A morphometric analysis of the Late Pleistocene Human Skeleton from the Moh Khiew Cave in Thailand

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A morphometric analysis of the Late Pleistocene Human Skeleton from the Moh Khiew Cave in Thailand Hirofumi Matsumuraa,, Surin Pookajornb,y a

Department of Anatomy, Sapporo Medical University, S1, W17, Sapporo 060-8556, Japan. Department of Archaecology, Silpakorn University, Bangkok 10200, Thailand

b

Received 25 March 2004; accepted 5 May 2005

Abstract Few Late Pleistocene human remains have been found in Southeast Asia and the morphological features of the people of that age are still largely unknown due to the virtual lack of human remains in the area. Recent excavations at the Moh Khiew Cave in Thailand resulted in the discovery of a Late Pleistocene human skeleton in a relatively good state of preservation. An AMS radiocarbon date on the charcoal sample gathered from the burial gave a result of 25,8007600 BP, implying that the inhabitants of Moh Khiew Cave resided in a part of Sundaland during the last glacial age. In debates on the population history of Southeast Asia, it has been repeatedly advocated that Southeast Asia was occupied by indigenous people akin to present-day Australo-Melanesians prior to an expansion of migrants from Northeast Asia into this area. Morphometric analyses were undertaken to test the validity of this hypothesis. In the present study, cranial and dental measurements recorded from the Moh Khiew remains are compared with those of early and modern samples from Southeast Asia and Australia. These comparisons demonstrate that the Moh Khiew specimen resembles the Late Pleistocene series from Coobool Creek, Australia in both cranial and dental measurements. These results suggest that the Moh Khiew skeleton, as well as other fossil remains from the Tabon, Niah and Gua Gunung sites, represents a member of the Sundaland

Corresponding author. Fax: +81 11 618 4288. y

E-mail address: [email protected] (H. Matsumura). Deceased.

0018-442X/$ - see front matter r 2005 Elsevier GmbH. All rights reserved. doi:10.1016/j.jchb.2005.05.004

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population during the Late Pleistocene, who may share common ancestry with the present-day Australian Aborigines and Melanesians. r 2005 Elsevier GmbH. All rights reserved.

Introduction The Moh Khiew Cave is situated in an isolated limestone massive, 10 m above mean sea level and about 13 km inland from the coast in Krabi Province, Southern Thailand (Fig. 1). The cave is a rock shelter that covers an area 3 m wide and 30 m long. Excavations at this cave were undertaken in 1990 and 1991 by the second author and were reported in Pookajorn (1991, 1994). With reference to all archaeological evidence found at the Moh Khiew Cave, the cultural context of this cave was classified into five cultural levels, in his reports, as briefly summarised below. The first cultural level is predominantly characterised by the appearance of large unifacial pebble tools that correspond to the Late Palaeolithic of the Upper Pleistocene when pebble tools were used in preference to flake tools. The typology and raw material of the pebble tools from this cultural level are nearly identical to those found from the Lang Rongrien rock shelter located near the Moh Khiew Cave (Anderson, 1990), which was dated to more than 37,000 BP. However, the sample size from Lang Rongrien is so small (36 artefacts) that its apparent similarity with Moh Khiew is of dubious reliability (Bulbeck, 2003). The second cultural level produced abundant flake tools together with pebble tools and waste flakes made of fine-grained stone including chert, which can be distinguished from the first cultural level containing stone tools made of siliceous shale. This level also corresponds to the Late Palaeolithic. The burial, which is the focus of the current study, was found at the boundary of cultural levels 1 and 2. A relatively well preserved human skeleton was unearthed from this burial (Fig. 2). An AMS radiocarbon date on the charcoal sample gathered from this burial gave a result of 25,8007600 BP (University of Tokyo: TK-933Pr). Thus, the second cultural level cannot be older than 25,000 BP.

Fig. 1. Location of the Moh Khiew Cave Site in Krabi Province, Thailand.

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Fig. 2. Moh Khiew Cave skeleton from Krabi Province in Thailand: (1) Skeleton reconstructed in burial position; (2) right mastoid process; (3) various views of the facial skeleton and mandible.

The third cultural level mainly contains unifacial cobbles and bifacial stone tools similar to those found at Hoabinhian sites in Vietnam (Ha Van Tan, 1980). The fourth and fifth levels are characterised by Neolithic cultural remains including pottery and polished adzes. The Carbon 14 dates are around 6000 BP at the fourth level and 4000 BP at the fifth level, assigning these two cultural levels to the early Neolithic and the middle to late Neolithic periods of the Middle Holocene, respectively. During the last glacial age of the Late Pleistocene, lowered sea levels produced a huge area of adjoining land, the so-called ‘‘Sundaland’’, connecting mainland Southeast Asia and the western half of the Indo-Malayan archipelago which covered the present Malay Peninsula including Thailand (Chappell and Shackleton, 1986).

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Discoveries of human remains from the Sundaland of this period are scarce. Only the specimens from the Niah Cave in Sarawak (Brothwell, 1960; Kennedy, 1977), and the Tabon Cave in Palawan Island (Macintosh, 1978) are well known as Late Pleistocene fossil remains from this area. These specimens, however, are of nonadults or are poorly preserved and thus have not facilitated our understanding of the morphological characteristics of the Sundaland inhabitants and their relationship with present-day peoples. Among the debates on this issue, on the grounds of such insufficient evidence or later Early to Middle Holocene human remains as referred to later in this paper, it has often been asserted that Southeast Asia was occupied by indigenous people akin to present-day Australo-Melanesians before the expansion of migrants from Northeast Asia into this area (Callenfels, 1936; Mijsberg, 1940; Von Koenigswald, 1952; Coon, 1962; Jacob, 1967, 1975; Bellwood, 1987). The human skeleton unearthed from the Moh Khiew Cave is very important for researchers who attempt to understand such peopling of Sundaland during the Late Pleistocene. A description of the preservation and some palaeopathological comments were first provided by Choosiri (1993). In the present study, we describe the cranial and dental measurements recorded in the Moh Khiew skeleton, and report the results of statistical comparisons with early and modern samples from East Asia and the Southwest Pacific to test the validity of the hypothesis that people akin to AustraloMelanesians occupied this region in the past.

