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HOMO - Journal of Comparative Human Biology 63 (2012) 94109

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HOMO - Journal of Comparative Human Biology

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Cementum annulations, age estimation, and demographic dynamics in Mid-Holocene foragers of North India
G. Robbins Schug a,, E.T. Brandt b, J.R. Lukacs c
a b c

Department of Anthropology, Appalachian State University, Boone, NC 28608, USA Department of Anthropology, University of Arkansas, Fayetteville, AR 72701, USA Department of Anthropology, University of Oregon, Eugene, OR 97403, USA

a r t i c l e

i n f o

a b s t r a c t
One of the principal problems facing palaeodemography is age estimation in adult skeletons and the centrist tendency that affects many age estimation methods by articially increasing the proportion of individuals in the 3045-year age category. Several recent publications have indicated that cementum annulations are signicantly correlated with known age of extraction or death. This study addresses the question of how demographic dynamics are altered for an archaeological sample when cementum-based age estimates are used as opposed to those obtained via conventional macroscopic methods. Age pyramids were constructed and demographic proles were compared for the early Holocene skeletal population from Damdama (India). The results demonstrate that the use of cementum annulations for age estimation in only a subset of the skeletal sample has a signicant impact on the demographic prole with regard to specic parameters such as mean age at death and life expectancy at birth. This conrms the importance of using cementum annulations to rene age estimates in archaeological samples, which, when combined with a fertility-centred approach to demography, can provide new insights into population dynamics in the past. 2012 Elsevier GmbH. All rights reserved.

Article history: Received 18 April 2008 Accepted 22 January 2012

Introduction Palaeodemographic proles are the foundation of bioarchaeological research, used primarily to compare prehistoric populations (Waldron, 1994) and address bioarchaeological hypotheses (Cohen
Corresponding author. Tel.: +1 828 262 7505; fax: +1 828 262 2982. E-mail address: Robbinsgm@appstate.edu (G. Robbins Schug). 0018-442X/$ see front matter 2012 Elsevier GmbH. All rights reserved. doi:10.1016/j.jchb.2012.01.002

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and Armelagos, 1984; Cohen and Krane-Cramer, 2007; Steckel and Rose, 2002). However, there are aws that severely limit the usefulness of these proles. The most signicant issue may be that the construction of these proles depends upon accurate and precise age and sex estimates for adult skeletons (Bocquet-Appel and Masset, 1982). However, many techniques currently used for ageing adults are imprecise and need improvement (Lovejoy et al., 1985; Mays et al., 1998: 50; Hoppa and Vaupel, 2002). Counts of tooth cementum annulations have demonstrated promise as an accurate method for estimating age in humans. It has been suggested that techniques that provide increased accuracy in age estimation could signicantly alter demographic proles for archaeological samples, but the degree to which that is true has yet to be tested. It is also possible that the broad age categories often in use for demography in archaeological samples could mute the effect of more accurate age estimates. This paper examines the usefulness of cementum annulations for rening age estimates and demographic proles in archaeological samples. Macroscopic methods for estimating age at death in the adult human skeleton are based on degenerative changes. The majority of the methods available rely on observations of a suite of complex morphological characteristics that are not always highly correlated. These traits are scored using discrete categories that limit statistical evaluation and increase rates of intraobserver and interobserver error (Bocquet-Appel and Masset, 1982; Lovejoy et al., 1985; Meindl and Russell, 1998). Age can also be estimated from the dentition, based on macroscopic evaluation of attrition, secondary dentine formation, and hypercementosis. These changes are clearly inuenced by lifestyle and subsistence factors, sometimes to an unknown magnitude in past populations. Histological age estimation methods have frequently been used in an attempt to reduce bias and inaccuracy. Bone remodelling at an adult femoral midshaft can provide an estimate of biological age (Kerley, 1965; Robling and Stout, 2000) until the number of accumulated osteons eventually reaches an asymptote as newly created osteons remove all observable traces of earlier osteons (Chamberlain, 2006: 111). Age can also be estimated based upon the histological evaluation of degenerative processes in the dentition, including root dentine translucency and sclerosis, the formation of secondary dentine, and periodontitis or gingival recession (Alt et al., 1998; Burns and Maples, 1976; Gustafson, 1950; Johanson, 1971; Kashyap and Rao, 1990; Lamendin and Cambray, 1981; Lamendin, 1992; LopezNicolas et al., 1990, 1993, 1996; Lopez-Nicolas and Luna, 1991; Lucy and Pollard, 1995; Lucy et al., 1995, 1996; Maples, 1978; Maples and Rice, 1979; Solheim, 1989, 1990, 1993; Solheim and Sundnes, 1980). Taphonomic processes limit the accuracy of many of these techniques in archaeological samples; for example, dentine translucency age estimates can be exaggerated by post-depositional mineralization inltrating the dentine tubules, causing substantially inated age estimates (Robbins et al., 2004). Cementum is an extracellular matrix composed of calcied collagenous Sharpeys brils, collagen, glycosaminoglycans, proteoglycans, and inorganic hydroxyapatite (Ten Cate, 1998). As the tooth erupts, acellular cementum develops slowly, eventually covering the coronal two-thirds of the root surface. Once the tooth has reached occlusion, subsequent layers of cementum are formed by the cementoblasts residing in the periodontal ligament. The cellular cementum, in the apical portion of the root, is formed more rapidly, being a dynamic tissue that responds to attrition and tooth movement, accumulating mass that performs an adaptive function of keeping the tooth in the occlusal plane. Cementum is formed in annulations, each composed of a pair of bands. While the exact origin of the annulations is unclear, their number appears to correlate with the age (in years) of the root of the tooth. Correlation between root age and apposition is considered strongest in the acellular cementum 1/3 of the distance from the roots apex. The annulations of acellular cementum are better predictors of age than cellular cementum because they are less inuenced by environmental factors and stresses. They are also more easily microscopically resolved (Lieberman and Meadow, 1992) because in this region, the cementum is less compressed than the cementum near the cemento-enamel junction and contains less cellular cementum than the root apex (Charles et al., 1986; Hillson, 1996; Naylor et al., 1985). Cementum is an attractive potential tool for age estimation in human skeletal samples because: (1) acellular cementum is deposited throughout the life span of the tooth and is rarely remodelled or destroyed in vivo in the absence of other pathological conditions (Lieberman and Meadow, 1992; Stein and Corcoran, 1994); (2) cementum occupies a protected location within the alveolus, which makes this delicate material less sensitive to the oral environment and to the effects of diagenesis; (3) since

