Hanoi (Van Dien Cemetery) and Ho Chi Minh City in Vietnam

(Pietrusewsky, 1992:13).Bachuc Village (Bac) 51 BAC-51 Victims of the 1978 Khmer Rouge massacre in Bachuc Village in western Angiang Province,

Vietnam (Pietrusewsky, 1992:13).Cambodia and Laos (Cml) 40 PAR-40 A combined sample of crania from various locations in Cambodia and Laos collected

between 1877 and 1920.Thailand (Thi) 50 SIR-50 Most of the specimens represent dissecting room cases from Bangkok.

M. Pietrusewsky / Quaternary International 211 (2010) 42–5444

Table 1 (continued )

Series name (abbreviation)a Samplesize

Collection locationand number of specimensb

Remarks

Burma (Bur) 16 ZUR-16 The crania in Zurich are from a series (Cat. Nos. 93-125) of skulls collected inMandalay, Myanmar (Burma), described in a catalogue dated ca 1900.

Sumatra (Sum) 39 BER-1; BRE-1; DRE-5; LEP-4;PAR-3; ZUR-25

The specimens in Zurich are designated ‘‘Battak’’; specific locations within theisland of Sumatra are not known.

Java (Jav) 50 BER-1; BLU-8; CHA-9; DRE-1;LEP-24; PAR-7

Crania were collected from several different localities in Java.

Borneo (Bor) 34 BER-2; BRE-2; DRE-6; FRE-4;LEP-8; PAR-12

A great many of the specimens are indicated as representing Dayak tribes, somehave elaborate decorations.

Sulawesi (Slw) 41 BAS-7; BER-10; DRE-4; FRE-7;LEP-5; PAR-8

An exact location is known for many of these specimens.

Lesser Sunda Island (Lsn) 61 BAS-5; BER-6; BLU-2;CHA-1; DRE-17; LEP-1;PAR-6; ZUR-7

Crania from Bali (13), Flores (9), Sumba (1), Lomblem (2), Alor (2), Timor (11),Wetar (2), Leti (4), Barbar (1), Tanimbar (13), Kai (2) and Aru (1) Islands of theLesser Sunda Islands.

Southern Molucca Islands(Sml)

65 FRE-47; DRE-17 Crania are from Ceram (48) and Buru (17) Islands of the Southern Molucca Islands.

Sulu (Sul) 38 LEP-1; PAR-37 The specimens in Paris were collected by Montano-Rey ca 1900.Philippines (Phl) 28 BER-9; DRE-19 Most specimens are from Luzon Island.

a Names of the cranial series and abbreviations used in the tables and figures that accompany this paper.b BAC, Bachuc Village, Angiang Province, Vietnam; BAS, Naturhistorisches Museum, Basel; BER, Museum fur Naturkunde, Berlin; BLU, Anatomisches Institut, Universitat

Gottingen, Gottingen; BRE, Uber-see Museum, Bremen; CHA, Anatomisches Institut der Chairte, Humboldt Universitat, Berlin; CHB, Chiba University School of Medicine, Chiba;CHE, Department of Anatomy, Chengdu College of Traditional Chinese Medicine, Chengdu, PRC; DRE, Museum fur Volkerkunde, Dresden; FRE, Institut fur Humangenetik u.Anthropologie, Universitat Freiburg; GYO, Gyokusendo Cave, Okinawa Island; HCM, Faculty of Medicine, Ho Chi Minh City, Vietnam; HKU, University of Hong Kong, Hong Kong;KAN, Kanegusuku Storage Room, Board of Education Cultural Division, Kanegusuku, Okinawa; KYO, Lab of Physical Anthropology, Faculty of Science, Kyoto University, Kyoto;KYU, Department of Anatomy, Faculty of Medicine, Kyushu University, Fukuoka; LEP, Anatomisches Institut, Karl Marx Universitat, Leipzig; NSM, National Science Museum,Tokyo; PAR, Musee de l’Homme, Paris; RYU, University of the Ryukyus, Naha, Okinawa Island; SAP, Department of Anatomy, Sapporo Medical College, Sapporo; SEN,Department of Anatomy, School of Medicine, Tohoku University, Sendai; SHA, Institute of Anthropology, College of Life Sciences, Fudan University, Shanghai; SIM, NationalMuseum of Natural History, Smithsonian Institution, Washington, D.C.; SIR, Department of Anatomy, Siriraj Hospital, Bangkok; TKM, Medical Museum, University Museum,University of Tokyo; TKO, University Museum, University of Tokyo, Tokyo; TPE, Academia Sinica, Nankang, Taipei; ZUR, Anthropologisches Institut, Universitat Zurich, Zurich.

Fig. 1. Map of Eastern Asia showing the location of the 38 male cranial series used in this analysis.

M. Pietrusewsky / Quaternary International 211 (2010) 42–54 45

M. Pietrusewsky / Quaternary International 211 (2010) 42–5446

are those from China (Anyang) and six which span the late Jomon toEdo Periods in Japan.

Anyang is a Bronze-Age site located in Henan Province, northernChina. The Anyang crania are from ‘sacrificial pits’ found in asso-ciation with the imperial tombs of the Shang Dynasty (11th centuryB.C.) (Li, 1977; Howells, 1983). The Anyang specimens were exca-vated prior to World War II and are currently curated at AcademiaSinica, Taipei, where they were examined by the author in 1983 and1991. Measurements from a random selection of 56 adult malecrania are used in this analysis.

The early cranial series from Japan include Late to Latest Jomon(3500–2000 BP), Yayoi (300 B.C.–300 A.D.), Kofun (4th–6th centu-ries), two medieval cranial series (Kamakura and Marunouchi), andan Edo Period (17th–19th centuries A.D.) cranial series. In additionto cranial series representing earlier and more modern Japanese,cranial series representing Ainu and Ryukyu Islanders are includedin this analysis.

2.2. Cranial measurements

Two multivariate statistical procedures, stepwise discriminantfunction analysis and Mahalanobis’ generalized distance statistic(Mahalanobis, 1936), are applied to a total of 28 standardmeasurements (see Table 2 for a description of the exactmeasurements) recorded in male crania. A more detailed discus-sion of these methods is provided by Pietrusewsky (2008).

2.3. Multivariate statistical procedures

2.3.1. Stepwise discriminant function (canonical) analysisThe major purpose of discriminant function (or canonical)

analysis is to maximize differences between groups which ismathematically achieved by producing a new set of transformed

Table 2Summary ranking of cranial measurements according to F-values received in thefinal step of discriminant function analysis (38 male groups, 28 measurements).