Material The Moh Khiew skeleton is currently housed at the Princess Maha Chakri Sirindhorn Anthropology Center in Bangkok, Thailand. Fig. 2 displays various aspects of the reconstructed skeleton. The calvaria is preserved, but was crushed and most of the temporal, frontal, and occipital regions were lost. The facial skeleton is in a relatively good condition, although the nasal bones were also lost. The mandible is nearly complete. The postcranial skeleton retains only the upper half of the body. The vertebral column and sternum are well preserved, but the ribs were broken into fragments. The right scapula is fragmentary, but the clavicle is complete. The bones of the left side were missing. The humeri remain only in parts of the shafts. This skeletal preservation has not facilitated sex determination because of the lack of the os coxae. Therefore, the sex must be determined by the remaining segments. The mastoid process is quite small and fine, the anterior part of the mandible is rounded without the mental tubercles, and the clavicle and humeri are slender and their muscle attachment areas are weakly developed. These characteristics indicate that the sex of this individual is probably female. Although the sex was not accurately determined from our observations, the current study attempts metric comparisons with other specimens regarding the sex as female. The occlusal surfaces of the dentition are heavily worn, and osteophytes are present at the rims of the vertebrae. Since other skeletal portions useful for age

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Table 1. Cranial, mandibular, and dental measurements (mm) of the Moh Khiew skeleton Martin no. & measurement

Martin no. & measurement

46. Bimaxillary breadth (108) 48. Upper facial height (67) 48(d). Cheek height 22 51. Orbital breadth 43 54. Nasal breadth 27 55. Nasal height (49)

65. Bicondylar breadth 66. Bigonial breadth 67. Bimental breadth 68. Mandibular length 69. Symphyseal height 69(1). Mand. height at mental 69(2). Mand. height at M2 69(3). Mand. body breadth 70. Ramus height 71. Ramus breadth 79. Mandibular angle

60. Alveolar length

62

61. Alveolar breadth

70

61(2). Bicanine breadth 62. Palatal length 63. Internal palate breadth 64. Palatal height

49 50 43 17

Buccolingual diameters 121 107 59 88 32 31

UI1 UI2 UC UP1 UP2 UM1

7.30 6.74 9.50 10.42 10.07 12.76

28

UM2

13.25

13

LI1

6.55

58 38 1151

LI2 LC LP1

6.93 8.73 8.84

LP2 LM1 LM2

9.00 11.34 10.89

( ) estimated value.

estimation such as cranial sutures are not observable due to the poor preservation, the age of this individual cannot be estimated at other than mature. Skeletal measurements were taken following Martin’s definitions (Martin and Saller, 1957; Bra¨uer, 1988). As shown in Table 1, measurements obtained from the Moh Khiew cranium are restricted to those from the facial skeleton and the mandible. The bimaxillary breadth is estimated as twice the measurement taken from the right half side of the maxilla, because the left side is incomplete. The upper facial height and nasal height are also estimated values, as the portion at nasion was missing. The estimate was made by adding 4 mm to the height between prosthion and the top of the frontal process of the maxilla (mnf: maxillonasofrontale) which is preserved at the height of frontomaxillary suture, since the vertical distance between the mnf and nasion was approximately 4 mm when we measured some representative female samples such as Jomon (n ¼ 18, mean ¼ 3.7 mm) and Japanese crania (n ¼ 15, mean ¼ 4.1 mm). Tooth crown measurements are also given in Table 1. Only the buccolingual diameters were recorded from the Moh Khiew dentition because the mesiodistal dimensions were obviously reduced from the original size due to heavy attrition. Cranial and dental measurements from representative Late Pleistocene and Holocene human specimens from East Asia and the Southwest Pacific regions were used as a basis for comparison with the Moh Khiew remains. A list of comparative cranial and tooth samples and data sources is given in Table 2. All the comparative data are from females.

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Locality

Period

Data (Skull)

N

Data (Dentition)

Coobool Creek Mai Da Dieu Middle Holocene Flores Liang Toge Guar Kepah Early to Middle Holocene Vietnam & Laos Ban Kao

Australia Vietnam Indonesia Indonesia Malaysia Vietnam, Laos

Late Pleistocene Early Holocene Middle Holocene Middle Holocene Middle Holocene Early to Middle Holocene

Brown (1989) Cuong (1986) — Jacob (1967) — —

9 1 — 1 — —

Brown (1989) Matsumura et al. (2001) Present study Present study Present study Present study

Thailand

Middle Holocene

Ban Chiang

Thailand

Early Metal age

Jomon Upper Cave 102,103 Liujiang

Japan China China

Middle Holocene Late Pleistocene Late Pleistocene

Sangvichien et al. (1969) Pietrusewsky and Douglas (2002) Ogata (1981) Wu (1961a, b) Woo (1959)

Australian Aborigines Australian Aborigines

Murray, Australia Swanport, Australia Papua New Guinea Bougainville Island Indonesia Philippine Hokkaido, Japan

Modern Modern

Papua New Guinea Nasioi Java Philippines Ainu N: sample size.

