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Bioarchaeology of Early Neolithic Skeletons from the Nankuanli East Site, Southwestern Taiwan Michael Pietrusewsky 1, Adam Lauer 1, Cheng-hwa Tsang 2, Kuang-ti Li2, Michele Toomay Douglas1. 1Department of Anthropology, University of Hawaii at Manoa, Honolulu, USA 2Institute of History and Philology, Academia Sinica, Taipei, Republic of Taiwan

Abstract

In this study, we examine the health and way of life of some of Taiwan’s earliest Neolithic peoples through studies of skeletons from the Nankuanli East (NKLE) site. The NKLE site is one of three oldest sites (ca. 4500-5000 BP) identified during salvage excavations in 2002-2003 in the Tainan Science Park (TSP) in Shanhua District, Tainan City, in southwestern Taiwan. Approximately 82 extended and supine burials and extensive archaeological materials including pottery, ornaments, shell-bracelet funerary objects and the remains of domesticated and wild animals were recovered from this site. The main subsistence base of these early Neolithic peoples included extensive marine exploitation, hunting, and collecting of wild plant resources as well as early farming involving the cultivation of small grains, root and fruit crops. The presence of foxtail millet has also been identified for this site. Twenty-three (15 male and 8 female) of the most complete and well-preserved burials from this site are used in this study. In addition to documenting two forms of dental modification, betel staining and tooth ablation, we examine the oral/dental and physiological health of these early inhabitants of Taiwan using several indicators including: adult stature, cribra orbitalia (CO), linear enamel hypoplasia (LEH), and dental pathology (dental caries, antemortem tooth loss (AMTL), alveolar defects, dental calculus, alveolar resorption, and dental attrition). Comparisons are made between males and females, with Iron Age skeletons from the Shisanhang (SSH) site in northwestern Taiwan, and with skeletal series outside of Taiwan. More than two-thirds of the teeth from the NKLE skeletons exhibit staining that is likely attributed to chewing of Areca nut. The frequency of betel-stained teeth is significantly greater in males than in females. With few exceptions, the most common pattern of tooth ablation in the NKLE skeletons is the bilateral removal of the maxillary lateral incisors and canines, a pattern that has been recorded among some Indigenous groups in Taiwan and in other Neolithic skeletons from Taiwan. This pattern is rarely observed outside Taiwan. No tooth ablation and very little betel staining are observed in the Iron Age skeletons from the Shisanhang site. Overall, very few differences are observed in the health of males and females from the NKLE site and between this early Neolithic site and the Iron Age site of Shisanhang. The estimated average stature for NKLE males is 160.9 cm and 155.2 cm for females. Regional comparisons of health indicators suggest that the earliest Neolithic inhabitants of Taiwan may have experienced more childhood physiological stress for at least one indicator (LEH) than that observed in other series. However, the frequency of another indicator of childhood health in the NKLE skeletons, CO, was significantly lower when compared to other skeletal series outside Taiwan. Overall, the frequencies of

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several indicators of oral infection (e.g., AMTL, dental caries, and alveolar defects) observed in the NKLE and SSH skeletons are among the lowest reported indicating generally good dental health for Taiwan’s prehistoric inhabitants. Future research, involving additional skeletons from the NKLE site as well as other sites from Taiwan, will expand on the research reported in this study. This study contributes to studies in anthropology that attempt to reconstruct past behaviors and health from archaeological human skeletons. Keywords: indicators of health, stature, cribra orbitalia, linear enamel hypoplasia, tooth ablation, betel staining, dental pathology, Nankuanli East, Neolithic and Iron Age Taiwan National Research Council of Taiwan supported this research.

Introduction

Archaeological salvage excavations at the Tainan Science Park (TSP), located in southwestern Taiwan, have identified at least 80 sites extending from the earliest Neolithic, approximately 4,500- 5,000 years BP, to the first Han Chinese settlements of a few hundred years BP. In addition to a rich record of cultural and faunal material, human skeletons representing approximately 2,000 individuals have been painstakingly and meticulously excavated from these sites. Two of the earliest Neolithic sites within the TSP complex are Nankuanli, and Nankuanli East (NKLE). The cultural horizon associated with these sites is designated the Tapenkeng (TPK) culture, Taiwan’s initial Neolithic culture known for its thick cord-marked pottery, which appeared approximately 3500-3000 BC (Chang 1969; Li 2013; Tsang 1995, 2005). In addition to cord-marked pottery, the TPK culture is characterized by stone adzes, net sinkers, and bark-cloth beaters (Chang 1969, Li 2013; Tsang 2005). Neolithic sites are generally located in coastal and riverine environments. The unique features of this new culture and its relatively rapid dissemination throughout Taiwan suggest its likely source was from neighboring coastal southeastern China (Bellwood 2005; Jiao 2007; Tsang, 2005, 2010, 2012).

The major subsistence base of these early Neolithic cultures included extensive marine exploitation, hunting, collecting wild plant seeds [e.g., “bitterwood (Picrasma quassiodes) and Chinese hackberry (Celtis sinensis)], as well as early farming. The discovery of rice and foxtail millet at Nankuanli and NKLE sites suggest early Neolithic people were already engaged in intensive agriculture (Li 2013). While cereal crops such as rice and millet formed a part of the subsistence base, hunting, marine and riverine fishing, and collecting of marine resources were dominant attributes of this subsistence economy (Pietrusewsky and Tsang 2003). Foxtail millet is still cultivated by Taiwan’s Indigenous people (Tsang 2005).

Archaeological and historical linguistic evidence suggest that the Neolithic cultures on both sides of the Taiwan Strait have figured prominently in identifying the homeland of Austronesian speakers. The expansion of cereal (rice and millet) based economies from coastal regions of southern China to Taiwan, the Philippines, eastern Indonesia, and ultimately throughout Oceania has been documented (Bellwood 2005; Jiao 2007; Tsang 2010, 2012).

The introduction of iron, commencing approximately 2000 years BP, gave rise to several regional Iron Age cultures in Taiwan, including the Shisanhang cultural tradition (Tsang 2000). The Shisanhang ceramic tradition included many different forms of pottery, including bottles, bowls, basins and globular cooking pots. In addition to tools made of stone, bone, and antler, there is also evidence of an iron-smelting furnace and a workshop indicating the people possessed knowledge of metal working at Shisanhang. There is also evidence of long-distance trade with the Chinese during Tang and Song Dynasties for exotic goods, such as agate beads and ornaments, bronze artifacts, Chinese coins, glass beads and bracelets, gold ornaments, porcelain, and silver objects, etc. (Tsang 2000).

There is now widespread acceptance that the indigenous people living in Taiwan prior to the major immigration of Han Chinese, beginning in the 17th century, are the direct descendants of people associated with these early Neolithic cultures of Taiwan. Previous bioarchaeological research, especially in the Western Hemisphere and western Eurasia, has reported an increased prevalence of dietary stress and disease following the transition to agricultural and economic intensification (e.g., Cohen and