Step No. Measurementa,b F-value d.f.B/d.f.Wc pd

1 Nasio-occipital length (M-1d) 17.792 37/1716 *2 Nasal height (H-NLH) 15.335 37/1715 *3 Nasal breadth (M-54) 11.847 37/1714 *4 Minimum cranial breadth (M-14) 11.610 37/1713 *5 Bijugal breadth [M-45(1)] 9.207 37/1712 *6 Basion–bregma height (M-17) 9.044 37/1711 *7 Biorbital breadth (H-EKB) 8.107 37/1710 *8 Bifrontal breadth (M-43) 10.334 37/1709 *9 Alveolar length (M-60) 7.438 37/1708 *10 Maximum cranial breadth (M-8) 7.380 37/1707 *11 Malar length inferior (H-IML) 7.146 37/1706 *12 Cheek height [H-WMH] 6.748 37/1705 *13 Basion–nasion length (M-5) 6.592 37/1704 *14 Orbital height left (M-52) 6.250 37/1703 *15 Biauricular breadth (M-11) 4.801 37/1702 *16 Mastoid height (H-MDL) 4.200 37/1701 *17 Nasion–bregma chord (M-29) 3.741 37/1700 *18 Lambda–opisthion chord (M-31) 3.722 37/1699 *19 Bistephanic breadth (H-STB) 3.512 37/1698 *20 Maximum frontal breadth (M-10) 5.831 37/1697 *21 Maximum cranial length (M-1) 3.387 37/1696 *22 Orbital height left (M-52) 3.344 37/1695 *23 Minimum frontal breadth (M-9) 3.086 37/1694 *24 Bregma–lambda chord (M-30) 3.125 37/1693 *25 Alveolar breadth (M-61) 3.013 37/1692 *26 Bimaxillary breadth (M-46) 2.251 37/1691 *27 Mastoid width (H-MDB) 1.987 37/1690 *28 Biasterionic breadth (M-12) 1.824 37/1689 *

a M¼Martin and Saller (1957).b H¼Howells (1973).c d.f.B/d.f.W ¼ degrees of freedom between/degrees of freedom within.d * Probability p� 0.01; n.s.¼ not significant.

variables (discriminant functions or canonical variates) betweenwhich correlation is removed (Tatsuoka, 1971). These transformedvalues are extremely useful in relating human groups to each otheras well as revealing much about the underlying differences incranial form. Typically, the first few functions, or canonical variates,account for most of the variation among groups. In this analysis, theoriginal measurements were selected in a stepwise manner suchthat, at each step, the measurement that adds most to the separa-tion of the groups was the one entered into the discriminantfunction in advance of the others (Dixon, 1992:363). This procedureallows identification of those variables that are most responsiblefor the observed differentiation between individuals of the variousgroups. Interpretations of discriminant functions and the patternsof group separation are based on an inspection of standardizedcanonical coefficient values.

At the end of the stepping process, each individual specimen isclassified into one of the original groups based on the discriminantscores it receives through the calculation of posterior (regularclassification) and/or typicality (jackknifed classification) proba-bilities (Van Vark and Schaafsma, 1992:244–255). Jackknifed clas-sification represents a common cross-validation procedure inmultiple discriminant analysis, where cases are classified withoutusing misclassified individuals in computing the classificationfunction. The ‘correct’ and ‘incorrect’ classification results providea general guide for assessing the homogeneity or heterogeneity ofthe original series. Another useful feature of this procedure is thatgroup means can be plotted on the first few canonical variates, thusallowing visualization of intergroup relationships. The computerprogram, BMDP-7M (Dixon, 1992; Dixon and Brown, 1979) is usedto perform the stepwise discriminant function analysis. The two-dimensional and three-dimensional canonical plots are made usingthe SYGRAPH module of the SYSTAT computer programs (Wilkin-son, 1992).

2.3.2. Mahalanobis’ generalized distanceMahalanobis’ generalized distance, or the sum of squared

differences, provides a single quantitative measure of dissimilarity(distance) between groups using several variables while removingthe correlation between the variables (Mahalanobis, 1936). Thesignificance of these distances is determined using the method ofRao (1952:245), a procedure recommended by Buranarugsa andLeach (1993:17).

The diagrams of relationship, or dendrograms, were constructedby applying the average linkage within group clustering algorithm,or Unweighted Pair Group Method Algorithm(UPGMA) (Sneath andSokal, 1973) to the distances. This latter algorithm combines clus-ters so that the average distance among all cases in the resultingcluster is as small as possible and the distance between two clustersis taken to be the average among all possible pairs of cases in thecluster. The NTSYS-pc computer software program was used toconstruct the dendrogram (Rohlf, 1993).

3. Results

The results of applying stepwise discriminant function analysisand Mahalanobis’ generalized distance to 28 cranial measurementsrecorded in 1753 male crania, the largest number of measurementscomparable to all 38 cranial series, are presented.

3.1. Stepwise discriminant function analysis

A summary of the measurements (Table 2), ranked according tothe F-values [tests of equality of group means using classical one-way analysis of variance] received in the final step of discriminantfunction analysis provides an indication of the discriminatory

M. Pietrusewsky / Quaternary International 211 (2010) 42–54 47

power of the original variables. Among the variables that areranked the highest (i.e., they contribute the most to the discrimi-nation produced) in this analysis are the nasio-occipital length(M-1d), nasal height (H-NLH), nasal breadth (M-54), and minimumcranial breadth (M-14).

Eigenvalues, which represent the amount of variance, accountedfor by each function or variate, expressed as the percentage of totaldispersion, and level of significance (Rao, 1952:323) for the 28canonical variates is presented in Table 3. The eigenvalues providean indication of the proportion of dispersion accounted for by eachcanonical variate. In this analysis, the first four canonical variatesaccount for 66.0% of the total variation. All the eigenvalues aresignificant at the 1% level, indicating significant heterogeneity forthese canonical variates.

Canonical coefficients, those values by which an individual’smeasurements may be multiplied to obtain its score, for 28measurements, for the first three canonical variates are given inTable 4. Bifrontal breadth (M-43), biorbital breadth (H-EKB), nasalbreadth (H-NLH), and nasio-occipital length (M-1d) (those vari-ables with the highest coefficients regardless of sign) are the mostimportant variables in producing group separation in the firstcanonical variate. This first variate emphasizes breadths of thefrontal, orbital, and nasal regions of the cranium. Orbital height(M-52), cheek height (H-WMH), maximum frontal breadth (M-10)and basion–nasion breadth (M-5) are most responsible for groupseparation produced in the second canonical variate. Bifrontalbreadth (M-43), biorbital breadth (H-EKB), orbital breadth (M-51a)and maximum cranial length (M-1) are primarily responsible forthe discrimination produced in the third canonical variate.