4 28

—

N 8 1 4 1 9 7

—

38 2 1

Pietrusewsky and Douglas (2002) Matsumura (1989) — —

33

Brown (1989) —

53 —

Brown (1989) Brown (1989)

50 23

Modern

—

—

Kanazawa et al. (2000)

57

Modern Modern Modern Modern

— — — —

— — — —

Bailit et al. (1968) Mijsberg (1931) Potter et al. (1981) Matsumura (1989)

210 — —

88 43 148 30

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Table 2. Comparative samples (females)

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Skeletal remains from two Late Pleistocene sites from China were used for comparison of the cranial metric data. The first is the Upper Cave (Shandingdong) (Nos. 102 and 103) series excavated from Zhoukoudian Cave, located southwest of Beijing (Wu, 1961a, b), and the second is the Liujiang cranium excavated from the Guangxi Zhuang Autonomous Region in southern China (Woo, 1959). As to the sex of the Upper Cave 102 and Liujiang crania, the present comparison treats these skulls as females, although those sexes remain in doubt (Wu and Poirier, 1995; Wolpoff, 1999). The teeth associated with these skeletal series were not used for comparison because the mandibular teeth are missing. The Mai Da Dieu specimen is a nearly complete female skull of the early Hoabinhian period (ca. 8000–10,000 BP) excavated from the Mai Da Dieu Site in Thanh Hoa Province, northern Vietnam (Cuong, 1986). This specimen is housed at the Institute of Archaeology in Hanoi. The Coobool Creek sample, collected from a site at Coobool Crossing on the Wakool River, is a Late Pleistocene Australian series. Descriptions of both the cranial and dental measurements were provided by Brown (1989). Middle Holocene Flores tooth samples were derived from skeletal remains from various sites in the western part of Flores Island. They are from the Liang Momer, Liang Toge, Liang X and Gua Alo sites, which range in date from ca. 7000 BP to ca. 4000 BP (Jacob, 1967). Among these early Flores specimens, as far as the female samples are concerned, the Liang Toge site produced a nearly complete skull, which can be used for cranial metric comparison. The Guar Kepah series was discovered from a shell midden site in Lenggong district, that belongs to the Hoabinhian cultural stage (Callenfels, 1936; Mijsberg, 1940), although some pottery was also excavated and the precise stratigraphy is unclear (Bellwood, 1997; Bulbeck, 2000). These early Flores and Malay specimens are stored at the Department of Palaeontology, Nationaal Natuurhistorisch Museum in Leiden. The early to middle Holocene Vietnamese and Laos sample consists of the remains from the Tam Hang site and the Tam Pong site in Laos (Fromaget, 1938; Huard et al., 1938), and those of the Bac Son cultural period from the Lang Cuom, Pho Binh Gia, Khac Kiem and Kio Phay sites in Northern Vietnam (ca. 10,000 BP–ca. 6000 BP; Mansuy and Colani, 1925; Huard et al., 1938). These specimens are housed at the Laboratorie d’Anthropologie Biologique, Musee de l’Homme Paris. The middle Holocene Thai specimens were excavated from the Ban Kao site in Kanchanaburi Province (ca. 2100–1600 BC, Sangvichien et al., 1969), which are kept at the Museum of Prehistory, Mahidol University in Bangkok. In addition, cranial and dental metric data of Early Metal Age Ban Chiang series (Pietrusewsky and Douglas, 2002) excavated from Northeast Thailand (Gorman and Charoenwongsa, 1976) were used for comparison. The Jomon specimens, excavated from various regions in Japan, were from the collection in the Department of Anatomy of Sapporo Medical University, the University Museum of the University of Tokyo, the Department of Anthropology of the National Science Museum, Tokyo, and the Laboratory of Physical Anthropology of Kyoto University. All of these samples date from the Middle to Final Jomon phases (ca. 4000–2300 BP).

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In addition to these remains, data recorded for several modern female samples were also used for the dental metric analysis.

Methods Twelve measurements of the crania and mandibles were selected for statistical comparison, because only these variables were measured in common for Moh Khiew and the published series (Table 3). Regarding the Coobool Creek specimens, the data of bicondylar breadth are reported only for the males. As a substitute, 92% of the mean value of males was assumed as the female mean value, because the sexual differences of bicondylar breadth in some representative population samples indicate a difference of between 91% and 93% (eg. Australian Aborigines: 92%, Brown, 1989; early Jomon: 91%, Ogata, 1981; Ban Kao: 93% Sangvichien et al., 1969). In cases where the mandibles were missing, only seven cranial measurements were used for analysis. As for the odontometric analysis, a total of 14 buccolingual crown diameters of the maxillary and mandibular teeth were used, since the mesiodistal measurements were not recorded from the heavily worn teeth of the Moh Khiew specimen. Table 4 shows dental measurements for the comparative female samples. Using these data, the following statistical analyses were performed to examine the affinities of the Moh Khiew specimen to each sample. Mahalanobis’ generalised distances were calculated using the cranial measurements to evaluate affinities in cranial morphology. The variance–covariance cranial matrices used for the distance computations were based on the pooled data from the modern Australian Aboriginal series provided by Peter Brown’s research data source (http://www-personal.une.edu.au/pbrown3/palaeo.html). Since the overall tooth sizes fluctuated between early and modern human populations due to techno-cultural factors such as cooking techniques, many researchers recommend using differences in proportion or shape alone for odontometric taxonomy (eg. Corruccini, 1973; Kieser, 1990). However, a recent odontometric study by Schnutenhaus and Ro¨sing, (1998) criticises, through their large-scale analysis using world-wide population samples, use of tooth measurements for taxonomy of modern populations. Although the dental proportion affinity may discord with such global taxonomy, many studies suggest that odontometric analysis provides useful information of population relationships within a few neighbouring taxonomic groups (e.g. Corruccini, 1973; Doran and Freedman, 1974; Hanihara, 1976, 1977; Harris and Nweeia, 1980; Hanihara, 1992). The present first author’s studies (Matsumura, 1994; Matsumura and Hudson, 2005) also demonstrate differences in tooth size proportion between Northeast Asians and AustraloMelanesians. In order to determine the relationship of the Moh Khiew specimen with its surrounding groups including Australo-Melanesians, therefore, similarities in proportion of dental measurements were estimated by a Q-mode correlation coefficient based on crown diameters, which entirely eliminates the size factor.