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teeth are the most taphonomic-resistant material of the human skeleton, cementum analysis allows fragmentary, incomplete, and poorly preserved individuals to be included in a palaeodemographic analysis. The use of cementum annulations in human age estimation is a relatively recent extension of a method used by wildlife biologists to ascertain demographic parameters of faunal assemblages (Stott et al., 1982). Tooth cementum annulations are regularly used in archaeofaunal analysis of archaeological sites to determine not only age at death, but season of death (Burke, 1993; Klevezal, 1996; Lieberman, 1994; Wall-Schefer, 2007), something rarely done with hominin material (Berryman, 2001; Klevezal and Shishlina, 2001; Rau, 2007; Wall-Schefer, 2007). Seasonality is determined by examining band width and colour in the outer-most annulation; thicker bands appear to be deposited during the warmer months of the year. Thinner, darker bands are apparently secreted during the colder months. It is thought that a seasonal variation in nutritional status and growth rate may lead to differences in cell density, collagen bre proportions and/or degree of mineralization (Hillson, 1996; Lieberman, 1993). While cementum annulations have become the most commonly used technique for ageing large animals (Calvert and Ramsay, 1995), research involving human tooth samples has produced mixed results, with correlations of cementum annulations and age at death widely ranging from r = 0.42 (Kasetty et al., 2010) to r = 0.98 (Wittwer-Backofen et al., 2004). Some of the reasons why the results vary so widely include: the type of tooth sample used (archaeological, forensic, chemically-preserved cadaver, or live extraction), sample size and condition (level of damage, pathology, etc.), sample preparation method used (mineralized versus decalcied, stained versus unstained, transverse versus longitudinal cuts), type of technology used (microscopy technique, photography package, use of image projection, increment-counting, and photoshopping software), the experience level of the observer(s), and nally, whether unreadable slides are included in or discarded from the study. Thus far, there have been few cementum annulation studies that have focused specically on human teeth discovered in archaeological contexts (Klevezal and Shishlina, 2001; Wittwer-Backofen, 2008; Wittwer-Backofen et al., 2004; Roksandic et al., 2009). These studies have elucidated the nding that microscopic analysis of archaeological teeth can present particular challenges that are not often encountered with modern teeth. Archaeological cementum may differ in composition due to the longterm effects of weathering and leaching of the organic matrix (Geusa et al., 1999). Stutz (2002) advises that band-like diagenic artefacts may be mistaken for cementum growth layers, and suggests a modied polarized microscopy in order to ensure differentiation between the two. Klevezal and Shishlina (2001) achieved moderate success in estimating season of death in three of ve Bronze Age samples from Russia (circa 2000 BCE). They experienced difculty nding adequate areas of undamaged cementum for analysis, and attributed this to the occurrence of postmortem taphonomic changes. In another study, Roksandic et al. (2009) began with a sample of 116 Serbian teeth dating from 8500 to 5500 BCE. Of this sample, 93 teeth were discarded (80%) because advanced diagenesis prevented adequate readability of the cementum annulations. Alternatively, in a German sample aged 450680 CE, 104 of 121 individuals yielded countable teeth (Wittwer-Backofen et al., 2004), and an overall correlation of 0.72 was obtained between estimated age using the complex method (a suite of macroscopic methods) and ageing via cementum annulations. Materials and methods The sample for this study was excavated from the Indian Mesolithic site of Damdama, which is located near the conuence of the Ganga and Jumna Rivers in the modern state of Uttar Pradesh (Fig. 1). This site, dated to 52505550 BCE (uncalibrated ASM dates), is remarkable because it is one of three Mesolithic sites that is not simply a lithic scatter; it provides deep stratied deposits from which faunal and oral remains, hearths, and human burials were recovered (Lukacs and Pal, 1993). The entire Damdama skeletal collection consists of 46 individuals. Of this group, 18 adults were selected for the present study. Based upon macroscopic age estimates made during the 1992 eld season, these 18 adults were assigned to the following age categories: 9 young adults (1629 years), 7 middle adults (3045 years), and 2 older adults (4660 years) (Lukacs and Pal, in preparation; Robbins et al., 2004; Table 4). Lukacs used the following methods to estimate age at death: dental eruption