Table 3Eigenvalues, percentage of total dispersions, cumulative percentage of dispersion,and level of significance for 28 canonical variates resulting from stepwise discrim-inant function analysis (38 male groups, 28 measurements).

Canonicalvariate

Eigenvalue % Dispersion Cumulative %dispersion

d.f.a pb

1 1.20084 26.4 26.4 64 *2 0.78632 17.2 43.6 62 *3 0.63974 14.1 57.7 60 *4 0.37895 8.3 66.0 58 *5 0.30527 6.7 72.7 56 *6 0.23188 5.1 77.8 54 *7 0.14688 3.3 81.1 52 *8 0.11856 2.6 83.7 50 *9 0.11142 2.4 86.1 48 *10 0.09717 2.1 88.2 46 *11 0.07926 1.8 90.0 44 *12 0.06725 1.5 91.5 42 *13 0.06281 1.3 92.8 40 *14 0.05433 1.2 94.0 38 *15 0.04579 1.0 95.0 36 *16 0.04257 1.0 96.0 34 *17 0.03850 0.8 96.8 32 *18 0.03481 0.8 97.6 30 *19 0.02422 0.5 98.1 28 *20 0.02061 0.5 98.6 26 *21 0.01582 0.3 98.9 24 *22 0.01087 0.3 99.2 22 *23 0.00949 0.2 99.4 20 *24 0.00869 0.2 99.6 18 *25 0.00861 0.1 99.7 16 *26 0.00551 0.2 99.9 14 *27 0.00505 0.0 99.9 12 *28 0.00133 0.1 100.0 10 *

a d.f.¼ degrees of freedom¼ (pþ q� 2), (pþ q� 4)..b *Probability p� 0.01 when eigenvalues are tested for significance according to

criterion [N�½ (pþ q)] loge (lþ 1), where N¼ total number of crania, p¼ numberof variables, q¼ number of groups, l¼ eigenvalue, all of which are distributedapproximately as chi-square (Rao, 1952:373).

A summary of the jackknifed classification results (Table 5)indicate that Mongolia, Southern Moluccas, Atayal, Cambodia-Laos,Chengdu, Ainu, Amami Islands, and Jomon are among the serieswith the best classification results (i.e. more than 45% of the casesare correctly assigned to their original groups). The poorest jack-knifed classification results (less than 12% of the cases correctlyclassified to the original group) are found for Sulawesi, Sakishima,Hangzhou, Edo, and Marunouchi series.

The most frequent (first 10) misclassifications for each of the 39groups are given in Table 6. The most frequent misclassifications arefound among the modern cranial series from China and north-eastern Asia. Typically, Shanghai, Hangzhou, and Nanjing arereclassified among these same Chinese cranial series indicatinga great deal of overlap among the eastern Chinese series. Several ofthe Chinese specimens are reclassified to one of the Japanese seriesand occasionally, to one of the Southeast Asian series. Hainan IslandChinese are frequently reclassified as one of the Southeast Asianseries. Notably, the Taiwan and Hainan Island Chinese are reclas-sified as the Bronze-Age series from Anyang. While nearly 40% ofthe Anyang specimens are correctly classified, others are mis-classified to Korean, Chinese, Ryukyu Island, or Taiwan Aboriginalseries. Ten of Manchurian crania are reclassified to one of theChinese series and nine are reclassified as Japanese. In these results,many of the Korean crania are reclassified as Hainan Island, Taiwan,Anyang, Tohoku, or Kyushu. Most of the Mongolian specimens arereclassified to their original group.

Examining some of the Japanese series, many of the Yayoispecimens are misclassified to other Japanese series that tempo-rally post-date the Yayoi Period. Likewise, a substantial number ofthe Kofun and medieval Kamakura crania are misclassified to one ofthe Ryukyu Island series or other modern Japanese cranial series.

Further inspection of the Ryukyu Island series (Amami, Okinawa,and Sakishima) demonstrates that many of these crania are reclas-sified to one of the Ryukyu Island cranial series or to one of theJapanese (Kamakura, Yayoi) series. The classification results for

Table 4Canonical coefficients of 28 cranial measurements for the first three canonicalvariates that result from stepwise discriminant function analysis of 38 male groups.

Canonicalvariate 1

Canonicalvariate 2

Canonicalvariate 3

Maximum cranial length, M-1 �0.06380 0.00397 �0.13038Nasio-occipital length, M-1d 0.19550 0.05226 0.03524Basion–nasion length, M-5 0.06464 �0.08309 �0.04394Basion–bregma height, M-17 0.00108 �0.01375 0.07186Maximum cranial breadth, M-8 �0.01839 0.00379 �0.03237Maximum frontal breadth, M-10 0.00658 0.11299 �0.02623Minimum frontal breadth, M-9 �0.04383 �0.05625 �0.06549Bistephanic breadth, H-STB 0.00890 �0.08166 0.00902Biauricular breadth, M-11b 0.03037 0.06017 �0.01840Minimum cranial breadth, M-14 0.06326 0.05047 0.07929Biasterionic breadth, M-12 �0.00745 0.00815 �0.03563Nasal height, H-NLH �0.10560 0.05579 0.04337Nasal breadth, M-54 �0.22983 0.07377 �0.00882Orbital height left, M-52 �0.03445 0.22709 �0.03625Orbital breadth left, M-51a �0.01822 0.01934 �0.19102Bijugal breadth, M-45(1) 0.04272 �0.04438 �0.06723Alveolar length, M-60 �0.06764 �0.07726 �0.08318Alveolar breadth, M-61 �0.01895 �0.01339 0.02816Mastoid height, H-MDL �0.01832 �0.00842 0.02493Mastoid width, H-MDB 0.02172 �0.00683 �0.00660Bimaxillary breadth, M-46 �0.00168 0.00693 �0.00249Bifrontal breadth, M-43 0.25212 0.02823 0.31428Biorbital breadth, H-EKB �0.25124 �0.07014 �0.19965Malar length inferior, H-IML �0.12438 0.04196 0.02469Cheek height, H-WMH 0.04362 0.19139 0.02226Nasion–bregma chord, M-29 �0.06716 �0.04277 0.03802Bregma–lambda chord, M-30 �0.01662 0.00558 0.03485Lambda–opisthion chord, M-31 �0.02148 �0.04094 0.03951

Table 5Classification results (regular and jackknifed) arranged by groups with the best tothe poorest showing percentage of correctly assigned cases.