46

Sample

Bimaxill. Upper breadth facial height

Coobool Creeka Australian Aboriginesa Liang Togea Mai Da Dieua Ban Chianga Ban Kaoa Jomona Upper Cave 102a Upper Cave 103a Liujianga a

48

51

54

55

60

61

65

66

69

70

71

Orbital breadth

Nasal breadth

Nasal height

Alveolar length

Alveolar breadth

Bicondyl. breadth

Bigonial breadth

Symphys. Ramus Height height

Ramus breadth

99.0 90.5

68.6 65.6

43.7 42.3

28.0 26.4

50.0 47.3

60.4 59.1

67.8 64

115.1** 112.8

102.0 91.9

36.7 32.5

55.5 50.6

34.9 32.4

97 88 100.2 98.0 97.9 106.4 101.0 97.1

70 58 64.2 58.0 64 69.0 68.5 65.9

42 38 38.2 38.0 39.3 40.5 45.0 43.1

22 27 26.3 27.0 25.3 26.0 25.5 26.8

49 45 47.3 44.5 46.4 46.5 51.0 45.8

59 54 52.1 49.0 51.1 57.5 58.3 51.0

57 57 64.5 65.0 61.4 72.6 66.0 65.0

122 114 117.9 121.0 119.7

105 89 98.5 94.0 97.5

33 29 31.1 32.0 30.4

54 58 56.7 60.0 55.7

36 38 33.4 34.0 34.4

Data sources shown in ‘‘Data (skull)’’ of Table 2.

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Martin no.

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Table 3. Cranial and mandibular measurements (mm) of comparative female samples

Sample

a

Maxillary dentition

Mandibular dentition

I1

I2

C

P1

P2

M1

M2

I1

I2

C

P1

P2

M1

M2

1 9 4 1 7

7.30 7.46 8.05 7.32 6.89

6.74 7.14 6.83 6.43 6.55

9.50 8.62 9.20 7.97 8.73

10.42 9.67 10.25 9.36 9.11

10.07 9.64 10.03 9.05 8.98

12.76 11.99 12.43 12.12 11.26

13.25 12.03 12.27 11.54 11.39

6.55 6.21 6.04 5.47 6.07

6.93 6.90 6.52 6.61 6.46

8.73 8.00 7.99 7.37 7.74

8.84 8.63 8.48 8.45 8.30

9.00 8.65 8.88 8.42 8.61

11.34 11.28 11.17 10.92 11.19

10.89 10.34 11.09 10.26 10.72

33 9 50 23 55 88 142 43 210 30

7.00 8.00 7.80 7.80 7.76 7.48 6.84 6.90 7.00 6.83

6.60 7.40 6.70 6.70 6.90 6.67 6.08 6.30 6.38 6.29

8.00 9.00 8.80 8.80 8.60 8.58 7.55 7.80 7.71 7.56

9.40 10.00 10.00 10.00 9.89 9.93 9.25 9.40 8.96 8.99

9.20 10.00 9.90 9.80 9.93 9.97 9.10 9.20 8.75 8.71

11.40 12.80 12.60 12.60 12.13 11.69 10.79 11.20 11.37 11.26

11.10 12.90 13.00 12.60 12.12 11.76 10.74 10.90 10.97 11.00

5.60 6.30 6.40 6.20 6.51 6.40 5.47 5.50 5.68 5.39

6.10 6.60 6.60 6.40 6.82 6.55 5.84 6.00 6.10 5.90

7.50 8.30 8.00 8.00 7.93 7.88 6.84 7.20 7.11 7.02

8.10 8.80 8.70 8.80 8.34 8.11 7.53 7.70 7.50 7.59

8.40 9.00 8.80 8.60 8.78 8.75 7.97 8.20 8.02 8.03

10.90 11.50 12.20 12.00 11.05 10.83 10.15 10.70 10.83 10.71

10.30 11.60 11.60 11.60 10.93 10.63 9.86 10.20 10.06 10.02

Data sources shown in ‘‘Data (Dentition)’’ of Table 2.

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Moh Khiew Cave Guar Kepah Middle Holocene Flores Mai Da Dieua Early to Middle Holocene Vietnam & Laos Ban Chianga Coobool Creeka Australian (Murray)a Australian (Swanport)a Papua New Guineaa Nasioia Philippinesa Javaa Jomona Ainua

n

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Table 4. Buccolingual tooth crown diameters (mm) of the comparative samples (females)

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Finally, a cluster analysis using the unweighted pair-group method (UPGMA: Sneath and Sokal, 1973) was applied to these distance matrices for visual display of the relationships between samples. While a dendrogram of a cluster analysis is often useful for emphasising the closeness of certain samples or to stress distantly linked samples, this method does not allow sufficient recognition of samples intermediately positioned between major clusters because such samples will occasionally aggregate with one or the other major cluster. In order to further aid interpretation of the sample affinities, therefore, two-dimensional plots using the multidimensional scaling (MDS) method of Torgerson (1958) were provided simultaneously. Deviation diagrams were also drawn using the standardised measurements and the results were compared between the Moh Khiew specimen and the other samples.