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Fig. 1. Map of the Mesolithic site of Damdama on the Gangetic Plain, India.

timing (Moorrees et al., 1963), dental attrition (Scott, 1979; Smith, 1984), the auricular surface (Lovejoy et al., 1985), the pubic symphysis (McKern and Stewart, 1957), cranial sutures (Meindl and Lovejoy, 1985), and epiphyseal suture closure (McKern and Stewart, 1957). Sex of the individuals (10 males and 8 females) was also estimated (Table 4) using the shape of the sciatic notch (Stewart, 1979), the diameter of the humeral and femoral heads (Stewart, 1979), mandibular and cranial morphology (Krogman, 1962: 115), as well as metric and morphological observations of the postcranial skeleton (Steele and Bramblett, 1988). Details on macroscopic age and sex

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Table 1 Dental sample collected from Damdama assemblage. Individual 6b 7 8 11 12 13 15 16a 17 18a 20a 28 30a 30b 32 34 36a 37 Total 18 Teeth available LRP3, URM2 LLtC, URM2, URM3 LLtI2, ULtM3 URM2 ULtM3 LRC, LRM3 URM3 ULtI2 ULtM1, ULtM3 ULtM2 ULtM1 URI1, LRM3 URM3 ULtC, LRM1 ULtI1, ULtP4, LRM3 ULtM2, ULtM3 LRP4 URM3 Total 29

Individuals are numbered using the original grave numbers. U: upper; L: lower; R: right; Lt: left; I1: central incisor; I2: lateral incisor; C: canine; P3: third premolar; P4: fourth premolar; M1: rst molar; M2: second molar; M3: third molar.