Regular classification results Jackknifed classification results

Group % Group %

Mongolia 82.0 Mongolia 76.0Burma 75.0 Southern Molucca Islands 63.1Southern Molucca Islands 70.8 Atayal 55.6Cambodia-Laos 65.0 Cambodia-Laos 55.0Atayal 63.9 Chengdu 47.2Amami Islands 61.3 Ainu 46.0Jomon 60.8 Amami Islands 45.2Bachuc Village 60.8 Jomon 45.1Ainu 58.0 Bachuc Village 41.2Chengdu 52.8 Anyang 39.3Okinawa Islands 52.8 Manchuria 38.0Hong Kong 50.0 Okinawa Islands 37.7Sulu 50.0 Lesser Sunda Islands 34.4Taiwan Chinese 48.9 Thailand 34.0Anyang 48.2 Hong Kong 32.0Thailand 46.0 Taiwan Chinese 29.8Manchuria 44.0 Sulu 28.9Lesser Sunda Islands 42.6 Vietnam 24.5Sakishima Group 41.7 Java 24.0Kanto Japanese 40.0 Kyushu Japanese 23.5Java 38.0 Hainan Island 23.4Kyushu Japanese 37.3 Shanghai 22.0Philippines 35.7 Kanto Japanese 22.0Vietnam 34.7 Tohoku Japanese 20.8Kamakura 34.6 Burma 18.8Hainan Island 34.0 Sumatra 17.9Sumatra 33.3 Philippines 17.9Kofun 32.3 Kofun 17.7Nanjing 30.6 Borneo 17.6Yayoi 30.6 Kamakura 17.3Borneo 29.4 Korea 15.6Tohoku Japanese 28.3 Yayoi 14.5Korea 28.1 Nanjing 12.2Shanghai 28.0 Sulawesi 9.8Sulawesi 24.4 Sakishima Group 8.3Hangzhou 22.0 Hangzhou 8.0Marunouchi 14.8 Edo 5.5Edo 9.1 Marunouchi 3.7

M. Pietrusewsky / Quaternary International 211 (2010) 42–5448

Jomon and Ainu indicate a great deal of reciprocity between thesecranial series. Of the Southeast Asian series, the Southern MoluccaIsland series has the best classification results with a majority of thecases being correctly classified to this group. Misclassificationsamong the remaining Southeast Asian series are generally to otherSoutheast Asian series. Several of the Vietnam and Thailand speci-mens are misclassified to Northern or Northeast Asian series.

When 38 group means are plotted on the first two canonicalvariates (Fig. 2) three separate clusters emerge. The cranial seriesfrom mainland and island Southeast Asia form a separate constel-lation in the lower left quadrant of this diagram. Four modernChinese cranial series, Chengdu, Shanghai, Nanjing, and Hangzhou,seen in the upper portion of this diagram, anchor another group towhich Mongolia, Manchuria, and Hong Kong are marginallyattracted. The last major cluster evident in Fig. 1 includes Taiwanand Hainan Island Chinese cranial series, Anyang, Korea, and all thecranial series from Japan, including the Ryukyu Islands. HainanIsland, Anyang, Taiwan, and Korea occupy more marginal positionsin this constellation. The Jomon and Ainu series form an isolatedgroup. The cranial series that are somewhat intermediate betweenthe clusters containing Northeast Asian (Chinese) and Japanesecranial series include Manchuria, Yayoi and Kofun.

3.2. Mahalanobis’ generalized distance

Applying the UPGMA clustering algorithm to the distances for38 groups results in the dendrogram presented in Fig. 3. Two major

divisions are evident in this dendrogram, one contains all thecranial series representing Southeast Asia and the second includesall the series from Eastern and Northern Asia. Within the SoutheastAsian division there is good separation between mainland andisland Southeast Asian cranial series. In the second major division,Shanghai, Hangzhou, Nanjing, Chengdu, Hong Kong, and Man-churia unite in a separate branch. Taiwan, Hainan, Korea, andAnyang form another branch in this division, one that unites withthe Japanese and Ryukyu Island cranial series. Also evident in thisdivision is a sub-branch that contains the more modern Japanese(Kanto, Kyushu, and Tohoku), Edo, and Marunouchi cranial series,which then unites with another cluster that includes Kamakura,Kofun, Yayoi, and the three Ryukyu Island cranial series. The Atayaldo not show a convincingly close relationship with any of the otherseries compared in this analysis. The Ainu and Jomon series falloutside the second major branch that contains all the cranial seriesfrom Northern and Eastern Asia. The most isolated cranial series inthis analysis is Mongolia.

Inspection of the 10 smallest distances for each of the 38 groups(Table 7) gives additional information on biological relatedness thatis not readily apparent in the diagrams that result from the appli-cation of clustering algorithms. Using distance size as a measure ofcloseness, the groups closest to the Southeast Asian series generallyinclude other Southeast Asian series, but not exclusively so. Forexample, Korea and Hainan Island appear among the groups closestto several of the Southeast Asian series, including Sumatra, Sula-wesi, Philippines, Vietnam, Bachuc, Thailand, and Burma. Likewise,among the groups found to be closest to Vietnam and Thailand areseveral that are not from Southeast Asia. Finally, Vietnam appearsamong the groups closest to most of the Chinese cranial series.

Further inspection of these results indicates that the groupsclosest to Manchuria include six Japanese and three Chinese series.Similarly, five modern, or relatively modern, cranial series fromJapan (Edo, Kyushu, Tohoku, Marunouchi, and Kanto) are among thegroups closest to Korea. The groups closest to Anyang includeHainan, Korea, and Taiwan followed by six cranial series from Japan.Korea, Taiwan and Anyang are among the series closest to HainanIsland. Hainan, Anyang, and Korea are the groups most closelyrelated toTaiwan. As was the case for Taiwan and Hainan, the groupsclosest to the Taiwan Aboriginal group, Atayal, are from Korea, Japan,Hainan, and Taiwan. Three cranial series from Japan (Yayoi, Kofun,and Marunouchi) and four from China (Hainan, Chengdu, Shanghai,and Nanjing) are among the groups closest to Mongolia.

The cranial series closest to Yayoi include Kofun and twomedieval series from Japan (Marunouchi and Kamakura) followedby two Ryukyu Island series. The groups closest to Kofun, includeYayoi and two medieval series from Japan followed by the threeRyukyu Island series. Among the groups that are most often closestto the Ryukyu Island series are Kamakura, Yayoi, Kofun and theMarunouchi. Further inspection of these distances reveals a Jomon-Ainu connection. Among the groups closest to the Ainu and Jomonseries are those from Japan such as Yayoi, Kofun, Kamakura andMarunouchi. None of the Chinese or mainland Northeast Asianseries is among the groups closest to either Jomon or Ainu.