Results Comparison of cranial and mandibular measurements Mahalanobis’ generalised distances between the comparative samples were calculated using the twelve cranial and mandibular measurements. The results are given in Table 5. The closest sample to the Moh Khiew specimen is the Late Pleistocene Coobool Creek from Australia, and the next closest is the modern Australian Aborigines. The Early to Middle Holocene Southeast Asian samples such as Liang Toge from Flores, Mai Da Dieu from Vietnam and Ban Kao from Thailand are distant from the Moh Khiew specimen. Fig. 3 represents a dendrogram resulting from a cluster analysis applied to the distance matrix of Table 5. The close similarities of the Moh Khiew specimen with the Coobool Creek and modern Australians Aborigines are confirmed. The Early to Middle Holocene samples of Liang Toge and Mai Da Dieu are secondarily clustered

Table 5. Mahalanobis’ generalized distance matrix based on 12 cranial and mandibular measurements

Coobool Creek Australian Aborigine Liang Toge Mai Da Dieu Ban Kao Ban Chiang Jomon

Moh Khiew

Coobool Aust. Creek Abor.

15.18 26.81

12.01

54.28 61.03 63.65 28.81 39.41

53.72 53.98 63.43 32.60 39.27

37.45 28.05 68.10 36.33 32.23

Liang Toge

Mai Da Dieu

Ban Kao Ban Chiang

45.33 102.61 58.38 36.35

56.03 39.62 27.73

15.15 23.70

6.89

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Fig. 3. Dendrogram of a cluster analysis applied to the Mahalanobis’ distance matrix of Table 5, based on 12 cranial and mandibular measurements.

Fig. 4. Two-dimensional plots of MDS applied to the Mahalanobis’ distance matrix of Table 5, based on 12 cranial and mandibular measurements.

with these Australian and Moh Khiew samples, while all the other Middle Holocene samples are separated from them and from other major clusters. The close affinities of Moh Khiew to Coobool Creek and modern Aborigines are further confirmed by the two-dimensional ordination of MDS values (Fig. 4). A comparison of the deviation patterns of the 12 cranial and mandibular measurements among the Coobool Creek and Mai Da Dieu with the Moh Khiew specimen is provided in Fig. 5. In order to standardise the measurements, data from modern Japanese (Morita, 1950) were used. The morphometrical affinity between the Moh Khiew specimen and the Coobool Creek sample is confirmed in this diagram. These two cranial samples are characterised by relatively wide orbital

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Fig. 5. Standardised deviations of 12 cranial and mandibular measurements of the Moh Khiew specimen and two comparative Late Pleistocene and Early Holocene samples, using modern Japanese as the basis for comparison.

Fig. 6. Standardised deviations of 12 cranial and mandibular measurements of the Moh Khiew specimen and Middle Holocene samples, using modern Japanese as the basis for comparison.

breadths, large alveolar lengths and breadths, wide bigonial breadths, and a wide ramus. Deviation diagrams of three Middle Holocene samples (Ban Chiang, Ban Kao, and Jomon) are provided in Fig. 6 together with that of the Moh Khiew specimen. In this comparison, Ban Chiang, Ban Kao and the Jomon exhibit patterns that contrast with that of Moh Khiew, and closely resemble those of one another.

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Comparison of cranial measurements Mahalanobis’ generalised distances were computed using only the seven cranial measurements to permit inclusion of those samples that lack mandibles. The results are given in Table 6. In this comparison, too, the distances place the Moh Khiew specimen closest to Coobool Creek. It is noteworthy that the next closest affinities to the Moh Khiew specimen are provided by Upper Cave No.103 followed by No.102 from Zhoukoudian. On the other hand, the Late Pleistocene Liujiang specimen from southern China represents the specimen most distant from the Moh Khiew cranium. Fig. 7 is a dendrogram from a cluster analysis applied to the distance matrix of Table 6. The Moh Khiew skeleton is grouped with the early and modern Australian Aborigines, and Upper Cave 103. The Mai Da Dieu and Liang Toge specimens are separated from the Moh Khiew skeleton. Fig. 8 provides a two-dimensional ordination of the distance matrix of Table 6. The close affinities between the Moh Khiew Cave, Coobool Creek and Upper Cave 103 specimens are repeatedly expressed in this schema. Comparison of dental measurements Table 7 gives a distance matrix transformed from the Q-mode correlation coefficients, which were computed using the 14 buccolingual diameters. Of the 15 samples, the Moh Khiew specimen shows the smallest distance from the samples from Early Holocene Vietnam and Laos; the next closest groups are the Middle Holocene Flores and Nasioi. These are followed by Coobool Creek, Papua New Guinea and the two modern Australian Aborigine groups. The Moh Khiew specimen is quite distant from the Jomon, Ainu, and any modern Southeast Asians who were analysed. Fig. 9 displays the results of a cluster analysis applied to the distance matrix of Table 7. All Australian and Melanesian samples form one major cluster, while the Jomon, the Ainu and modern Southeast Asians form another. Together with Middle Holocene Flores and the Early to Middle Holocene Vietnam and Laos samples, the Moh Khiew specimen is affiliated with the Australo-Melanesian samples. These groups are located on the left side of the horizontal axis on the two-dimensional scale drawn in Fig. 10, and are separated from the modern Southeast Asians, the Jomon and the Ainu, the latter of which are all found on the right side of the axis. The close affinity of the Guar Kepah sample is stressed on this two-dimensional plot, which was not expressed on the dendrogram of Fig. 9. On the basis of means and standard deviations from modern Japanese (Matsumura, 1994), deviation diagrams of the 14 buccolingual crown diameters for the Moh Khiew specimen and six prehistoric samples are illustrated in Fig. 11. The large size of the Moh Khiew teeth is comparable with that of the Coobool Creek dentition. In particular, the Moh Khiew specimen has quite large maxillary canines. The Coobool Creek, Early to Middle Holocene Vietnam and Laos, and Middle Holocene Flores samples, which cluster with the Moh Khiew specimen in Fig. 7, also bear proportionally large upper canines. From the viewpoints of both tooth size

Coobool Creek

Aust. Abor.