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Armelagos 1984; Cohen and Crane-Kramer 2007; Larsen 2002, 2006; Rose et al. 1991; Steckel and Rose 2002). This research demonstrated that the frequencies of indicators of health such as linear enamel hypoplasia (LEH), cribra orbitalia (CO), and dental pathology increase significantly in response to malnutrition, infectious diseases, and changes in diet following the transition to agriculture. Likewise, dental attrition is found to generally decline following the transition from foraging to farming. Bioarchaeological research in mainland Southeast Asia, however, has revealed a different and more complex pattern. Unlike the pattern associated with wheat and maize agriculture, deterioration in health associated with the transition to sedentism and the intensification of agriculture in Southeast Asia is not so readily apparent. Among the reasons advanced to explain this discrepancy is the relative low cariogenicity of rice and the retention of broad-spectrum subsistence strategies associated with agricultural intensification (Domett and Tayles 2007; Douglas and Pietrusewsky 2007; Halcrow et al. 2013; Oxenham 2004; Oxenham et al. 2006; Pietrusewsky and Douglas 2002). A preliminary investigation of a sample of Neolithic skeletons from NKLE, which makes comparisons with later Iron Age skeletons from Taiwan and other prehistoric series outside of Taiwan, has provided the first glimpses of the health and lifestyle of these early inhabitants (Pietrusewsky et al. 2014a). Using several dental indicators of health, it was observed that the earliest inhabitants of Taiwan might have experienced considerable childhood stress. However, relatively low frequencies of many of the dental pathologies observed in the NKLE skeletons indicated generally good dental health for these early prehistoric inhabitants of Taiwan. With the exception of more dental attrition in males, there were few significant differences between sexes for these indicators of health. Comparisons of the early Neolithic and later Iron Age skeletons suggested only two indicators of dental health (alveolar resorption and dental calculus) were significantly different, consistent with the transition to more advanced food processing techniques. Likewise the reduced amount of dental attrition observed in the Iron Age skeletons compared to those of the early Neolithic, was expected given an increased reliance on softer processed foods and the use of metal tools during the Iron Age. The higher than expected frequency of LEH observed in the early Neolithic inhabitants of Taiwan suggests they may have experienced considerable nonspecific physiological stress, including the possibility of infectious diseases and/or metabolic disorders, during early childhood. However, the overwhelming similarities in dental health of the Neolithic and Iron Age inhabitants of Taiwan suggest that, despite being separated by several thousand years, these prehistoric people shared very similar subsistence bases including farming, fishing and the hunting, and the gathering of marine and terrestrial resources. Regional comparisons indicated that the earliest Neolithic inhabitants of Taiwan experienced more childhood physiological stress than observed in other prehistoric skeletons from outside Taiwan. Overall, however, the frequencies of several indicators of dental infection in the Neolithic and Iron Age skeletons are among the lowest reported suggesting the dental health for Taiwan’s prehistoric inhabitants was good. Our discussion of the reasons for the differences and similarities focused on possible differences in subsistence economies, diet, hygiene, and cultural behaviors. Taking this as background, we now expand our examination of the health of Taiwan’s earliest Neolithic inhabitants. In addition to the indicators used in the earlier study (linear enamel hypoplasia (LEH), dental caries, antemortem tooth loss - AMTL, alveolar defects, dental calculus, alveolar resorption, and dental attrition) we evaluate two additional indicators of physiological health, adult stature and cribra orbitalia (CO). We

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further document two forms of dental modification, betel staining and tooth ablation, observed in the NKLE skeletons. Given the general observation that differences in diet may explain differences in male and female health, we again make comparisons between the male and female skeletons from NKLE. As well, we examine whether there are differences in health between the early Neolithic and Iron Age skeletons that support, or refute, the observation that health generally deteriorates with the transition from Neolithic to Iron Age in Taiwan. Finally, limited comparisons of the NKLE skeletons and those from surrounding regions are made that provide broader regional context for these data.

Material

Nankuanli East (NKLE) Site—Early Neolithic

The NKLE site is located five hundred meters east of the Nankuanli site on the flood plains of Shanhua District, Tainan City, in southwestern Taiwan (Figure 1). Salvage excavations at NKLE in 2002 and 2003 recovered approximately 82 burials (42 adults and 40 subadults) and extensive archaeological materials that are very similar to those recovered from the neighboring Nankuanli site. The predominant mortuary pattern identified includes extended supine burials associated with pottery, bead and pendant ornaments, shell-bracelet funerary objects, and other artifacts (Figure 2). In addition to ceramic ware (i.e., jars and bowls), the archaeological record includes the presence of foxtail millet (Setaria italic L.), dogs, deer, wild boars, fish, shellfish, reaping knives made of shell, fishing sinkers, ground adzes and projectile points (Bellwood 2006; Li 2013; Tsang 2005). Although terrestrial mammals, especially dog, deer, and wild boar, were important, fish represent the most abundant potential food source to the inhabitants of NKLE (Li 2013).

Because the skeletons from NKLE site experienced extensive postmortem taphonomic changes due to 7-8 meters of overlying sediment, very few skulls or bones from the postcranial skeleton are complete. Further, given the manner in which these skeletons were excavated and stabilized (requiring time consuming extraction of the bones from plaster), only the skeletons of 24 of the most complete NKLE burials were available for the present study (Table 1). Eighteen individuals (9 male and 9 females) died as young adults (20-35 years of age) and six (5 male and 1 female) as middle-aged (35-50 years of age) adults.

Shisanhang (SSH) Site—Iron Age More than 200 hundred well preserved human skeletons, many in the flexed position with heads facing southwest, were excavated from the Shisanhang (SSH) site. The site, which dates from approximately 1800 – 500 years BP, is located on the northwestern coast of Taiwan near the modern Bali District, New Taipei City, near the mouth of the Danshui River (Tsang 2000; Yang 1961). Our sample consists of 23 of the best preserved and most complete adult skeletons (Pietrusewsky and Tsang 2003). Eighteen of the specimens examined died as young adults (20-35 years of age) and five were middle-aged adults (35-50 years) at the time of their death (Table 1). Comparative Skeletal Assemblages Selected skeletal assemblages from the surrounding regions are used for comparison (Table 2). The approximate location of these series and those from Taiwan are shown in the map in Figure 3.

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Methods

Standard osteological methods were used to estimate sex and age-at-death (Buikstra and Ubelaker 1994; Pietrusewsky and Douglas 2002). The skeletal and dental indicators of health investigated include those attributable to non-specific systemic stress during growth and development (e.g., stature, cribra orbitalia, LEH) and those that suggest specific stressors (e.g., dental disease). We also report two forms of dental modification observed in the NKLE dentitions, tooth ablation and betel staining. Fisher’s exact test (FET) was used to test for significant differences of the discontinuous data and Student’s t-test for continuous data (Thomas 1986). Stature Given their fragility, measurements of the length of limb bones were recorded while the skeletons were in their natural state of articulation (see Figure 3). The maximum length of the femur was used to estimate stature for adult males and females. Because of the lack of population-specific stature formulae for estimating adult stature in prehistoric skeletons from Taiwan, estimates use several different stature formulae for modern Asian (e.g., Trotter and Gleser 1958; Sangvichien et al. 1985, n.d., Shao 1989) and non-ethnic formulae (Sjøvold 1990). Female stature was estimated using comparable formulae (Pearson 1899; Sangvichien et al. 1985; Sjøvold 1990). Cribra orbitalia (CO) Commonly attributed to iron deficiency anemia (Stuart-Macadam 1989, 1991; Oxenham and Cavill 2010) or megaloblastic anemia (Walker et al. 2009), cribra orbitalia (CO) is characterized by the sieve-like lesions that develop in the orbital roofs (Figure 4). In this study we report the frequency of cribra orbitalia by individual. Linear enamel hypoplasia (LEH) Linear enamel hypoplasia (LEH), one of the most common forms of dental enamel hypoplasia observed in archaeological skeletal remains, is characterized by one or more transverse grooves of varying depths on the tooth crown surface (Figure 5). Although a single cause can rarely be assigned to a specific hypoplastic defect, this indicator has been linked to malnutrition, metabolic disorders, acute and chronic infections, physical trauma, and hereditary conditions that affect the mother and/or growing child during infancy and early childhood, conditions that can disrupt enamel production (Goodman et al., 1984; Goodman and Rose 1990, 1991; Hillson 2008). Because defects are more frequent in the incisor and canine teeth, we report any manifestation of LEH, on a presence/absence basis for these teeth only. Dental Modification and Dental Pathology  Tooth Ablation The intentional removal of teeth, especially of the anterior teeth (Figure 6), represents a common dental modification observed in archaeological human skeletons. Distinguishing between antemortem (before death) and postmortem (after death) tooth loss is relatively straightforward. Empty sockets that exhibit no remodeling of the alveolar bone generally indicate postmortem tooth loss while the presence of alveolar bone resorption is generally indicative of antemortem tooth loss. The loss of teeth before death may be attributed to a number of processes, including dental pathology (caries,

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alveolar defects, periodontal disease), accidental trauma, unerupted and/or impacted teeth, and cultural modification. Although most antemortem tooth loss in archaeological remains is generally the result of pathogenesis, the identification of tooth ablation in archaeological remains frequently relies on the recognition of a repetitive pattern of missing teeth in the presence of otherwise good dental health. Criteria used to identify tooth ablation include the spacing of teeth, symmetry and non-randomness of loss, the absence of significant dental pathology, presence of roots, fractures of the labial portion of the alveolus and ethnographic accounts. Betel Staining Chewing of areca, or betel, nut (Areca catechu), with or without the use of pepper leaf (Piper betle) and/or slaked lime, is a common cultural behavior observed among people living today in South Asia, Southeast Asia, Taiwan, and the western Pacific,. Regular use of betel quid causes a bright orange/red stain of the teeth and gums. The staining observed in the NKLE teeth is usually light to dark brown/reddish brown in color.