4. Discussion of results

4.1. Origins of Southern and Northern Asians

Because the present craniometric study does not include theearlier fossil record for Eastern Asia from the time of Homo erectusto late Pleistocene, the results of the present analysis cannotdirectly test models of continuity (Etler, 1996) or discontinuity(Brown, 1998, 2001) to explain the origins of the modern peoples ofEastern Asia. However, the results of the present analysis provide

Table 6Some of the jackknifed classification results obtained from stepwise discriminant function analysis showing the cases reclassified at the end of the stepping process (numbersin parentheses represent the number of crania originally assigned to each group). See Table 1 for explanation of abbreviations.

Shanghai (50) Hong Kong (50) Chengdu (53) Hangzhou (50) Nanjing (49) TaiwanChinese (47)

Hainan Island (47) Atayal (36)

HAN 12 HK 16 CHE 25 SHA 11 CHE 6 TAI 14 HAI 11 ATY 20SHA 11 HAN 5 NAN 8 NAN 5 HAN 6 ANY 5 ANY 5 KOR 2NAN 4 NAN 3 SHA 4 HK 4 NAN 6 HAI 4 TAI 4 HK 1KAN 4 MAR 3 HK 3 CHE 4 MAN 5 TOH 4 THI 4 HAI 1THI 4 SHA 2 HAN 2 HAN 4 SHA 4 BAC 3 BUR 3 EDO 1MOG 3 TAI 2 PHL 2 MOG 3 HK 4 CHE 2 KOR 2 KAM 1HK 2 HAI 2 VTN 2 SUM 3 ANY 3 MAN 2 KYU 2 TOH 1CHE 2 MAN 2 HAI 1 MAN 2 KOR 3 KAN 2 JAV 2 KYU 1AIN 2 AMA 2 MAN 1 ANY 2 YAY 2 KAM 2 SLW 2 AIN 1JAV 2 THI 2 MOG 1 BOR 2 VTN 2 PHL 2 VTN 2 OKI 1

Manchuria (50) Anyang (56) Mongolia (50) Korea (32) KantoJapanese (50)

Edo (55) Kamakura (52) Kofun (62)

MAN 19 ANY 22 MON 38 HAI 5 KAN 11 KYU 5 KAM 9 KOF 11CHE 4 KOR 4 SHA 2 KOR 5 MAR 4 ATY 4 AMA 6 KAM 7EDO 3 NAN 3 CHE 1 TAI 2 TOH 3 KAN 4 KYU 5 AMA 6HK 2 TAI 3 HAN 1 ANY 2 KYU 3 TOH 4 SAK 5 JOM 5TAI 2 HAI 3 NAN 1 TOH 2 JOM 3 MAN 3 KOF 4 SAK 4TOH 2 AMA 3 HAI 1 KYU 2 SHA 2 EDO 3 TOH 3 HK 3SAK 2 HK 2 YAY 1 CHE 1 HK 2 YAY 3 LSN 3 YAY 3MAR 2 ATY 2 AMA 1 NAN 1 HAN 2 SAK 3 ANY 2 TOH 3NAN 1 KYU 2 MAR 1 ATY 1 ATY 2 MAR 3 KAN 2 OKI 3HAI 1 SAK 2 JAV 1 MAN 1 OKI 2 CHE 2 EDO 2 MAR 3

Yayoi (62) TohokuJapanese (53)

KyushuJapanese (51)

Ainu (50) AmamiIslands (31)

Jomon (51) OkinawaIsland (53)

SakishimaGroup (24)

YAY 9 TOH 11 KYU 12 AIN 23 AMA 14 JOM 23 OKI 20 OKI 5MAR 8 KYU 5 KAN 6 KAN 3 KAM 3 AIN 6 AMA 5 MAN 2KOF 7 AIN 5 ATY 3 TOH 3 YAY 2 OKI 4 VTN 4 KAM 2KAM 5 KOR 4 KOR 3 JOM 3 OKI 2 KAM 3 TAI 3 TOH 2KAN 3 ATY 3 EDO 3 MAR 3 SHA 1 KAN 2 HAI 3 AMA 2AMA 3 MAN 3 TOH 3 KAM 2 TAI 1 KOF 2 KAM 2 SAK 2THI 3 KAM 3 HAI 2 KOF 2 ATY 1 YAY 2 YAY 2 SHA 1CHE 2 KAN 2 KAM 2 SUM 2 ANY 1 MAR 2 KYU 2 HAI 1NAN 2 EDO 2 KOF 2 HK 1 KYU 1 MOG 1 JOM 2 KAN 1MOG 2 KOF 2 JOM 2 MAN 1 AIN 1 TOH 1 SAK 2 KYU 1

Marunouchi (27) Burma (16) Sumatra (39) Java (50) Borneo (34) Sulawesi (41) Lesser SundaIslands (61)

SouthernMoluccaIslands (65)

YAY 4 BUR 3 SUM 7 JAV 12 BOR 6 CML 6 LSN 21 SML 41ATY 3 BAC 2 BUR 3 CML 6 VTN 4 SUM 4 BOR 5 LSN 6TOH 3 CML 2 BOR 3 SLW 4 SUM 3 SLW 4 SLW 5 SUL 3KYU 2 HAN 1 SML 3 SUL 4 LSN 3 SUL 3 SUM 4 ATY 2OKI 2 HAI 1 PHL 3 LSN 3 SUL 3 THI 3 BUR 3 SUM 2SAK 2 TOH 1 CHE 2 BAC 3 KOF 2 SHA 2 CML 3 SLW 2SHA 1 AIN 1 MOG 2 THI 3 JAV 2 ATY 2 HK 2 PHL 2HK 1 JOM 1 EDO 2 HK 1 SLW 2 JAV 2 KAM 2 VTN 2NAN 1 SUM 1 JAV 2 CHE 1 HAN 1 BOR 2 SAK 2 SAK 1HAI 1 SLW 1 LSN 2 HAN 1 KYU 1 LSN 2 PHL 2 JAV 1

Sulu (38) Philippines (28) Vietnam (49) BachucVillage (51)

Cambodia-Laos (40)

Thailand (50)

SUL 11 PHL 5 VTN 12 BAC 21 CML 22 THI 17CML 6 VTN 5 PHL 4 VTN 4 LSN 3 BAC 5BOR 4 BAC 3 ATY 3 SUL 3 SUL 3 KOR 3SLW 4 SHA 2 OKI 3 THI 3 SML 2 HK 2JAV 3 HK 2 LSN 3 HK 2 BAC 2 TAI 2LSN 2 SLW 2 BAC 3 TAI 2 NAN 1 KAN 2HAN 1 CML 2 SHA 2 KYU 2 KOR 1 KOF 2KOR 1 HAN 1 HAN 2 BUR 2 JOM 1 JAV 2YAY 1 HAI 1 TAI 2 JAV 2 BUR 1 SML 2AIN 1 ATY 1 KOR 2 BOR 2 SUM 1 SHA 1