estimates are given in the forthcoming monograph on the Damdama skeletal collection (Lukacs and Pal, in preparation). Due to the heavy encrustation of the Damdama skeletons with a calcareous matrix, the selection of teeth for this analysis was largely inuenced by which teeth could be harvested without inicting damage to either the tooth or the supporting alveolar bone. The selection of teeth for histological ageing should ideally be determined by preservation, root morphology, and absence of pathology. Teeth that have remained within the alveolus throughout the depositional period are protected, and thus less likely to suffer damage from diagenesis and other taphonomic processes (Stott et al., 1982). Single rooted teeth are preferred for histological study because they are generally easier to position and section, though there may be few differences in accuracy between anterior and posterior teeth in evaluating root dentine translucency (Drusini et al., 1989) and cementum annulations (Klevezal and Shishlina, 2001; Maples, 1978). In archaeological specimens, where preservation, recovery, taphonomy and diagenesis are issues, single rooted teeth are not always available and posterior teeth may be substituted (Klevezal and Shishlina, 2001). The tooth sample used in this study was initially selected for palaeodietary analysis; hence, this sample includes both anterior and posterior teeth. A total of 29 teeth were harvested from the skeletons with permission from the Department of Ancient History, Culture, and Archaeology, University of Allahabad (Table 1). Eight of the teeth were extracted along with portions of the surrounding alveolar bone (30%), while the remainder are isolated teeth. The sample consists of 4 incisors, 3 canines, 3 premolars and 19 molars (of which 11 are third molars). There are 20 maxillary teeth and 9 mandibular, divided nearly equally between the left side (14) and the right side (15). The teeth did not require xation due to their archaeological derivation. Prior to sectioning, all teeth were analysed for the presence of pathological conditions that may have a signicantly negative impact on age estimation, such as periodontitis, alveolar resorption, passive eruption, adjacent antemortem tooth loss, and root caries that can expose the tooth root to the oral environment. Table 2 provides details on the condition and pathological prole of the teeth used in this sample. Attrition had reached the level of dentine exposure in 17 teeth (67%) and pulp exposure in 7 teeth (26%). There were 13 teeth (48%) which had large interproximal wear facets, and 1 tooth (3.7%) exhibited an antemortem interproximal groove at the cemento-enamel junction, which probably resulted from some habitual idiosyncratic behaviour such as tooth picking (not included

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G. Robbins Schug et al. / HOMO - Journal of Comparative Human Biology 63 (2012) 94109 Table 2 Pathological prole. Ind. # 6b URM2 7 8 11 12 13 15 17 18a 28 30a 30b 32 34 36a 37 TT# Prop. Alvoelar bone URM2 URM2 URM2 URM3 Caries Dentine exp. LRP3, URM2 Pulp exp. Interproxwear LRP3, URM2 LLtI2, ULtM3 LLtC p.m. damage 99

LLt12, ULtM3 URM2 ULtM3 ULtI1, LRM3

LLtI2, ULM3 URM2 ULtI1, LRM3

LRM3 URM3 ULtM2 LRM3 LRM3 LRM1 ULtI1 LRP4

ULtI1 ULtI1, ULtM3

LRM3

ULtP4, LRM3 ULtM2, ULtM3 LRP4 URM3 8 0.3 2 0.07

ULtM2 UR1I, LRM3 URM3 ULtC, LRM1 ULtI1 ULtM2, ULtM3 URM3 17 0.67

URI1

ULtC, LRM1

ULtI1 LRP4 8 0.3

7 0.26

13 0.48

Ind #: number of individuals affected by pathological condition; exp.: exposure; Interprox.: interproximal; p.m.: postmortem; U: upper; L: lower; R: right; Lt: left; I1: central incisor; I2: lateral incisor; C: canine; P3: third premolar; P4: fourth premolar; M1: rst molar; M2: second molar; M3: third molar; TT: total teeth; Prop.: proportion.

in Table 2). There was clear evidence of postmortem damage to the enamel of 8 teeth (30%). Two teeth had occlusal caries (7.4%), but the root was unaffected. In this sample most of the teeth were missing portions of the periodontal ligament, making periodontitis impossible to judge with complete accuracy. Thus, the possible effects of periodontitis on the results of this study are unknown. Estimating age at death from cementum increments requires thin sectioning and histological analysis. Some authors have obtained good resolution of cementum annulations when teeth were decalcied, microtome sectioned, and stained with haematoxylin (Charles et al., 1986, 1989; Condon et al., 1986; Hillson, 1996). However this procedure is often too harsh for archaeological specimens and tends to produce macerated sections in ancient teeth (Charles et al., 1986, 1989; Condon et al., 1986; Klevezal and Shishlina, 2001; Lieberman and Meadow, 1992). For this study, each tooth was embedded in Spurrs resin (mixed for medium hardness) and polymerized for 24 h at 60 , using a protocol similar to that of Stein and Corcoran (1994). The teeth were sectioned in the bucco-lingual plane near the centre of the root, using a Buehler Isomet low speed saw with a diamond-impregnated blade. Longitudinal sections were used because they are considered more conservative for archaeological samples. Due to post-depositional mineralization, the teeth had to be sectioned at 200 m and then ground to a nal thickness of approximately 100 m using a series of sandpapers (grit 200600) and 9 m diamond paste on a Buehler Minimet automatic polisher. (A trial run had indicated that the