Because there is now a substantial epidemiological literature demonstrating a link between chewing betel nut and cariostasis (e.g., Howden 1984; Möller et al. 2009; Schamschula et al. 2010) and other oral-dental pathologies including periodontal disease in living people (e.g., Chatrchaiwiwatana 2006; Trivedy et al. 2002), and a documented increase in dental attrition (IARC 2004; Kumar et al. 2004), we report the frequency of stained teeth for our skeletal series. Betel stained teeth and dental pathology were scored on a per tooth basis and are reported in adults (>15 years) with sex estimates. Antemortem Tooth Loss Antemortem tooth loss (AMTL), the loss of teeth before death, can be attributed to several pathological processes including periodontal disease, carious lesions, and alveolar defects. The principal criterion for recognizing AMTL is alveolar bone remodeling following the loss of teeth. Tooth ablation, or the ritual removal of teeth during the lifetime of an individual, is not included in the reported frequencies of AMTL. Dental Caries Dental caries is the result of demineralization of the tooth structures caused by organic acids produced by bacterial processes involved in the fermentation of dietary carbohydrates (Hillson 2008:313). Dental caries are scored on a presence/absence per tooth basis (Figure 7). Alveolar Resorption and Alveolar Defects Two distinct types of alveolar bone loss were recorded. The first type, referred to in this study as alveolar resorption, is related to the removal of alveolar bone due to inflammation of the supporting tissues of the teeth known associated with periodontal disease (Hillson 2008). The second type of alveolar bone loss is concentrated around the apex of the tooth roots, originating from infections of pulp, so-called periapical inflammation (or periapical dental abscess (Dias and Tayles 1997; Hillson 2008; Oxenham et al. 2006). We have adopted the term alveolar defect for this type of bone loss in the present paper. In this study, alveolar resorption is scored as the amount (none, slight, moderate, or marked) of tooth root exposed above the alveolar bone margin. Alveolar defects were scored on a per tooth/socket basis.

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Dental Calculus Calcified or mineralized dental plaque is a common dental disease (Hillson 2008). Dental calculus accumulates at the base of a living plaque deposit that attaches to the tooth surface (Figure 8). Levels of calculus and location are affected by oral hygiene habits, diet (carbohydrate consumption), and systemic disease (Hillson 2008; White 1997). We report the frequencies of dental calculus observed on the tooth crown and cemento-enamel junction on a graded scale ranging from absence, slight, moderate, and marked (Brothwell 1981: Figure 6.14b). Dental Attrition Dental attrition results from tooth-on-tooth contact, often producing wear facets on the occlusal and proximal surfaces of the teeth in function (Hillson 1996, 2008). In this study we record occlusal attrition on an absent, slight (enamel wear), moderate (dentin exposure), and marked (pulp exposure) gradient.

Results

Stature Estimates of living stature for 13 males and eight females from the NKLE site using Asian, Chinese, and non-ethnic formulae are presented in Tables 3 and 4. Here, we provide a summary of adult statures using non-ethnic formulae (Sjøvold 1990) only. Male statures range from 154.3 cm to 167.8 cm, the mean is 160.1 cm. Female statures range from 146.1 cm to 162.4 cm, with a mean of 155.2 cm. Comparisons of the statures of these Neolithic skeletons with other skeletal series within and outside Taiwan as well as with indigenous groups living in Taiwan are discussed later. Cribra Orbitalia (CO)

Although there is a slightly higher frequency of CO in females (40.0%) than males (25.0%), this frequency is not statistically significant. The combined frequency of CO in the NKLE skeletons is 30.8% (Table 5). Dental Enamel Hypoplasia While not statistically significant, a higher frequency of LEH is observed in adult male teeth (57.1%) than in adult female teeth (44.1%). The frequency of LEH in the NKLE maxillary and mandibular incisor and canine teeth, sexes combined, is 51.3% (Table 5). Tooth Ablation Tooth ablation was observed in all adult (14 males, 8 females) and one male subadult skeleton from NKLE (Pietrusewsky et al. 2014b). The overall frequency of tooth ablation was 24.7% in males and 23.7% in females (Table 6). Only the maxillary teeth were targeted for removal. The most common pattern of tooth ablation in the NKLE skeletons was the bilateral ablation of the maxillary lateral incisor [I2] and canine [C] teeth [CI2⏐I2C], occurring in 8/8 (100%) of the observable females and in 12/14 (85.7%) of the males. In addition to this pattern of ablation, the right P3 was observed to be ablated in one young adult male [P3CI2⏐I2C]. Another pattern of ablation, observed in a middle-age male, was the bilateral ablation of maxillary lateral incisors [I2⏐I2]. In all cases, the sockets for the ablated teeth exhibit complete healing of the alveolar bone. Three adults

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(two females and one male) exhibit root remnants of ablated teeth suggesting the ablation of the crown portion of these teeth occurred well before death. Betel Staining The cultural practice of chewing betel (Areca) nut influences many of the dental health indicators and has proven to complicate interpretations of oral health archaeological remains. As mentioned, chewing the betel quid (the nut, leaf and possibly lime) is cariostatic and also affects the prevalence of calculus, attrition, and periodontal disease. A statistically higher frequency of betel stained teeth was observed in the male (75.5%) compared to the female (52.3%) skeletons from NKLE (Table 6). Because the staining was observed in all teeth, the discoloration is likely due to chewing betel nut rather than the intentional application of betel for aesthetic purposes. Dental Pathology Antemortem Tooth Loss (AMTL) Very similar frequencies of AMTL were observed in males (0.2%) and females (0.4%). The overall frequency of AMTL in the NKLE dentitions, sexes combined, is extremely low (0.3%) (Table 6). Dental Caries There is no significant difference in the frequency of caries observed in males (1.7%) and females (2.1%) (Table 6). The overall frequency of carious teeth from NKLE, sexes combined, is low (1.9%). Alveolar Resorption No significant difference is reported for the frequency of resorption in adult male (10.4%) and female (6.1%) alveoli from NKLE. The overall frequency of advanced (moderate and marked expressions) levels of resorption of the alveolus in the NKLE series is 8.8% (Table 6). Alveolar Defects There is no significant difference in the frequency of alveolar defects in the adult male sockets (1.3%) and female sockets (0.5%). The frequency of alveolar defects, or periapical inflammation, in the NKLE series is 1.0% of tooth sockets (Table 6). Dental Calculus A significantly higher frequency of calculus was observed in female teeth (13.9%) compared to male teeth (4.4 %). The overall frequency of advanced (moderate and marked) calculus build-up in the NKLE teeth is 8.3% (Table 6). Dental Attrition Advanced attrition was significantly higher in males (11.7%) than females (2.2%) (Table 6). The overall incidence of advanced dental attrition in the NKLE adults is 7.9%.

Discussion

Differences between Sexes As expected, NKLE adult males, on average, were approximately 5 cm taller than adult females from this site. No significant differences between the sexes were found for two indicators of early childhood health, LEH and CO. While not significant, a slightly higher frequency of linear enamel hypoplasia was observed in NKLE males compared to females. Among the conditions associated with LEH are malnutrition, acute and chronic infection, trauma, and hereditary anomalies (Goodman and Rose 1991).

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Likewise, the relatively high frequency of CO observed in these remains suggests the existence of childhood physiological stress, conditions associated with a variety of anemic conditions and other disorders such as rickets and scurvy as well as nutritional deficiencies associated with parasitic infections and diarrheal disease (Walker 1986; Walker et al. 2009).

Overall, the dental health of the NKLE skeletons is good as demonstrated by the relatively low frequencies of dental pathology in these remains. With the exception of betel staining, dental attrition, and dental calculus, no statistically significant differences were observed between males and females.

The significantly higher frequency of advanced dental attrition observed in the males correlates with significantly higher frequency of betel staining in males. Although not significant, the slightly lower rates of caries and AMTL observed in males compares favorably with the known cariostatic effects of chewing betel nut. The significantly lower frequency of advanced dental calculus observed in males may support the plaque prevention properties of betel nut and/or reflect differences in hygiene and diet of these early inhabitants. Overall, there are relatively few differences in the health of these early Neolithic inhabitants with respect to sex. Early Neolithic and Iron Age Taiwan Comparisons

Based on previous research in bioarchaeology, it is predicted that subsistence economies of Iron Age Taiwan will be associated with an increase in systematic stress and certain dental indicators of health when compared to the early Neolithic communities in Taiwan. Thus, frequencies of childhood stress (CO and LEH), and dental pathology (e.g., dental caries, AMTL, alveolar defects, and dental calculus) should increase from early Neolithic to later Iron Age Taiwan. Likewise, is it expected that adult stature should decrease with the transition to more intensified agriculture.