M. Pietrusewsky / Quaternary International 211 (2010) 42–54 49

background for examining some of the current models that attemptto explain the population history of these regions. As discussed inmore detail earlier, using primarily archaeological and linguisticevidence, Bellwood (2005) and others have argued strongly fora population displacement to account for the people who nowinhabit Southeast Asia. Specifically, Bellwood has maintained thatthe indigenous inhabitants of mainland and island Southeast Asiawere replaced by an immigrant group of people of a more northern

origin, or, to use his terminology, ‘‘Australoids’’ were displaced by‘‘Mongoloids’’. Such a scenario should, at least in theory, result inthe presence of a somewhat hybridized population living in thisregion. An alternative model, the so-called continuity model,argues that the present day inhabitants of Southeast Asia evolvedwithin this region from the Late Pleistocene onward.

The present craniometric comparisons, especially the diagramof relationship presented in Fig. 3, demonstrate a clear separation

Fig. 2. Plot of 38 group means on the first two canonical variates using 28 cranialmeasurements [see Table 1 for explanation of group abbreviations].

M. Pietrusewsky / Quaternary International 211 (2010) 42–5450

between cranial series in Northern and Eastern Asia from those inSoutheast Asia. The connecting point between these two divisionsis a group comprised of Hainan, Taiwan, Anyang, and Korean cranialseries. This striking separation between North/East Asian cranialseries provides more support for models of long-term in situ

Fig. 3. Diagram of relationship (dendrogram) based on a cluster analysis (UPGMA) of Magroups.

evolution in these two regions rather than those that advocatereplacement and discontinuity to account for the modern peoplesof Eastern Asia. Similar conclusions have been reached by Turner(1987, 1990) using dental morphology and Hanihara (1993) andPietrusewsky (2006) using craniometric data.

4.2. Origins of the modern Chinese and the peoples of Eastern Asia

More than a quarter of a century ago, Howells (1983) concludedthat modern Chinese were already in place in northern China ora source not too distant from there in Northeast Asia perhaps asearly or earlier than 5000 B.C., a people who share a common originwith the modern inhabitants of the Korean Peninsula and Japanesearchipelago. Others (e.g., Bowles, 1984) have suggested that theorigins of the Han Chinese, the most populous of the modernEastern Asians, has been linked invariably to the fertile middle andlower basins of the Yellow (Huanghe) and Yangzi rivers and severalsmaller river basins in present day China.

Again, while the results of the multivariate analysis presentedhere cannot speak directly of the origins of the Chinese and thepeoples of Eastern Asia, the patterns of relationship demonstratedin these results provide context for interpreting various originscenarios. In these results, the cranial series from China, Korea, andJapan form a distinct constellation well removed from the cranialseries representing Southeast Asia, a finding that reinforcesHowells’ (1986) earlier assertion that the Chinese, Koreans, andJapanese represent historically expanded populations arising froma common northern hearth. Closer inspection of the assemblagecontaining the East Asian series in the results of present multivar-iate analyses of craniometric data further indicates that the cranial

halanobis’ generalized distances using 28 cranial measurements recorded in 38 male

Table 7The smallest Mahalanobis’ distances for 38 male cranial groups using 28 measure-ments. All distances are significant p� 0.01 unless indicated otherwise.a

Sumatra Java Borneo SulawesiBOR 2.686** SLW 2.452** SUM 2.686** CML 2.444**LSN 2.965* CML 3.596* LSN 3.033* JAV 2.452**SLW 3.269** LSN 3.673 SLW 3.272** LSN 2.919*JAV 4.453 SUL 4.426 SUL 4.641* SUM 3.269**VTN 4.860 SUM 4.453 JAV 4.866 BOR 3.272**HAN 4.901 THI 4.461 VTN 5.061 SUL 3.561**EDO 5.031 BOR 4.866 CML 5.560 THI 4.605PHL 5.348* PHL 5.754 PHL 5.864** PHL 4.642**SHA 5.623 VTN 6.026 SML 6.110 VTN 5.914SUL 5.696 BAC 6.502 EDO 7.506 KOR 6.456

Lesser SundaIslands

SouthernMolucca Islands

Sulu Philippines

SLW 2.919* LSN 5.124 CML 3.401** VTN 2.175**SUM 2.965* BOR 6.110 SLW 3.561** LSN 4.594BOR 3.033* SUM 6.213 JAV 4.426 SLW 4.642**JAV 3.673 SLW 6.664 BOR 4.641* THI 4.849*PHL 4.594 SUL 7.039 SUM 5.696 SUM 5.348*VTN 4.905 PHL 8.154 LSN 5.930 KOR 5.641*SML 5.124 CML 9.480 BUR 6.717** HAI 5.746EDO 5.763 VTN 10.091 SML 7.039 JAV 5.754SUL 5.930 JAV 10.126 BAC 7.761 BOR 5.864**CML 6.198 EDO 11.210 PHL 7.958 HK 5.877

Vietnam Buchuc Village Cambodia–Laos ThailandPHL 2.175** THI 3.338 SLW 2.444** BAC 3.338THI 4.318 HAI 5.095 SUL 3.401** VTN 4.318KOR 4.364* VTN 5.226 JAV 3.596* JAV 4.461SUM 4.860 PHL 5.910 BOR 5.560 SLW 4.605HAI 4.893 JAV 6.502 THI 6.176 HAI 4.759LSN 4.905 SLW 6.652 LSN 6.198 PHL 4.849*EDO 4.950 CML 7.257 SUM 6.374 KOR 4.938BOR 5.061 KOR 7.302 BAC 7.257 HK 5.456HK 5.222 BUR 7.563* BUR 7.401** CML 6.176BAC 5.226 SUL 7.761 VTN 8.203 KAN 6.224

Burma Shanghai Hong Kong ChengduSUL 6.717** HAN 0.593** HAN 3.708 NAN 2.639*SUM 6.962** NAN 2.339** NAN 4.116 HAN 3.906CML 7.401** CHE 4.034 SHA 4.170 SHA 4.034SLW 7.453** HK 4.170 VTN 5.222 MAN 6.118BAC 7.563* SUM 5.623 MAR 5.356 YAY 7.118JAV 7.610* MAR 5.787 THI 5.456 VTN 7.187THI 8.182* EDO 6.137 PHL 5.877 KOR 7.320HAI 8.810* THI 6.228 EDO 6.097 MAR 7.606BOR 9.224** KOR 6.472 KOR 6.302 EDO 7.689EDO 9.511 MAN 6.506 MAN 6.446 HK 7.827