Table 3 Dental emergence timing for children in Western Australia (years). Maxilla Central incisor Lateral incisor Canine Third premolar Fourth premolar First molar Second molar Adapted from Halikis (1961). Males 7.10 8.00 10.80 10.00 10.92 6.30 11.50 Females 6.20 7.20 9.80 10.50 11.50 6.10 11.10 Mandible Central incisor Lateral incisor Canine Third premolar Fourth premolar First molar Second molar Males 7.60 8.30 11.60 10.40 11.20 6.40 11.10 Females 6.40 7.70 10.70 11.30 12.30 6.30 11.70

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Fig. 2. Cementum annulation count (individual 30b upper right canine).

sections were easily fractured and occasionally disintegrated when cut at 100 m or less.) The teeth were then stained with 2% Alizarin Red, following the procedure used by Charles et al. (1989). Acellular cementum was evaluated only in regions where the periodontal ligament remained intact. On single-rooted specimens, cementum annulations were counted on the external surface of the root, 1/3 of the distance from the root apex to the cemento-enamel junction, to avoid encounters with cellular cementum (Klevezal and Shishlina, 2001). When posterior teeth were used, root pads (interradicular surfaces of multi-rooted teeth) were avoided. The areas chosen for evaluation were photographed using a Polaroid digital camera (DMC1) at 1600 1200 resolution through a Zeiss 2000C stereomicroscope at 6.5 magnication (Fig. 2). The annulations were counted on the digital images. Counts were recorded by Robbins in a blind evaluation of each section, which had previously been assigned a random number. Each observation was repeated by Robbins after one week to evaluate intra-observer error. Age estimates were obtained by adding the number of annulations to the age at which the tooth generally completes eruption and reaches the occlusal plane among Aboriginal children in western Australia (Halikis, 1961). This source was chosen based upon having a similar subsistence pattern to the study sample (Table 3). A Pearsons correlation was employed to examine the relationship between the macroscopic age estimations and those of the cementum annulations. The sample was divided into younger adults (1629) and middle adults (3055), and a Students t-test for paired samples was used to test for signicant differences between the sets of microscopic and macroscopic age estimates. These estimates were also examined for sexual differences. All statistical analyses were conducted using SPSS (Version 14.0). Results Age estimation Table 4 summarizes the results of age estimation using counts of cementum annulations. After histological processing, 5 teeth (17%) belonging to 4 individuals were unscorable and had to be eliminated from the analysis, decreasing the sample size to 14 individuals and 24 teeth (Table 5). The teeth were not scorable because there were no annulations visible in the cementum. In a Students t-test for paired samples, the level of intra-observer error was not signicant (P = 0.307). The Pearsons correlation between the macroscopic and cementum-based age estimates is r = 0.642, signicant at the = 0.01 level (P = 0.013). The mean difference of 5 years (S.D. = 3.77) between the two sets of estimates was not signicant (P = 0.776). This set of results is also presented in graphic format (Fig. 3). In both males and females, the mean of the cementum estimates was not signicantly different from that of the macroscopic estimates; and the estimates for males were not signicantly different from those of the females. However, ve females (35.7% of the total sample) produced estimates differing from the mean macroscopic estimate by 910 years. The cementum annulations provided estimates 910 years below the macroscopic estimates for three of the females (12, 13, 30a), based on

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G. Robbins Schug et al. / HOMO - Journal of Comparative Human Biology 63 (2012) 94109 Table 4 Estimates for age at death in histological sample. Ind. 6b 7 8 11 12 13 15 16a 17 18a 20a 28 30a 30b 32 34 36a 37 n Mean S.D. min. max. Sex M M M M F F F M F M F M F M M M F F Macroscopic methods (range) 3035 1820 2023 3545 3545 3545 2535 1824 1618 2025 1720 4550 3539 2733 1620 2529 1620 4555 Macroscopic methods (mean) 32.5 19.0 21.5 40.0 40.0 40.0 30.0 21.0 19.0 22.5 18.5 47.5 37.0 30.0 18.0 27.0 18.0 50.0 14.0 26.6 8.3 18.0 40.0 Estimate from cementum annulations 34.0 24.0 27.0 Not scorable 30.0 30.0 27.0 Not scorable 28.0 23.0 28.0 Not scorable 28.0 26.0 18.0 24.0 19.0 Not scorable 14.0 26.1 4.3 18.0 34.0 Absolute difference (yrs) 1.5 5.0 5.5 10.0 10.0 3.0 9.0 0.5 9.5 9.0 4.0 0.0 3.0 1.0 101