Contrary to expectations, adult stature estimates are taller in the Iron Age skeletons from SSH compared to the Neolithic NKLE skeletons, a difference that is almost statistically significant for males and is statistically significant for females (Table 7). As expected, CO is slightly greater in the Iron Age series than in the skeletons from NKLE, although the difference is not significant (Table 8). Contrary to expectations, the frequency of LEH is significantly lower in the Iron Age skeletons (Table 9). The expected increase in AMTL, caries, and alveolar defects in the Iron Age skeletons is not confirmed, none of the frequencies was significantly different in the two skeletal series (Table 11). Only the frequencies of alveolar resorption and dental calculus, indicators of periodontal disease, increased in the Iron Age series as expected (Table 12). The differences in these two indicators may also reflect differences in oral hygiene, diet, and the prophylactic effects of chewing betel nut between the two series. Finally, as predicted, advanced dental attrition was significantly less in the Iron Age skeletons, a change that is consistent with a transition from a more abrasive or fibrous diet and/or use of the teeth as tools in the early Neolithic to an increased reliance on softer, more processed foods and metal for tools in the Iron Age.

Of the nine indicators of systemic stress examined, only two were found to change as expected, suggesting there is no temporal increase in reliance on cereals or change in emphasis between millet and rice; rather, the subsistence base remained very broad, including marine, riverine and terrestrial resources. If anything, there was a decline in childhood stress over time, that may be linked to improved food processing techniques (e.g., boiling) because of improved ceramic vessels, better weaning foods, resulting in less weaning stress and less contamination of food in later Iron Age Taiwan.

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Significant differences between the Neolithic and Iron Age skeletons were observed for two different kinds of dental modification. Betel-stained teeth and tooth ablation observed in the NKLE skeletons, were either completely absent or rare in the SSH skeletons. In contrast to these mostly minor differences, which may be attributable to differences in diet and oral hygiene, is the overall similarity in dental health between the Neolithic skeletons from NKLE and the later Iron Age skeletons from SSH. The archaeological evidence suggests that Taiwan’s earliest Neolithic inhabitants, like those who settled at the NKLE site in southwestern Taiwan, possessed well-developed coastal adaptive strategies (Li 2013). While the NKLE settlers were among Taiwan’s first farmers, which included the cultivation of millet, their mode of subsistence was heavily dependent on intensive marine exploitation and the hunting of terrestrial mammals (dog, deer and wild boar) and birds (Li 2013; Tsang 1995). Likewise, although there is evidence that the diet of the Iron Age settlers at SSH, a site located near the sea and rivers, included rice, the collection of shellfish, and terrestrial hunting likely constituted an important part of their subsistence base (Tsang 2000). Despite the fact that the inhabitants of the NKLE and SSH sites are separated by at least 3000 years and assigned to different cultural periods in Taiwan, both appear to have relied heavily on marine resources in their subsistence strategies. The similarities in dental health for the early settlers at these sites, demonstrated in the present study, agree with this interpretation. Regional Comparisons

Obtaining comparative data recorded in skeletal series from outside of Taiwan that overlap temporally and spatially with those from NKLE and SSH has proven difficult. Part of the reason for the relative lack of data is that there have been relatively few studies that focus on indicators of health in skeletal series in surrounding regions, especially from directly across the Taiwan Strait in southern China. Another problem is the lack of standardization in reporting data in the studies conducted by different researchers. Skeletal series from Southeast Asia, Japan, and China that provide comparative data are given in Table 2. Generally, these sites have similar subsistence economies and temporally overlap the NKLE and SSH skeletons. Stature With the exception of some of the indigenous groups in Taiwan and the Jomon skeletons, the average adult male stature for NKLE generally falls below the average statures for most of the skeletal series from China and Southeast Asia (Table 7). Comparing the average statures of Taiwan’s indigenous groups, the NKLE adult male stature is most similar to Atayal, Saisiyat, and Puyuma groups (see map in Figure 1 for locations of these groups). In contrast, the average stature of NKLE adult females is comparable or exceeds female adult stature for many of the skeletal series from East and Southeast Asia. Among the indigenous groups, the stature of NKLE females is most similar to the Amis group. Cribra orbitalia There is considerable variability in the prevalence of cribra orbitalia (CO) among the comparative series (Table 8). Some of the highest frequencies of this early childhood indicator are those reported for early skeletal series from Vietnam and some of the lowest frequencies are those for Jomon, Yayoi, Northeast Thailand and early and middle Yangshao skeletons from China. The frequencies of CO in the NKLE skeletons are most similar to those reported for the skeletal series from China and Thailand. Significantly

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higher frequencies of CO are reported for Khok Phanom Di (Thailand) and all the Vietnamese series. LEH

Although the sample size is small, the moderately high frequency (51.3%) of LEH recorded in the NKLE teeth is similar to frequencies reported for Khok Phanom Di, Jomon, and Man Bac (Vietnam) series (Table 9), series whose subsistence base includes foraging, hunting and fishing, as well as plant cultivation. Significantly lower frequencies of LEH are reported for most of the remaining series including Yayoi agriculturalists, Non Nok Tha, Ban Chiang, and the Vat Komnou skeletal series. The relatively high frequency of this indicator suggests that the early Neolithic inhabitants in this region experienced relatively high levels of physiological stress during childhood. Dental Modification and Dental Pathology The distinctive pattern of tooth ablation observed in the NKLE skeletons has been reported in Neolithic (e.g., Lien 1987, 1989), Iron Age (e.g., Chiu 2010), and modern Indigenous skeletons from Taiwan (e.g., Nakahashi 2008). Likewise, tooth ablation has been reported in skeletons of archaeological sites in Mainland China, Japan, and Southeast Asia (e.g., Han and Nakahashi 1996). However, the bilateral removal of the maxillary canines and lateral incisors, the pattern observed in the NKLE skeletons, is relatively infrequent in these comparative series. The finding that the maxillary lateral incisor [I2⏐], but not the canine, in one subadult male (approximately 15 years of age) not included in this analysis supports a rite of passage explanation for this observed behavior. Although the cultural practice of chewing of betel nut (Areca catechu) is fairly prevalent in contemporary groups throughout Southeast Asia, Taiwan, and the Pacific, the lack of comparative data in archaeological remains from these regions hinders comparisons. Limited information (Table 10) indicates that betel stained teeth are common in the NKLE and Vat Komnou (Cambodia) skeletons but relatively infrequent in the Ban Chiang (Thailand) and Shisanhang skeletal series. The betel staining observed in the Bronze Age site Nui Nap in Thanh Hoa Province in Vietnam is likely due to the intentional application of betel nut residues to the teeth rather than as a result of chewing betel nut (Oxenham Locher et al. 2002). Unlike NKLE, the frequency of betel stained teeth in these skeletons is very similar in both sexes. The prevalences of oral/dental infection; AMTL, dental caries, and alveolar defects for the NKLE and SSH series are among the lowest reported (Table 11), suggesting generally good dental health for these prehistoric inhabitants of Taiwan. Significantly higher frequencies of AMTL are observed for the majority of the Neolithic and early metal age skeletons from Thailand, Vietnam, and Vat Komnou (Cambodia). A comparably high frequency of AMTL is observed for the Phum Snay site in Cambodia. The frequencies of carious lesions in the NKLE and SSH series, while among the lowest reported, are not significantly different from those reported for many of the Neolithic and Metal age series from Thailand (early Non Nok Tha) and Vietnam (Table 11). Significantly higher frequencies of carious lesions are reported for all remaining series. Equally low frequencies of alveolar defects observed in the NKLE and SSH series are reported for several Neolithic and Metal Age series from Thailand, Vietnam, and Cambodia. Significantly higher frequencies for this dental indicator are reported for some of the skeletal series from Thailand (e.g., Ban Chiang, Khok Phanom Di, late Non Nok Tha, and Noen U-Loke). Differences in diet (particularly carbohydrate content) and subsistence patterns have been linked to caries frequencies. Caries are generally rare until the adoption of agriculture (Larsen 1995, 1997; Larsen et al. 1991) and when fermentable carbohydrates

13

from cultivated crops were added to the diet (Hillson 2008). Although Tayles et al. (2000) have made the case that low rates of carious lesions may be associated with rice-dominated agricultural communities (as opposed to other grains such as corn), the unusually low frequencies of dental infections observed in the NKLE series may also reflect a non-agricultural subsistence economy, or a diet low in starches and sugars, and/or the effects of betel chewing. Several authors (e.g., Kelley et al. 1991; Larsen et al. 1991) have reported the cariostatic nature of marine diets. The coastal locations and archaeological evidence for both NKLE and SSH sites indicates that both fishing and exploitation of marine resources were important components of the subsistence patterns for these early inhabitants of Taiwan. It is also of note that the Neolithic and Metal Age inhabitants of Vietnam, described as foragers and agriculturalists whose subsistence base included marine resources, have similarly low frequencies of dental caries and alveolar defects as the early Neolithic and later Iron Age skeletons from Taiwan.