Hangzhou Nanjing Taiwan Chinese Hainan IslandSHA 0.593** HAN 1.637** HAI 3.127* KOR 2.573**NAN 1.637** SHA 2.339** ANY 3.769 TAI 3.127*HK 3.708 CHE 2.639* KOR 4.042* ANY 3.372CHE 3.906 MAN 3.900 OKI 5.760 EDO 4.416SUM 4.901 HK 4.116 VTN 6.312 THI 4.759MAR 5.579 KOR 4.759 MAN 6.398 VTN 4.893VTN 5.654 MAR 4.804* SAK 6.426 BAC 5.095EDO 5.870 EDO 5.035 EDO 6.555 OKI 5.107KOR 5.981 YAY 5.414 AMA 7.084 KYU 5.349YAY 6.021 VTN 6.124 KYU 7.341 SAK 5.508*

Atayal Manchuria Anyang MongoliaKOR 5.828* EDO 3.574 HAI 3.372 HAN 10.079EDO 5.952 NAN 3.900 KOR 3.547* YAY 10.155TOH 6.217 KOR 4.041* TAI 3.769 CHE 10.230HAI 6.380 MAR 4.961* AMA 3.939* SHA 10.572KYU 6.457 TOH 5.170 KAM 4.869 NAN 11.135SAK 7.334* YAY 5.757 EDO 5.302 SUM 13.023MAR 7.401* HAN 6.050 KYU 5.522 KOF 13.251TAI 8.031 CHE 6.118 SAK 5.694 MAN 14.584KAM 8.122 KYU 6.123 MAR 5.725 MAR 14.866VTN 8.132 SAK 6.201 KOF 5.946 SLW 14.875

Korea Kanto Japanese Edo KamakuraEDO 2.242** KYU 2.696* KYU 1.423** KOF 2.715HAI 2.573** EDO 2.699* TOH 1.760** KYU 2.865*TOH 3.068** MAR 2.743** MAR 1.895** MAR 2.876**KYU 3.092** TOH 3.011 KOR 2.242** AMA 2.930**ANY 3.547* KOR 3.889* KAN 2.699* YAY 3.004MAR 3.653** YAY 6.181 SAK 3.193** SAK 3.134**KAN 3.889* THI 6.224 KAM 3.301 EDO 3.301MAN 4.041* AIN 6.333 YAY 3.421 TOH 3.634TAI 4.042* KAM 6.520 MAN 3.574 OKI 4.217KOF 4.046 HAI 6.540 KOF 3.891 AIN 4.269

Kofun Yayoi Tohoku Japanese Kyushu JapaneseYAY 1.669** KOF 1.669** EDO 1.760** EDO 1.423**KAM 2.715 MAR 2.158** KYU 2.254** TOH 2.254**MAR 3.291** KAM 3.004 MAR 2.676** MAR 2.388**SAK 3.443** SAK 3.326** KAN 3.011 KAN 2.696*OKI 3.461 EDO 3.421 KOR 3.068** KAM 2.865*AMA 3.467* AMA 3.520* KAM 3.634 KOR 3.092**EDO 3.891 TOH 4.130 YAY 4.130 SAK 3.728**KOR 4.046 KOR 4.449 SAK 4.977* AIN 4.511KYU 4.530 OKI 4.724 KOF 5.031 KOF 4.530JOM 4.928 KYU 5.127 AIN 5.064 YAY 5.127

Ainu Jomon Amami Islands Okinawa IslandEDO 4.196 AIN 4.268 KAM 2.930** SAK 3.116**JOM 4.268 KAM 4.901 KOF 3.467* KOF 3.461KAM 4.269 KOF 4.928 OKI 3.499** AMA 3.499**KYU 4.511 YAY 5.133 YAY 3.520* KAM 4.217TOH 5.064 MAR 5.930 SAK 3.626** MAR 4.561*MAR 5.567 OKI 6.170 ANY 3.939* YAY 4.724YAY 5.597 TOH 6.731 MAR 4.608** EDO 4.892KAN 6.333 KYU 6.837 KOR 5.028** HAI 5.107KOF 6.512 EDO 7.019 EDO 5.036 KOR 5.151SAK 6.760 AMA 7.287 KYU 5.831 KYU 5.243

Sakishima Group MarunouchiOKI 3.116** EDO 1.895**KAM 3.134** YAY 2.158**EDO 3.193** KYU 2.388**YAY 3.326** TOH 2.676**MAR 3.358** KAN 2.743**KOF 3.443** KAM 2.876**AMA 3.626** KOF 2.291**KYU 3.728** SAK 2.358**KOR 4.328** KOR 2.653**TOH 4.977* OKI 4.561*

*Distances significant at 5% level; **distances not significant at 1% or 5% levels.a Where the quantity (ni nj/niþ nj) D2

ij is distributed as chi-square with p degreesof freedom (ni¼ sample size of group i; nj ¼ sample size of group j); D2

ij¼ square ofthe generalized distance between groups i and j, and p¼ number of variables.

M. Pietrusewsky / Quaternary International 211 (2010) 42–54 51

series from eastern and western China form a close association, onethat is removed from more marginal southern Chinese series suchas Hainan Island and Taiwan and the Northern Asian series fromAnyang and Korea. The separation between southern (e.g., Hainanand Taiwan) and northern Chinese samples in the present resultssupports earlier work in physical anthropology (e.g., Wu et al.,2007; Zhang, 1999) and molecular genetic studies (e.g., Kivisildet al., 2002; Shi et al., 2005) that has demonstrated systematicdifferences between northern and southern Chinese. Hainan Island,Taiwan, Anyang, and Korea form a group that is intermediatebetween one that includes the remaining Chinese and all of theJapanese cranial series, an association that suggests that this regionis the likely source of Eastern Asians on both sides of the Sea ofJapan. The inclusion of the Shang Dynasty Anyang cranial series inthis intermediate clustering adds an element of antiquity to thisassociation.

The only Taiwan Aboriginal cranial series included in thesecomparisons, the Atayal, assumes a marginal placement in thecanonical plots as well as in the diagram of relationship based onMahalanobis’ distances. The groups closest to the Atayal are thosefrom Korea and Japan. The isolation of the Atayal points moretoward the long-term isolation on the island of Taiwan of this

M. Pietrusewsky / Quaternary International 211 (2010) 42–5452

non-Han Chinese tribal series, a conclusion that reinforces onereached by Howells (1986).