5.1 3.8 0.0 10.0

Absolute difference = estimate from cementum annulations mean estimate from macroscopic methods; highest differences in bold. Ind.: individual; yrs: years; M: male, F: female; n: sample size; S.D.: standard deviation; min.: minimum; max.: maximum.

evaluations of the auricular surface and the pubic symphysis. Alternatively, they provided estimates 9 years above the macroscopic estimates for two females (17 and 20a), based on the eruption of the third molar and amount of dental attrition. In the young adult category (1629) the estimates from the cementum annulations were not signicantly different from the macroscopic estimates (P = 0.084) and the correlation between the two

Fig. 3. Macroscopic versus histological age estimates.

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Table 5 Comparison of age estimates by sex. n Females Males n: sample size. 7 7 Mean age (macroscopic) 28.93 24.36 Mean age (cementum) 27.14 25.14 Difference (P-value) 1.79 (0.601) 0.79 (0.586) Correlation (P-value) 0.608 (0.147) 0.767 (0.044)

Table 6 Age estimates for individuals 1629 years old. Individual 7 8 17 18a 20a 32 34 36a Mean Standard deviation t (P-value) Correlation (P-value) Macroscopic range 1820 2023 1618 2025 1720 1620 2529 1620 Macroscopic mean 19.0 21.5 19.0 22.5 18.5 18.0 27.5 18.0 20.50 3.27 Cementum annulations 24.00 27.00 28.00 23.00 28.00 24.00 19.00 24.00 24.63 3.02 2.014 (0.084) 0.693 (0.056)

sets of estimates was statistically signicant (r = 0.693, P = 0.05). However, in the middle adult category (3055) the cementum estimates were signicantly different from the macroscopic estimates (P = 0.02) and the two were not signicantly correlated (r = 0.514, P = 0.30) (Tables 6 and 7). Discussion Age estimation Despite their early Holocene derivation, the proportion of unscorable teeth in the sample (17%) was comparable with studies using modern teeth: 24% (Charles et al., 1986 and Avadhani et al., 2009), 21% (Berryman, 2001) and 23% (Jankauskas et al., 2001; Pundir et al., 2009). When strictly compared to the handful of studies applying this method to archaeological teeth, the 17% rate is relatively low: 14% (Wittwer-Backofen et al., 2004), 40% (Klevezal and Shishlina, 2001) and 80% (Roksandic et al., 2009). It is not surprising that ancient teeth tend to present greater analytical challenges. Roksandic et al.

Table 7 Test for signicant differences in individuals 3055 years old. Individual 6b 12 13 15 30a 30b Mean Standard deviation t (P-value) Correlation (P-value) Macroscopic range 3035 3545 3545 2535 3539 2733 Macroscopic mean 32.5 40.0 40.0 30.0 37.5 30.0 35.00 4.74 Cementum annulations 34.00 30.00 30.00 27.00 31.00 18.00 28.33 5.54 3.192 (0.02) 0.514 (0.30)