Levels of advanced alveolar resorption observed in the NKLE series are similar to the early and late Non Nok Tha and late Ban Chiang series in northeast Thailand (Table 12). The overall frequency of advanced dental calculus observed in the NKLE series is significantly lower than the frequencies reported for all the skeletons from Thailand, Vietnam, and Cambodia (Table 12). The frequencies of advanced alveolar resorption and dental calculus observed in the NKLE series are among the lowest reported. Higher frequencies of dental calculus have been attributed to a more alkaline oral environment, which may be associated with several factors including the chewing of betel nut (Areca catechu), a diet high in carbohydrates, or poor dental hygiene. With the exception of NKLE (and SSH), all skeletal series have somewhat higher frequencies of advanced dental attrition (Table 12). In addition to diet, cultural practices such as betel nut chewing, and the use of teeth in task-related activities may contribute to advanced dental attrition rates. Comparisons of the NKLE and SSH skeletons from Taiwan show these two skeletal series within the range of other coastal communities in the region with good dental health but with evidence of more childhood stress. Additional skeletal evidence is necessary to examine the possible causes of this, including evidence for infectious disease and anemia.

Conclusions

The average stature of Taiwan’s early Neolithic people from the NKLE site is estimated at 160 cm for adult males and 155 cm for adult females. The frequencies of two childhood indicators of health, CO and LEH, are similar in males and females. However, the relatively high frequency of these two indicators suggests the early Neolithic inhabitants experienced considerable physiological stress as children, most likely attributable to diseases and malnutrition. The most common pattern of tooth ablation, most likely associated with an age-based achievement, involved the removal the maxillary lateral incisors and canines on both sides of the jaw. This cultural behavior is expressed equally in males and females from the NKLE site.

Compared to females, significantly higher frequencies of betel-stained teeth and marked attrition are observed in the males from NKLE. Calcified dental plaque is significantly lower in males than females. No significant differences are found for the remaining dental pathologies. The chewing of betel nut and hygiene may account for some of the differences observed.

The prediction that there was a decline in health corresponding to the transition from early Neolithic to Iron Age in Taiwan is generally not supported by the results of

14

this study. Only two indicators, dental calculus and alveolar resorption, are found to be significantly greater in the Iron Age skeletons from SSH and, as predicted dental attrition is significantly less in the SSH skeletons. Contrary to expectations, adult stature is greater in the Iron Age skeletons and LEH is significantly lower compared to the early Neolithic skeletons from NKLE. For the remaining indicators, there are no significant differences between the early Neolithic and Iron age. While tooth ablation and betel stained teeth are common in the NKLE skeletons, they are rare or completely nonexistent in the skeletons form the SSH site. The similarities in dental health between the early Neolithic skeletons from NKLE and the later Iron Age skeletons from SSH, suggest they shared a similar subsistence economy, one that was likely based on fishing and the hunting and gathering of marine and terrestrial resources.

Comparisons of the NKLE skeletons with those from surrounding regions demonstrate that the average stature of NKLE males generally falls below many of the comparative series while the average stature of NKLE females is generally greater than most of the series from East and southeast Asia. Overall, the stature of NKLE adults is similar to modern indigenous groups in Taiwan.

The frequency of CO in the NKLE skeletons is relatively low and similar to skeletal series from China and Thailand. The relatively high frequency of LEH in the NKLE skeletons suggests that early Neolithic inhabitants of Taiwan may have experienced infectious diseases and/or metabolic disorders during early childhood.

Comparisons of the dental pathology in the prehistoric skeletons from Taiwan with other skeletal series suggest the early inhabitants of Taiwan enjoyed relatively good dental-oral health with low frequencies of dental caries, AMTL, alveolar defect, advanced alveolar resorption, and dental calculus. The NKLE teeth exhibit marked attrition comparable to other early skeletal series from Southeast Asia.

The nearly ubiquitous bilateral removal of the maxillary canines and lateral incisors represents a distinctive cultural behavior for the early Neolithic inhabitants at NKLE.

Further studies of the skeletons from the Nankuanli East site and other sites in the Tainan Science Park will hopefully advance the research reported here.

Acknowledgments Marween Yagin, Graphics Specialist, Center for Instructional Support, University of Hawaii at Manoa is responsible for the map. The National Research Council of Taiwan supported this research.

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List of Tables Table 1. Age and sex distribution for adult Nankuanli East (NKLE) and Shisanhang (SSH) skeletons. Table 2. Comparative adult (>15 years) skeletal series arranged temporally by region. Table 3. Summary of estimated statures for Nankuanli East males. Table 4. Summary of statures for Nankuanli East females. Table 5. Frequencies of LEH and CO in Nankuanli East skeletons. Table 6. Frequencies of tooth ablation, betel staining, and dental pathology in Nankuanli East adult teeth/sockets. Table 7. Stature comparisons. Table 8. Comparison of cribra orbitalia in adults (by individual). Table 9. Regional comparison of LEH in canine and incisor teeth. Table 10. Regional comparison of betel staining. Table 11. Regional comparison of dental infection in teeth/sockets (sides combined). Table 12. Regional comparison of alveolar resorption, calculus, and attrition (sides combined). List of Figures Figure 1. Map showing the approximate locations of the archaeological skeletal series examined in this study and the distribution of some of the present-day Indigenous groups of Taiwan. Figure 2. Inferior view of two burials (interred in the supine position) from the NKLE site showing associated ceramic, shell artifacts, and bone spear point (embedded in right scapula of F3-B3, a middle-aged male). NKLE F3-B4 was determined to be an adolescent male. Figure 3. The skeleton of a young adult female from NKLE (F5B6) showing in-tact long limb bones, which were measured to estimate stature. Figure 4. Lesions attributed to cribra orbitalia are visible in the superior orbits of this young male adult skeleton (F5B40) from NKLE. Figure 5.Multiple LEH are visible in the maxillary central incisors of this young adult male skeleton (E3-B1) from NKLE. Figure 6. Tooth ablation of the maxillary lateral incisors and canines of NKLE E3-B1, a young adult male. Figure 7. Mandibular teeth of NKLE F5-B33, young adult female, showing carious lesions in the left second molar. Many of the teeth are worn to the dentin. Figure 8. The mandibular teeth of NKLE F5-B33 exhibiting dental calculus and alveolar resorption.

Table 1. Age and sex distribution for adult Nankuanli East (NKLE) and Shisanhang (SSH) skeletons. NKLE SSH

Age/Sex Male1 Female2 Total Male3 Female4 Total Young Adult (20 - 35 yrs.)

9 9 18 13 5 18

Middle-aged (35 - 50 yrs.)

5 1 6 2 3 5

Total 14 10 24 15 8 23

                                                                                                               1 The male NKLE burials used in this study: NKLE-D3-B2, NKLE-E3-B1, NKLE-E3-B2, NKLE-E3-B5, NKLE-E4-B4, NKLE-F3-B3, NKLE-F5-B11, NKLE-F5-B19, NKLE-F5-B25, NKLE-F5-B26, NKLE-F5-B27, NKLE-F5-B29, NKLE-F5-B39, NKLE-F5-B40. 2 The female NKLE burials used in this study: NKLE-D3-B3, NKLE-D4-B1, NKLE-E6-B1, NKLE-F3-B1, NKLE-F5-B6, NKLE-F5-B9 (long bones only), NKLE-F5-B14, NKLE-F5-B20, NKLE-F5-B32, NKLE-F5-B33. 3 The male SSH burials used in this study: BM 11, BM18, BM 72, BM 75, BM 76, BM 77,CM 1, CM 11, CM 36, CM 40, EM 17, HM 36, HM 41, HM 47, HM 126. 4 The female SSH burials used in this study: BM 65, BM 86, CM 5, CM 30, HM 40, HM 45, HM 46, HM 78.