It is of particular note that the ‘‘Mongolian’’ cranial seriesrepresents the most differentiated and atypical series investigatedin this analysis. As has been noted by others (e.g., Brace and Tracer,1992) it is ironic that the term ‘Mongoloid’ was used by earlierresearchers to represent all the people of Eastern Asia. With theexception of the Mongolian cranial series, the remaining cranialseries representing Eastern Asia, including the prehistoric Jomon,modern Ainu, and Ryukyu Islanders of Japan, represent a singledivision of humanity, a division that is separate from anotherdivision that includes all of the mainland and island SoutheastAsian cranial series.

4.3. Population history of the Japanese archipelago

Turning next to the Japanese cranial series, there is relativelymarked differentiation between modern Japanese, pre-modernJapanese since Yayoi times, and an isolated group containing theJomon and Ainu cranial series. The Ryukyu Island series are mostclosely related to the Yayoi, Kofun and the medieval Kamakuracranial series from Japan. These results further indicate affinitiesamongst Marunouchi (a medieval series), Edo, and the modernJapanese cranial series from Tohoku, Kanto, and Kyushu. Themarked differentiation between the Japanese since Yayoi times andthe Jomon–Ainu series agrees with the general consensus thata major influx of new people into the Japanese archipelago (via theKorean Peninsula) began with the Yayoi (about 500 B.C.–300 A.D.)and Kofun (3rd–7th century A.D.) Periods (Imamura, 1996), a viewthat has been popularized in Kazuro Hanihara’s ‘dual structuremodel’ (Hanihara, 1991).

Again, as is evidenced in the diagram of relationship based ondistances (Fig. 3), a branch containing Taiwan, Hainan, and moreNorthern East Asian cranial series, Korea, and the Bronze-AgeChinese series from Anyang, is the first to connect to the Japaneseseries, a connection that is consistent with a Northeast Asian originof the Japanese beginning in the Yayoi Period.

Inspection of the closest distances further reveals that Edo,Tohoku, Marunouchi, Kanto, and Kyushu are among the seriesclosest to Manchuria, Anyang, and Korea. Korea is frequently closestto several of the Japanese series (e.g., Kanto, Edo, Kofun, Yayoi,Tohoku, and Kyushu). Likewise, closer inspection of the classifica-tion results indicates that several of the Yayoi specimens arereclassified as either Mongolia or Manchuria and several of theManchurian specimens are reclassified as Japanese. These andother classification results clearly point to biological connectionsbetween Northeast Asia (especially Korea, Manchuria, and Anyang)and the Japanese series. The Ryukyu Island misclassificationssuggest a much diversified population for the inhabitants of theseislands with affinities to Hainan Island, Taiwan, and severalNorthern Asian series.

The close relationship between Jomon and Ainu crania seriesfound in the present results supports numerous lines of evidenceincluding molecular and earlier studies of skull morphology (e.g.,Dodo, 1986; Turner, 1987; Dodo and Ishida, 1990; Yamaguchi,1992; Hanihara, 1993; Omoto et al., 1996; Pietrusewsky, 2000)that view the Jomon (people who inhabited the archipelago forapproximately 10,000 years beginning ca 12,000 years B.P.) as theancestors of the modern Ainu. An earlier date (16,000 years B. P.)for the beginning of the Jomon Period is discussed in Habu(2004).

The results of the present analysis provide little support fora close affinity between the Ryukyu Islanders, Jomon, and the Ainuand little evidence for a connection between these groups andthose in Southeast Asia. Several researchers (e.g., Turner, 1987,

1990; Hanihara, 1991) have suggested that Japan’s pre-agriculturalJomon populations; and by association the Ainu and RyukyuIslanders their presumed descendants, derive from a people livingin Southeast Asia during the Upper Paleolithic. Despite the lack ofa close relationship between the Ryukyu Island, Ainu, and Jomonseries, the results presented in this chapter indicate that all theprehistoric and modern cranial series from Japan, including thoserepresenting the Jomon, Ainu, and Ryukyu Islands, are members ofa greater East/Northeast Asian constellation, the geographicalancestral hearth of the earlier and later inhabitants of the Japanesearchipelago. Omoto and Saitou (1997) using genetic data havereached a similar conclusion.

5. Conclusions

The results of this multivariate craniometric analysis providea basis for summarizing some of the main points regarding thebiological relationships and possible origins of the inhabitants ofEastern Asia.

� The biological closeness of the modern cranial series fromNorthern and Northeastern Asia, including China, Korea, andJapan, indicate a recent common origin.� The Shang Dynasty cranial series from Anyang, Korea, and the

modern cranial series from Manchuria are among the seriesmost closely associated with the source population that ulti-mately was responsible for the modern Han Chinese, Korean,and Japanese, people who live on either side of the Sea of Japan.� The marked separation of East/North Asian and Southeast

Asian cranial series in this analysis supports models of long-term continuity within both regions rather than models thatadvocate population intrusion or replacement in Eastern Asiaand Southeast Asia.� Biological connections between post-Yayoi cranial series from

the Japanese archipelago and several mainland Northern Asiancranial series support scenarios that posit a major intrusion ofpeople from the Asian mainland beginning in the Yayoi Period.� The Ainu cranial series is closest to the prehistoric Jomon

suggesting an ancestral–descendant relationship betweenthese two groups, one that is well differentiated from the Yayoiand post-Yayoi Japanese cranial series.� The Ryukyu Island cranial series are more closely related to the

Yayoi, Kofun and Kamakura cranial series than they are tothe Jomon and Ainu cranial series, a finding that suggests thepresent day Ryukyu Islanders were greatly influenced bya population that began to expand into this arc of islands southof the main Japanese Islands beginning in the Yayoi Period.� Taiwan Aboriginals (Atayal) are well differentiated from Han

Chinese and only remotely related to other Eastern Asians.� Mongolia is the most differentiated Eastern Asian group

investigated in this analysis.

Acknowledgements

My thanks to Rona Ikehara-Quebral for her assistance with theanalysis of the data in this paper and to Rhea Hood, Joey Condit, andKaren Kadohiro for their help with the construction of the tablesand editing. Dr. Michele Toomay Douglas gave helpful advice andcomments on earlier drafts of this paper. Ms. Billie Ikeda helpedwith the figures used in this paper. My thanks to the Editors ofQuaternary International and two anonymous reviewers for theirhelpful comments and criticisms of earlier drafts of the paper.

M. Pietrusewsky / Quaternary International 211 (2010) 42–54 53

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