Liang Toge

Mai Da Dieu

U.Cave 102

U.Cave 103

10.82 25.64 44.35 46.42 20.47 15.79 46.92 36.12 23.65 30.57

5.34 31.92 26.01 31.32 5.26 27.09 33.24 23.22 21.84

28.23 14.79 45.43 11.75 28.18 36.44 28.32 22.40

43.74 68.10 24.24 51.05 79.23 50.28 33.87

70.16 39.96 46.21 32.86 28.41 24.64

37.10 28.01 22.62 14.74 22.12

27.50 44.93 32.44 25.85

Liujiang

23.25 20.18 10.77

Ban Kao

Ban Chiang

5.90 12.42

4.06

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Coobool Creek Australian Aborigine Liang Toge Mai Da Dieu Upper Cave 102 Upper Cave 103 Liujiang Ban Kao Ban Chiang Jomon

Moh Khiew

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Table 6. Mahalanobis’ generalized distance matrix based on seven cranial measurements

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Fig. 7. Dendrogram of cluster analysis applied to Mahalanobis’ distance matrix of Table 6, based on seven cranial measurements.

Fig. 8. Two-dimensional plots of MDS applied to Mahalanobis’ distance matrix of Table 6, based on seven cranial measurements.

proportion and absolute overall tooth size, the Jomon and Mai Da Dieu specimens, who have small teeth, are considerably different from the Moh Khiew specimen.

Discussion The present study of the Moh Khiew Cave specimen is also suggestive of the early indigenous people in this region. Our morphometric comparisons demonstrated close affinities between the Moh Khiew skeleton and Australian samples in both the cranial and dental segments. The Moh Khiew cranium exhibits the closest affinity to the Late Pleistocene Coobool Creek remains from Australia. Odontometrically, the

Guar Kepah

0.619 0.484

0.715

1.192 0.395

0.655 0.452

1.043 0.679

1.088

1.063 0.726 0.850

0.416 0.505 1.062

0.961 0.345 0.706

0.736 1.027 1.008

0.813

1.013

0.543

0.766

0.364

0.529 1.425 1.358 1.513 1.675

0.568 1.097 0.937 0.693 0.935

E–M: Early to Middle.

M-Hol. Mai Da E–M Flores Dieu Hol. Viet. & Laos

Ban Chiang

Coobool Murray Swanp. Creek Aust. Aust.

0.705 0.612 0.608

0.635 1.041

0.767

0.947

0.653

0.883

0.567

0.080

0.493

1.003

0.676

0.883

0.595

1.179

1.124

0.454 1.118 1.092 1.299 1.488

1.353 0.893 0.862 0.682 0.567

0.645 1.434 1.166 1.058 1.293

0.900 0.794 0.367 0.702 0.534

0.795 1.342 1.172 0.986 1.105

1.226 1.303 1.049 0.819 0.889

1.179 1.270 0.995 0.933 0.940

P. N. Guinea

0.180 0.788 0.970 0.757 1.221

Nasioi

Philip.

Java

Jomon

0.726 0.862 1.138 1.487

0.252 0.745 0.598

0.599 0.400

0.193

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Guar Kepah Middle Holocene Flores Mai Da Dieu E–M Hol. Vietnam & Laos Ban Chiang Coobool Creek Australian (Murray) Australian (Swanport) Papua New Guinea Nasioi Philippines Java Jomon Ainu

Moh Khiew

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Table 7. Distance matrix of Q-mode correlation coefficients (1–r) based on 14 buccolingual crown diameters

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Fig. 9. Dendrogram of cluster analysis applied to the distances of Q-mode correlation coefficients of Table 7, based on 14 buccolingual crown diameters. E–M Hol: Early–Middle Holocene.

Fig. 10. Two dimensional plots of MDS applied to the distances of Q-mode correlation coefficients of Table 7, based on 14 buccolingual crown diameters. E–M Hol. : Early–Middle Holocene.

Moh Khiew specimen resembles the Coobool Creek sample as well as some Early to Middle Holocene Southeast Asians. The deviation diagrams demonstrate that the Moh Khiew specimen shares common features such as the large alveolus, wide mandibular body and ramus, and buccolingually large teeth, in particular, maxillary canines, with the Coobool Creek sample. Furthermore, the visual impression of the

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Fig. 11. Standardised deviations of 14 buccolingual crown diameters, using modern Japanese as the basis for comparison.