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(2009) reported clear evidence of advanced diagenesis, which prevented line counts in the majority of their sample (93 of 116 teeth): broken, faint or erased lines, pitting, and obscuring artefacts. Klevezal and Shishlina (2001) reported large amounts of brittle cementum and non-visible annulations, usually nding undamaged portions in the interroot pads. Additionally, each of the above authors, including those of this study, used slightly different methods of slide preparation in their samples. In this study, the mean difference between the age estimates based on cementum annulations and those based on macroscopic indicators was 4.89 years with a range of 9 years, well within the margin of error for those methods. This mean difference was similar to that obtained (6.510 years) in a previous test of macroscopic versus cementum estimates (Jankauskas et al., 2001). The Pearsons correlation between the two sets of age estimates (r = 0.642), signicant at the = 0.01 level (P = 0.013), is similar to that (r = 0.717) reported by Wittwer-Backofen et al. (2004). The magnitude of the discrepancies in age estimates in this study is not particularly surprising given that (1) the discrepancy is within the margin of error for the macroscopic methods, (2) macroscopic methods used for younger individuals are based on development and methods for middle adult individuals are based on senescence, (3) idiosyncratic and cultural factors may be responsible for different levels of dental attrition or changes to pelvic morphology for males and females in this sample, (4) discrepancies are not uncommon among dental versus skeletal ageing criteria, (5) high inter-observer error rates in the application of the macroscopic methods, (6) the use of discrete categories to describe several complex morphological changes may have led to falsely enhanced dichotomies between old and young individuals, and (7) the expected difference between chronological (cementum annulations) and biological (macroscopic methods) age. Points #3 through #5 (above) could potentially provide explanations for the large age estimation discrepancies between the two methods for ve females in this study. Pelvic indicators of the three middle age adult females aged them 910 years older than the cementum estimates did. Alternatively, dental characteristics of the younger two females aged them 9 years younger than the cementum estimates. Concerning the middle age adult females, the results could be interpreted in light of mobility. Could higher levels of mobility possibly result in an overestimate of age? Possibly, but Cox and Mays (2000) reported that the pelvic age indicators of McKern and Stewart (1957), when tested on a documented archaeological sample, underaged 80% of middle age adult females by at least 10 years. A large number of researchers studying cementum annulations have reported a signicant decrease in accuracy of the method with age, with a tendency to underage middle adult individuals, especially after the age of 40 (Aggarwal et al., 2008; Berryman, 2001; Charles et al., 1986; Dias et al., 2010; Jankauskas et al., 2001; Klevezal and Shishlina, 2001; Kvaal and Solheim, 1995; Miller et al., 1988; Obertova and Francken, 2009; Stein and Corcoran, 1994). It may also be important to note that of the four individuals (of 18) that were dropped from the current analysis due to unscorable teeth, three of the four are middle age adult individuals. While the true reason for the age estimation discrepancy remains uncertain, the most parsimonious explanation is that cementum annulation analysis may have underestimated the age of the three middle age adult females. Two younger females were originally placed in the 1620 age group based upon the estimated eruption time and attrition level of their maxillary molars. These qualities can be difcult to measure accurately in ancient populations due to the level of temporal and cultural divergence from modern populations. Homo sapiens has experienced a general trend towards an overall reduced size of the dentition and its supporting structures (musculature and alveolar bone), reecting changes in diet, food preparation and technology. The most rapid dental reduction occurred between the early and late Upper Palaeolithic periods, particularly in the anterior mandibular teeth and maxillary molars (Hillson, 1996). It is not clear whether eruption patterns were similarly affected by this trend; however, attrition patterns certainly were. Both individuals were aged nine years older than the above methods via cementum annulations. In a plethora of studies, estimates of age using cementum annulations correlate signicantly with known age in individuals under the age of 35 (Aggarwal et al., 2008; Dias et al., 2010; Jankauskas et al., 2001; Kvaal and Solheim, 1995; Meinl et al., 2008; Stein and Corcoran, 1994). In this case, it seems more plausible that these two young females were under-aged by the macroscopic dental observations.

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Table 8 Revised age and sex estimates for the Damdama sample. Age Male n 04 59 1014 1519 2024 2529 3034 3539 4044 4549 5054 5559 6064 Adult Indet. Total 0 0 0 4 7 1 2 1 3 2 0 1 0 3 0 24 prop. 0.17 0.29 0.04 0.08 0.04 0.13 0.08 0.04 0.13 1.00 Female n 0 0 0 1 1 3 5 1 1 1 2 1 0 1 1 18 prop. 0.06 0.06 0.17 0.28 0.06 0.06 0.06 0.11 0.06 0.06 0.06 1.00 Indeterminate Sex n 2 2 4 prop. 1.00 1.00 Total n 2 0 0 5 8 4 7 2 4 3 2 2 0 4 3 46 prop. 0.04 0.11 0.17 0.09 0.15 0.04 0.09 0.07 0.04 0.04 0.09 0.07 1.00 Macro. n 2 0 0 8 5 1 5 3 6 3 1 2 1 6 3 46 prop. 0.04 0.17 0.11 0.02 0.11 0.07 0.13 0.07 0.02 0.04 0.02 0.13 0.07 1.00

Macro.: distribution of ages from the original estimates made using macroscopic methods; n: sample size; prop.: proportion; Indet.: indeterminate.