Table 2. Comparative adult (>15 years) skeletal series arranged temporally by region. Site Location Approx. Dates

(years BP) References

Taiwan Nankuanli East SW Taiwan 5000 Tsang (2005)

Shisanhang NW Taiwan 1500-1000 Tsang (2000)

China Jihau China 9000-7800 Pechenkina et al. (2013)

Early Yangshao1 China 6900-6000 Pechenkina et al. (2013)

Middle Yangshao2 China 6000-5000 Pechenkina et al. (2013)

Anyang N China 3385-3112 Li (1977)

Japan

Middle/Late Jomon Japan 5000 - 3000 Temple (2007a) Temple & Larsen (2013)

Late to Final Jomon Japan 3000 - 2500 Temple (2007a) Temple & Larsen (2013)

Yayoi Japan 2500 - 1700 Temple & Larsen (2007)

Thailand

Early Non Nok Tha3 NE Thailand 5000-4000 Douglas & Pietrusewsky (2007)

Khok Phanom Di C Thailand 4000 - 3500 Tayles (1999)

Early Ban Chiang NE Thailand 4100 - 2900 Douglas (1996); Douglas & Pietrusewsky (2007); Pietrusewsky & Douglas (2002)

Ban Lum Khao NE Thailand 3400-2500 Domett (2001, 2004); Domett & Tayles (2006)

Late Non Nok Tha4 NE Thailand 4100-1800 Douglas & Pietrusewsky (2007) Late Ban Chiang Thailand 2900-1800 Douglas (1996); Douglas & Pietrusewsky (2007);

Pietrusewsky & Douglas (2002) Nong Nor C. Thailand 3100-2500 Tayles et al. (1998) Noen U-Loke NE Thailand 2300-1600 Domett & Tayles (2006); Higham (2002)

Vietnam Con Co Ngua N Vietnam 6000-5500 Oxenham (2000); Oxenham Nguyen et al. (2002);

Oxenham et al. 2006) Man Bac N Vietnam 3500-3800 Oxenham et al. (2011)

Metal Age/Dong Son Period

N Vietnam 2500 - 1700 Oxenham (2000); Oxenham Nguyen et al. (2002); Oxenham et al. 2006)

Cambodia Vat Komnou S Cambodia 2500-1500 Ikehara (2010)

Phum Snay NW Cambodia

2350-1800 O’Reilly et al. (2004); Domett & O’Reilly (2009)

1 Skeletons associated with the Early Yangshao Culture from Jiangzhai and Shijia sites in Shaanxi Province, China. 2 Skeletons associated with the Middle Yangshao Culture from Guanjia, Xipo, and Xishan sites in Henan Province, China. 3 Burials from Early Periods 1-3 and Middle Periods 1-3. 4 Burials from Middle Periods 4-8.

Table 3. Summary of estimated statures for Nankuanli East males. Burial Stature (N1 = 15) Regression Side/Bone ID. Age2 (yrs.) Cm. Formula length (cm) F5B25 YA 167.8 ± 4.49 Sjøvold (1990)3 L Femur (450) 169.32 ± 3.8 Trotter and Gleser (1958)4 167.86 ± 3.48 Shao (1989)5 165.9 ± 5.4 Sangvichien et al. (1985, n.d.)6 F5B40 YA 154.2 ± 4.49 Sjøvold (1990) L Femur (400) 158.57 ± 3.8 Trotter and Gleser (1958) 156.36 ± 3.48 Shao (1989) 157.29 ± 5.4 Sangvichien et al. (1985, n.d.) F5B39 MA 159.68 ± 4.49 Sjøvold (1990) L Femur (420) 162.87 ± 3.12 Trotter and Gleser (1958) 160.87 ± 3.8 Shao (1989) 160.75 ± 5.4 Sangvichien et al. (1985, n.d.) F5B29 MA+ 162.39 ± 4.49 Sjøvold (1990) L Femur (430) 165.02 ± 3.8 Trotter and Gleser (1958) 163.23 ± 3.12 Shao (1989) 162.48 ± 5.4 Sangvichien et al. (1985, n.d.) F5B27 MA 166.46 ± 4.49 Sjøvold (1990) L Femur (445) 168.25 ± 3.8 Trotter and Gleser (1958) 166.77 ± 3.12 Shao (1989) 165.07 ± 5.4 Sangvichien et al. (1985, n.d.) F5B19 MA 165.10 ± 4.49 Sjøvold (1990) L Femur (440) 167.17 ± 3.8 Trotter and Gleser (1958) 165.69 ± 3.12 Shao (1989) 164.2 ± 5.4 Sangvichien et al. (1985, n.d.) F5B11 YA+ 157.0 ± 4.49 Sjøvold (1990) L Femur (410) 160.72 ± 3.8 Trotter and Gleser (1958) 158.66 ± 3.48 Shao (1989) 159.02 ± 5.4 Sangvichien et al. (1985, n.d.) F3B3 YA 155.6 ± 4.49 Sjøvold (1990) L Femur (405) 159.65 ± 3.8 Trotter and Gleser (1958) 157.51 ± 3.48 Shao (1989) 158.15 ± 5.4 Sangvichien et al. (1985, n.d.) E4B4 YA 158.33 ± 4.49 Sjøvold (1990) L Femur (415) 161.8 ± 3.8 Trotter and Gleser (1958) 159.81 ± 3.48 Shao (1989) 159.88 ± 5.4 Sangvichien et al. (1985, n.d.)

1 N = number of individuals. 2 Age in years. 3 Sjøvold (1990) non-ethnic formulae: 2.71(Femur) + 45.86 ± 4.49 4 Trotter and Gleser (1958) Asian formulae: 2.15 (Femur) + 72.57 ± 3.8 5 Shao (1989): YA (21-30 years): 2.30 (Femur) + 64.36 ± 3.48; MA (41- 50 yrs): 2.36 (Femur) + 61.75 ± 3.12 6 Sangvichien et al. (1985, n.d.) 1.7289 (Femur) + 88.132 + 5.4

Burial

Stature (N1 = 15) Regression Side/Bone

ID. Age2 (yrs.) Cm. Formula length (cm) E3B5 YA 157.0 ± 4.49 Sjøvold (1990) L Femur (410) 160.72 ± 3.8 Trotter and Gleser (1958) 158.66 ± 3.48 Shao (1989) 159.02 ± 5.4 Sangvichien et al. (1985, n.d.) E3B2 YA+ 155.62 ± 4.49 Sjøvold (1990) L Femur (405) 159.65 ± 3.8 Trotter and Gleser (1958) 157.51 ± 3.48 Shao (1989) 158.15 ± 5.4 Sangvichien et al. (1985, n.d.) E3B1 YA 156.4 ± 4.49 Sjøvold (1990) L Femur (408) 160.29 ± 3.8 Trotter and Gleser (1958) 158.2 ± 3.48 Shao (1989) 158.67 ± 5.4 Sangvichien et al. (1985, n.d.) D3B2 YA+ 165.10 ± 4.49 Sjøvold (1990) L Femur (440) 167.17 ± 3.8 Trotter and Gleser (1958) 165.56 ± 3.48 Shao (1989) 164.2 ± 5.4 Sangvichien et al. (1985, n.d.)