Moh Khiew cranium and mandible accords well with those of the Coobool Creek remains in terms of the broad, straight and rectangular inferior orbital margin, the wide nasal aperture, the prominent canine eminences, the strong alveolar prognathism, the robust massive mandible with the weakly protruding mentum, and the convexly curved basal line of the corpus like a rocker jaw. As mentioned in the introduction, few Late Pleistocene human remains have been found in Southeast Asia. In East Malaysia, the Niah Cave in Sarawak is the wellknown site of the earliest appearance of human remains in Southeast Asia. The socalled ‘Deep Skull’ from Niah Cave has an associated radiocarbon date of ca. 40,000 BP (Kennedy, 1977). Brothwell (1960) examined this sub-adult skull and concluded that it bears the closest similarity to the Tasmanians. Tabon Cave on Palawan Island is the site which has produced the oldest human skeletal remains in the Philippines. The remains consist of a frontal bone and two mandibular fragments. AMS dating of this frontal bone resulted in a date of ca.16,500 BP. Macintosh (1978) considered this specimen to be linked to Australo-Melanesian populations. From Indonesia, two skulls were found in 1888 and 1890 at Wajak in Gunung Lawa, central Java. Dubois (1922), who first examined these specimens, labelled them ‘‘Proto Australians’’. Later, Weidenreich (1945) regarded the Wajak skulls to be akin to the terminal Pleistocene specimen from Keilor, Australia. Wolpoff et al. (1984), in their analysis of those specimens, supported these conclusions. In contrast, Jacob (1967) supported a southwest Pacific affinity for these skulls, whereas in fact he found the so-called ‘‘Mongoloid’’ features. Storm (1995) examined more details of the Wajak skeletons and found similarities to modern Indonesians rather than to Australians. Hence, the assessment of the morphological affinity of the Wajak remains differs between researchers. In addition to this problem, the chronological framework of these fossils has been unclear until recently. The age of these fossils has long been regarded as Late Pleistocene, but

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more recent AMS dating of the skeleton indicates that a Middle Holocene date (ca. 6500 BP) is more appropriate (Storm, 1995). A skeletal morphology similar to modern Australo-Melanesians is manifested not only in these Late Pleistocene fossils, but also in some Early Holocene remains from Southeast Asia. In the northern Malay Peninsula, several sites have produced EpiPalaeolithic and Hoabinhian cultural human skeletons. Gua Gunung Runtuh in the Lenggong Valley in the State of Perak was excavated by Zuraina (1994), and revealed a 10,000–11,000 year-old human skeleton (Jacob and Soepriyo, 1994). There is, however, a possibility that this skeleton is younger than the estimated date because the radiocarbon date was obtained from freshwater shellfish in a limestone rich region, which are likely to be in order of 2000 years younger than the estimation (Bulbeck, 2003). Analyses of measurements obtained from these remains by Matsumura and Zuraina (1995, 1999) demonstrated that this male individual was marked by Australo-Melanesian characteristics in both the dentition and limb bones. From Lenggong district, Evans (1918) discovered a fragment of human jaw with some teeth at Gua Kajang. Furthermore, Gordon, Evans and Callenfels collected fragmentary human bones at Gua Kerbau in Gunung Pondok. Duckworth (1934) interpreted these remains to have features of the present-day Australo-Melanesians. From then Wellesley Province in northern Malaysia, Callenfels (1936) and Mijsberg found human remains from a shell midden site at Guar Kepah, and identified the specimens as ‘‘Palaeo-Melanesians’’ (Mijsberg, 1940) of early Holocene date. As far as the dental metrics are concerned, the present study also demonstrated that Guar Kepah has a relatively close similarity to Melanesians such as in Papua New Guinea as shown in Table 7. The Gua Cha remains excavated by Sieveking (1954) and later by Adi (1985), from a rock shelter located in the State of Kelantan, were recognised as dating to the Hoabinhian and Neolithic cultures (ca. 10,000–2000 BP). Trevor and Brothwell (1962) concluded that the Gua Cha skeletons also have a close affinity with Melanesians. On the other hand, Bulbeck (2000) studied nonmetric dental traits of the Gua Cha specimens, and found Australian and then Polynesian affinities overall but an interesting similarity between the Gua Cha and Jomon teeth. From northern Vietnam, nearly complete skulls of the early Hoabinhian period (ca. 8000–10,000 BP) were excavated from the Mai Da Nuoc and Mai Da Dieu sites in Thanh Hoa Province. A comparison of the Mai Da Dieu female specimen in the current study does not reveal a resemblance to Australians. These results stand at odds with those obtained from the Mai Da Nuoc male specimen described by Cuong (1986) as possessing a cranial morphology similar to Australian Aborigines, despite possessing some East Asian features. From most of these studies of Late Pleistocene and Early Holocene human remains, the general view has been that Southeast Asia was occupied by indigenous people belonging to the Australo-Melanesian lineage before the expansion of migrants of North or East Asian lineage through southern China into this area (Callenfels, 1936; Mijsberg, 1940; Von Koenigswald, 1952; Coon, 1962; Jacob, 1967 1975; Bellwood, 1987). This population history scenario for Southeast Asia is known as the ‘Immigration’ or ‘Two Layer’ model. The Two Layer hypothesis is supported by a wide range of genetic, linguistic, and archaeological evidence. Classic genetic