Table 4. Summary of estimated statures for Nankuanli East females. Burial Stature (N1 = 8)

Regression Formula2 Side/Bone length (cm) ID. Age3 (yrs.) Cm. Ft./In. D3B3 YA 162.39 ± 4.49 Sjøvold (1990) L Femur (430) 160.25 ± 3.0 Sangvichien et al. (1985, n.d.) 156.48 Pearson (1899) D4B1 YA 149.65 ± 4.49 Sjøvold (1990) R Femur (383) 148.11 ± 3.0 Sangvichien et al. (1985, n.d.) 147.34 Pearson (1899) F3B1 MA 156.97 ± 4.49 Sjøvold (1990) L Femur (410) 155.08 ± 3.0 Sangvichien et al. (1985, n.d.) 152.59 Pearson (1899) E6B1 YA 156.97 ± 4.49 Sjøvold (1990) L Femur (410) 155.08 ± 3.0 Sangvichien et al. (1985, n.d.) 152.59 Pearson (1899) F5B33 YA+ 152.91 ± 4.49 Sjøvold (1990) L Femur (395) 151.21 ± 3.0 Sangvichien et al. (1985, n.d.) 149.67 Pearson (1899) F5B32 YA 146.13 ± 4.49 Sjøvold (1990) L Femur (370) 144.76 ± 3.0 Sangvichien et al. (1985, n.d.) 144.81 Pearson (1899) F5B20 YA 161.04 ± 4.49 Sjøvold (1990) L Femur (425) 159 ± 3.0 Sangvichien et al. (1985, n.d.) 155.51 Pearson (1899) F5B9 YA 151.55 ± 4.49 Sjøvold (1990) L Femur (390) 149.91 ± 3.0 Sangvichien et al. (1985, n.d.) 148.7 Pearson (1899)

1 N = number of individuals. 2 Sjøvold (1990): 2.71 (Femur) +45.86 ± 4.49; Sangvichien et al. (1985, n.d.): 2.5815 (Femur) + 49.2412 ± 3.0007; Pearson (1899): 1.945(Femur) + 72.844 3 Age in years.

Table 5. Frequencies of LEH and CO in Nankuanli East skeletons. Indicator Male Female Total P Value & A/O1 % A/O2 % A/O2 % Significance2 LEH3 48/84 57.1 30/68 44.1 78/152 51.3 0.1419 CO4 4/16 25.0 4/10 40.0 8/26 30.8 0.6645

1 A/O = affected/observed. 2 Two-tailed P values from Fisher’s Exact Test (FET); values in bold indicate statistical significance. 3 Frequencies of LEH or maxillary and mandibular canine and incisor teeth only. 4 Frequency of cribra per orbit.

Table 6. Frequencies1 of tooth ablation, betel staining, and dental pathology in Nankuanli East adult teeth/sockets. Indicator Male Female Total P Value & A/O2 % A/O2 % A/O2 % Significance3 Tooth ablation4 54/219 24.7 32/135 23.7 86/354 24.23 0.2182 Betel staining 259/343 75.5 102/195 52.3 361/538 67.1 <0.0001 AMTL 1/421 0.2 1/277 0.4 2/698 0.3 1.0000 Caries 6/345 1.7 5/233 2.1 11/578 1.9 0.7628 Alveolar resorption5 26/250 10.4 9/148 6.1 35/398 8.8 0.1991 Alveolar defect 4/314 1.3 1/208 0.5 5/522 1.0 0.6527 Calculus5 14/315 4.4 30/216 13.9 44/529 8.3 0.0002 Attrition6 40/342 11.7 5/231 2.2 45/573 7.9 0.0001

1 Frequencies include right and left sides combined, where applicable. 2 A/O = affected/observed. 3 Two-tailed P values from Fisher’s Exact Test (FET); values in bold indicate statistical significance. 4 Ablation of maxillary teeth. 5 Frequencies of moderate and severe expressions. 6 Advanced (pulp exposure or wear to the root).

Table 7. Stature comparisons. Site Male Female Reference Stature Formulae Taiwan Nankuanli East 160.1 155.2 this study Sjøvold (1990) Nankuanli East 160.9 152.9 this study Sangvichien (1985, n.d.)

Shisanghang 164.5 159.9 Pietrusewsky & Tsang (2003) Sjøvold (1990) Tsou 164.0 153.9 Chai (1967) living stature Atayal 160.1 149.8 Chai (1967) living stature Saisiyat 158.7 150.4 Chai (1967) living stature Bunun 157.2 146.2 Chai (1967) living stature Paiwan 156.6 148.0 Chai (1967) living stature Amis 164.6 155.9 Chai (1967) living stature Puyuma 160.0 149.6 Chai (1967) living stature Rukai 158.2 148.7 Chai (1967) living stature China Yangshao Neolithic 169.3 153.9 Pechenkina et al. (2013) Trotter & Gleser 1958 (M); Pearson 1899 (F)

Longshan Neolthic 167.2 151.8 Pechenkina et al. (2013) Trotter & Gleser 1958 (M); Pearson 1899 (F)

Western Zhou 166.5 150.1 Pechenkina et al. (2013) Trotter & Gleser 1958 (M); Pearson 1899 (F)

Han Dynasty 165.8 148.1 Pechenkina et al. (2013) Trotter & Gleser 1958 (M); Pearson 1899 (F)

Japan Middle Jomon 160.8 152.0 Pechenkina et al. (2013) Fuji (1960) Late/Final Jomon 156.3 149.2 Pechenkina et al. (2013) Fuji (1960) All Jomon 157.6 149.7 Temple (2007a) Fuji (1960) S Honshu Yayoi 169.8 159.6 Temple (2007a) Stevenson (1929) N Kyushu Yayoi 165.6 158.0 Temple (2007a) Stevenson (1929) Thailand Early Non Nok Tha 164.7 152.0 Douglas & Pietrusewsky (2007) Sangvichien (1985, n.d.) Khok Phanom Di 162.2 154.3 Tayles (1992) Sangvichien (1985, n.d.) Early Ban Chiang 165.4 153.9 Douglas & Pietrusewsky (2007) Sangvichien (1985, n.d.) Late Non Nok Tha 166.5 155.0 Douglas & Pietrusewsky (2007) Sangvichien (1985, n.d.) Nong Nor 167.2 156.1 Tayles et al. (1998) Sangvichien (1985, n.d.) Ban Lum Khao 164.7 154.7 Domett & Tayles (2006) Sangvichien (1985, n.d.) Late Ban Chiang 166.0 154.4 Douglas & Pietrusewsky (2007) Sangvichien (1985, n.d.) Ban Na Di 167.1 156.2 Pietrusewsky & Douglas (2002) Sangvichien (1985, n.d.) Noen U-Loke 169.3 154.6 Domett & Tayles (2006) Sangvichien (1985, n.d.) Vietnam Con Co Ngua 170.5 158.1 Oxenham (2000) Sjøvold (1990) Man Bac 161.4 154.0 Oxenham et al. (2011) Sangvichien (1985) Metal Age/Dong Son 166.1 152.2 Oxenham (2000) Sjøvold (1990) Cambodia Vat Komnou 165.3 154.8 Ikehara (2010) Sangvichien (1985, n.d.) Phum Snay 167.7 161.1 Domett & O’Reilly (2009) Sangvichien (1985, n.d.)

Table 8. Comparison of cribra orbitalia in adults (by individual).

Sample A/O % P Value &

Significance1

Taiwan

Nankuanli East 4/16 25.0

Shisanhang 8/18 44.4 0.2966

China

Jiahu 7/28 25.0 1.0000

Early Yangshao 6/66 9.1 0.0985

Middle Yangshao 13/95 13.7 0.2646

Thailand

Early Non Nok Tha 6/31 19.4 0.7156

Early Ban Chiang 2/23 8.7 0.2053

Late Non Nok Tha 1/27 3.7 0.0556

Khok Phanom Di 31/57 54.4 0.0492

Late Ban Chiang 4/10 40.0 0.6645

Vietnam

Co Con Ngua2 46/57 80.7 0.0008

Man Bac 19/26 73.1 0.0039

Metal Age Vietnam2 39/53 73.6 0.0035

Cambodia

Vat Komnou 5/12 41.7 0.4319

Japan

Jomon3 18/209 8.6 0.0569

Yayoi4 8/91 8.8 0.0790

1 Two-tailed P values from Fisher’s Exact Test (FET) of NKLE frequencies versus other sites; values in bold indicate statistical significance. 2 Data from Oxenham (2000:388) 3 Frequency of CO for Middle, Late, and Final Jomon skeletons from Honshu Island (Temple 2010). 4 Frequency is based on a pooled Yayoi series from northern Kyushu and southwestern Honshu (Temple 2010).

Table 9. Regional comparison of LEH in canine and incisor teeth.