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markers and recent mtDNA analyses (Ballinger et al., 1992; Cavalli-Sforza et al., 1994; Omoto and Saitou, 1997; Tan, 2001) have found many biological similarities between Chinese and Southeast Asian samples. Linguistic and archaeological studies have linked the premodern expansion of the Austronesian and Austroasiatic language families with the dispersal of rice cultivating populations during the Neolithic period (Renfrew, 1989, 1992; Bellwood, 1991, 1993, 1996, 1997; Bellwood et al., 1992; Blust, 1996; Glover and Higham, 1996; Higham, 1998, 2001; Bellwood and Renfrew, 2003; Diamond and Bellwood, 2003). Both linguistic and archaeological considerations suggest that southern China and Taiwan were the ultimate sources of these language and population dispersals. On the contrary, some recent cranial and dental studies suggest different views. Turner’s studies (Turner, 1989, 1990, 1992) of nonmetric tooth traits showed that both the early and modern Southeast Asians display the so-called ‘‘Sundadont’’ dental complex, as do teeth of Australian Aborigines. This may indicate that Australian Aborigines and Southeast Asians had originated from a common ancestral population, which occupied the late Pleistocene Sundaland. On the other hand, cranial studies by Hanihara (1993,1994) advocated that the Proto-Malay like the present-day Dayak widely inhabited Southeast Asia during the late Pleistocene. He regards the Proto-Malay as an original source for present-day Southeast Asians including the Jomon Japanese. These researchers consider that the present-day Southeast Asians have evolved by local adaptation without large admixture with North/East Asians. In addition, Hanihara’s assessment does not favour the hypothesis of the occupation of an Australian lineage in early Southeast Asia. Our study of the Moh Khiew skeleton is not concerned with the problem of whether the present-day Southeast Asians hybridised with North/East Asians or not, but it is suggestive of the early indigenous people in this region. Many of the prehistoric specimens discussed above were not used for the present comparison with the Moh Khiew female, because most of them are fragmentary, non-adult, unknown sex or males. The results of morphometric comparison of the Moh Khiew specimen with several female samples suggest that, along with other fossil specimens from Tabon, Niah, and some Early Holocene Southeast Asians including the Gua Gunung Runtuh, it was a member of a population of Sundaland dwellers during the Late Pleistocene, who may share common ancestry with the present-day Australian Aborigines and Melanesians. Until the end of the last glacial stage, lowered sea level reduced the water barrier between Australia and Southeast Asia (Bowdler, 1992). Against this geographic background, during the Late Pleistocene, genetic separation between the inhabitants of Australia and Southeast Asia is considered to have still been incomplete. Some traits exhibited in this skeletal and dental morphology may have been retained in early Southeast Asia. Oota et al. (2001) analysed mtDNA recovered from the Moh Khiew specimen and found continuity with the Semang Negrito foragers living in the Malay Peninsula. Although the present study does not deal with Orang Asli people such as the Semang or Senoi, earlier researches stressed biological links between the Semang and Melanesians (Hughes, 1965; Birdsell, 1972; Carey, 1976). Further, Ballinger et al. (1992) found similar haplotypes in Semang mtDNA sequences to those found among

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Papua New Guineans. Dental and cranial studies by Rayner and Bulbeck (2000) and Lauer (2002) found a close relationship between the Malay Hoabinhian series and the Semang and Senoi, as well as with Melanesians and Africans. These findings accord with our conclusion that the Moh Khiew specimen had a close affinity with the Australo-Melanesians, although the relationship of this specimen with the Orang Asli people was not revealed. The Minatogawa skeletons, discovered from Okinawa Island, Japan, are other well-known representatives of the Late Pleistocene inhabitants of East Asia (Suzuki, 1982). This series was not compared with the Moh Khiew specimen in the current study because comparable data were not available from the Minatogawa female samples. Nevertheless, the features observed in the Minatogawa male crania, such as the lower alveolar prognathism and smaller dentition, are different from those of the Moh Khiew specimen. Furthermore, as stated by Suzuki (1982), the Minatogawa remains, which share many common cranial features with the Jomon people, are considered to be the direct ancestors of the Jomon people in Japan. The dissimilarity between the Moh Khiew and Jomon samples, revealed in our study, supports this perspective. It may be worth pointing out in passing that, in the cranial metric comparison, the Moh Khiew cranium also has a close affinity to the Upper Cave crania from Zhoukoudian. Concerning the female crania, Weidenreich, (1939) regarded the No. 102 cranium as a primitive Melanesian, and the No. 103 skull as an ancestor of the Eskimos. However, few later studies support this view (e.g. Howells, 1989; Cunningham and Wescott, 2002). Some researchers regard the morphology of these crania as being archaic features common to Late Pleistocene human fossils from around the world (Brown, 1996; Wolpoff, 1999); nevertheless, some other scholars assert their linkage to the Australian Aborigines (Kamminga and Wright, 1988; Stringer, 1999) or to the Easter Islanders (Wright, 1992). Although the comparisons in the current study were made using limited portions of the skulls, the closeness of the Moh Khiew specimen to the Upper Cave specimens supports the latter view, implying that the Sundaland inhabitants had a genetic linkage with the people in northern mainland East Asia in the Late Pleistocene.

Conclusions Statistical comparisons of cranial and dental measurements reveal that the Moh Khiew specimen is most similar to Australian samples, especially the Late Pleistocene series from Coobool Creek. These findings further suggest that the Moh Khiew skeleton, as well as other fossils from Tabon and Niah, are representative of an early group of people who occupied Sundaland during the Late Pleistocene, and may share common ancestors of Australian Aborigines and Melanesians. Further, craniometric similarities are also found between the Moh Khiew specimen and crania recovered from the Upper Cave (No.103) at Zhoukoudian, China.

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Acknowledgements The first author wishes to express his great sadness over the death of the coauthor, Prof. Surin Pookajorn, who excavated the Moh Khiew skeleton, after this manuscript was initially submitted. We are grateful to Dr. John de Vos, Nationaal Natuurhistorisch Museum, Netherlands, for permission to investigate the skeletal collections in his charge. We also wish to express our gratitude to Dr. Nancy Tayles, Department of Anatomy and Structural Biology, Otago School of Medical Sciences, University of Otago, and Dr. David Bulbeck, School of Archaeology and Anthropology, Australian National University for their invaluable assistance with the preparation of this paper. Thanks are also due to Dr. Mark Hudson for editing this manuscript. This study was supported in part by a Grant-in -Aid in 2002 (No. 14540664) and 2003 (No. 15405018) from the Japan Society for the Promotion of Science.

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