Sample A/O1 % P Value &

Significance2

Taiwan

Nankuanli East 78/152 51.3

Shisanhang 69/186 37.1 0.0111

Japan

Middle to Late Jomon3 187/330 56.7 0.2801

Late to Final Jomon4 97/165 56.9 0.2137

Yayoi5 292/965 30.3 <0.0001

Thailand

Early Non Nok Tha 18/160 11.3 <0.0001

Khok Phanom Di 130/297 43.8 0.1340

Late Non Nok Tha 6/153 3.9 <0.0001

Early Ban Chiang 34/254 13.4 <0.0001

Late Ban Chiang 10/81 12.3 <0.0001

Vietnam

Man Bac 181/279 64.9 0.0074

Cambodia

Vat Komnou 13/120 10.8 <0.0001

1 A/O =affected/observed; % = frequency of occurrence. Includes maxillary and mandibular teeth. 2 Two-tailed P values from Fisher’s Exact Test (FET) of NKLE frequencies versus other sites; values in bold indicate statistical significance. 3 Temple (2007b:Table 5) for Middle to Late Jomon skeletons from the Kitamura and Ota sites on Honshu Island, Japan. 4 Temple (2007b:Table 5) for Late to Final Jomon skeletons from the Tsukumo and Yosekura sites on Honshu Island. 5 Frequency is based on a pooled Yayoi series from Temple (2010).

Table 10. Regional comparison of betel-stained teeth. Male Female Total P Value & A/O1 % A/O % A/O % Significance2 Nankuanli East 259/343 75.5 102/195 52.3 361/538 67.1 Ban Chiang3 47/569 8.3 31/464 6.7 78/1033 7.6 <0.0001 Vat Komnou 131/174 75.3 98/131 74.8 229/305 75.1 0.0155 Nui Nap4 140/275 50.9 120/218 55.0 281/558 50.4 <0.0001 Shisanhang 10/360 2.9 4/145 2.8 14/505 2.8 <0.0001

                                                                                                               1 A/O = Affected/Observed. 2 Two-tailed P values from Fisher’s Exact Test (FET) of NKLE frequencies versus other sites; values in bold indicate statistical significance. 3 Combined frequency for Early and Late burials from the Ban Chiang site (Pietrusewsky & Douglas 2002). 4 Bronze-age site in Vietnam (Oxenham et al. 2002).

Table 11. Regional comparison of dental infection in teeth/sockets (sides combined).

Antemortem Tooth Loss

Carious Lesions Alveolar defect

Sample A/O1 % P Value & Sig.2

A/O % P Value & Sig A/O %

P Value & Sig.

Taiwan

Nankuanli East 2/698 0.3 11/578 1.9 5/522 1.0

Shisanhang 2/633 0.3 1.0000 6/585 1.0 0.2319 4/598 0.7 0.7410

China

Anyang 177/5100 3.5 0.0488

Atayal3 50/378 13.2 <0.0001

Japan

Middle-Late Jomon4 54/1448 3.7 0.0361

Late-Final Jomon 277/2746 10.1 <0.0001

Yayoi, Japan5 185/1725 10.7 <0.0001

Thailand

Early Non Nok Tha 45/900 5.0 <0.0001 11/666 1.7 0.8304 11/767 1.4 0.6101

Khok Phanom Di 183/2047 8.9 <0.0001 139/1282 10.8 <0.0001 122/2047 6.0 <0.0001

Early Ban Chiang 64/1012 6.3 <0.0001 60/826 7.3 <0.0001 44/850 5.2 <0.0001

Ban Lum Khao 59/1138 5.2 <0.0001 39/874 4.5 0.0081 15/1138 1.3 0.6340

Late Non Nok Tha 80/766 10.4 <0.0001 22/536 4.1 0.0340 16/599 2.7 0.0453

Man Bac 19/935 2.0 0.0014 64/581 11.1 <0.0001 13/935 1.4 0.6230

Late Ban Chiang 25/312 8.0 <0.0001 10/235 4.3 0.0840 22/234 9.4 <0.0001

Noen U-Loke 69/1334 5.2 <0.0001 46/956 4.8 0.0032 34/571 6.0 <0.0001

Vietnam

Co Con Ngua6 69/1430 4.8 <0.0001 14/951 1.5 0.5374 22/1430 1.5 0.3892

Metal Age Vietnam 46/1518 3.0 <0.0001 26/1152 2.3 0.7262 39/1518 2.6 0.0343

Cambodia

Vat Komnou 23/647 3.6 <0.0001 25/497 5.0 0.0059 10/523 1.9 0.2982

Phum Snay 2/267 0.7 0.3073 17/188 9.0 <0.0001 3/181 1.7 0.4309

1 A/O =affected/observed teeth. 2 Two-tailed P values from Fisher’s Exact Test (FET) of NKLE frequencies versus other sites; values in bold indicate statistical significance. 3 Frequency from Turner (1979). 4 Calculated from data in Table 6 in Temple (2007b). 5 Calculated from data in Table 4 in Temple and Larsen (2007); Southern Honshu and Northern Kyushu Yayoi only. 6 The number of affected teeth was determined from Oxenham (2000) and Oxenham et al. (2006).

Table 12. Regional comparison of alveolar resorption, calculus, and attrition (sides combined).

Alveolar Resorption1 P Value Calculus1 P Value Attrition2 P Value

Sample A/O3 % & Sig.4 A/O % & Sig. A/O % & Sig.

Taiwan

Nankuanli East 35/398 8.8 44/529 8.3 45/573 7.9

Shisanhang 130/508 25.6 <0.0001 162/560 28.9 <0.0001 4/584 0.7 <0.0001

Thailand

Khok Phanom Di 112/1272 8.8 0.5289

Early Non Nok Tha 39/540 7.2 0.3929 146/497 29.4 <0.0001 35/642 5.5 0.0604 Early Ban Chiang 92/657 14.0 0.0113 158/663 23.8 0.0113 124/754 16.4 <0.0001 Late Non Nok Tha 53/454 11.7 0.1793 138/426 32.4 <0.0001 41/536 7.6 0.9111

Late Ban Chiang 24/239 10.0 0.6722 95/255 37.3 <0.0001 45/307 14.7 0.0023

Ban Lum Khao 104/861 12.1 0.0081

Noen U-Loke 152/977 15.6 <0.0001

Vietnam

Co Con Ngua5 211/951 22.2 <0.0001

Metal Age 168/1152 14.6 0.0003

Cambodia

Vat Komnou 81/461 17.6 0.0002 100/481 20.8 <0.0001 52/498 10.4 0.1651

1 Frequencies of moderate and marked expressions. 2 Advanced attrition (pulp exposure or wear to the roots). 3 A/O =affected/observed. 4 Two-tailed P values from Fisher’s Exact Test (FET) of NKLE frequencies versus other sites; values in bold indicate statistical significance. 5 Data from Oxenham (2000).

500 km

300 mi

N

0 20 40 60 km

TAIWAN

THAILAND

LAOS

PHILIPPINES

I N D O N E S I A

CAMBODIA

VIETNAM

JAPAN

CHINA (Jomon/Yayoi)

Khok Phanom DiNong Nor

Phum SnayVat Komnou

Con Co NguaMan BacDong Song

Ban Lum KhaoNoen U-Loke

Shisanhang Ban ChiangNon Nok Tha

Ban Nadi

Anyang

JiahuYangshao

NankuanliEast

Puyuma

SaisiyatAtayal

BununTsou

Yami

Rukai

Amis

Paiwan

 Figure 2. Inferior view of two burials (interred in the supine position) from the NKLE site showing associated ceramic, shell artifacts, and bone spear point (embedded in right scapula of F3-B3, a middle-aged male). NKLE F3-B4 was determined to be an adolescent male.

 

Figure 3. The skeleton of a young adult female from NKLE (F5B6) showing intact long limb bones, which were measured in-situ to estimate stature.

Figure 4. Lesions attributed to cribra orbitalia are visible in the superior orbits of this young male adult skeleton (F5B40) from NKLE.

 

Figure 6. Tooth ablation of the maxillary lateral incisors and canines of NKLE E3-B1, a young adult male.

Figure 7. Mandibular teeth of NKLE F5-B33, young adult female, showing carious lesions in the left second molar. Many of the teeth are worn to the dentin.

Figure 8. Figure 8. The mandibular teeth of NKLE F5-B33 exhibiting dental calculus and alveolar resorption.

Figure 5. Multiple LEH are visible in the maxillary central incisors of this young adult male skeleton (E3-B1) from NKLE.