This study reects a low intra-observer error rate (2.5 years), although the average error rate from several other studies indicates a precision of 6.75 years (Alt et al., 1998; Jankauskas et al., 2001; Pilloud, 2004; Rsing and Kvaal, 1998), a level that is still more precise than most macroscopic methods. Final results and discussion Demographic dynamics Mortality proles from archaeological samples are often biased towards compression of the adult age distribution within the 3045-year age category and underrepresentation of older individuals (60+). Recent approaches to this problem have focused on using Bayesian statistics or other multivariate approaches to increase precision and accuracy of existing macroscopic adult age estimation methods. Counts of cementum annulations have also been proposed as a potential solution for increasing accuracy of age estimation in bioarchaeological samples (Bocquet-Appel, 2008; Hoppa and Vaupel, 2002; Prince and Wittwer-Backofen, 2002; Wittwer-Backofen, 2008; Wittwer-Backofen and Buba, 2002; Wittwer-Backofen et al., 2004). For this analysis adult age estimates were revised using cementum annulations for 14/42 (31.8%) adult individuals included from the Damdama cemetery sample and these are presented along with the age estimates for the entire skeletal series (n = 48) in Table 8. We evaluated the demographic dynamics in our analysis and examined whether or not the prole supports our hypothesis that the use of cementum annulations will signicantly change the age pyramid in both the range of ages represented and the proportion of middle adult individuals. This in turn will lead to signicantly different demographic parameters such as mean age at death, life expectancy at birth, and measures of fertility. The age pyramids for the males and females (Fig. 4) demonstrate that the cementum annulations did allow 2 individuals out of 42 (4.7%) to be moved from the adult age category to a more specic category, increasing the representativeness of the demographic prole. The use of cementum annulations also signicantly altered the age pyramid for males, increasing the proportion of young individuals 2024 and expanding the range of ages represented from 1554 to 1559 years of age. The female age pyramid is also signicantly different, with more females in the 2034-year age range and fewer females in the 3049-year age range. In addition, some female individuals that had previously been aged 5564 years moved into the 5059-year age category. Taking the pooled sample into account,

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Fig. 4. Male (a), female (b), and pooled sex (c) age pyramids by estimation method.

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Table 9 Life table for Damdama (with age estimated using macroscopic methods). Age 04 59 1014 1519 2024 2529 3034 3539 4044 4549 5054 55+ Total Dx 2 0 0 5 8 4 7 2 4 3 2 2 39 dx 5.13 0.00 0.00 12.82 20.51 10.26 17.95 5.13 10.26 7.69 5.13 5.13 100.00 lx 100.00 94.87 94.87 94.87 82.05 61.54 51.28 33.33 28.21 17.95 10.26 5.13 qx 0.05 0.00 0.00 0.14 0.25 0.17 0.35 0.15 0.36 0.43 0.50 1.00 Lx 487.18 474.36 474.36 442.31 358.97 282.05 211.54 153.85 115.38 70.51 38.46 12.82 Tx 3121.79 2634.62 2160.26 1685.90 1243.59 884.62 602.56 391.03 237.18 121.79 51.28 12.82 ex 31.22 27.77 22.77 17.77 15.16 14.38 11.75 11.73 8.41 6.79 5.00 2.50

Dx : raw number of deaths; dx : percentage of total deaths; lx : proportion of survivors; qx : probability of death in age class; Lx : years lived in age class; Tx : years left in life; ex : life expectancy at beginning of age class.

two of our predictions were supported. The use of cementum annulations did shift individuals away from the 3045-year age category and also changed the range of ages represented in the prole. These two results support the hypothesis that the use of cementum annulations does alter the shape of the age pyramid and consequently changes some features of the demographic prole in a signicant manner, even when all of the individuals in a sample cannot be included in the analysis due to preservation, recovery, and other taphonomic issues. When a mortality-centred approach to demography was employed (a life table approach), few signicant differences were noted in the demographic proles. Mean age at death for the 14 individuals evaluated using both macroscopic estimates (mean = 26.64 4.155 years) and cementum annulations (mean = 26.14 2.145 years) did not change signicantly. The median life expectancy (eo ) was not signicantly different either; life expectancy at birth was 31 years for the pooled sex sample using age estimates from macroscopic methods and 28 years according to cementum annulations (Table 9). Our prediction that mean age at death and life expectancy would shift was not supported by the data. However, it has been previously demonstrated that mortality-centred approaches to demography are not the most sensitive indicators of changes to the age pyramid (McCaa, 1998, 2002) and thus we also examined the effect of changes to the age pyramid when a fertility-centred approach to demography is employed. Signicant differences in the demographic proles were obtained when cementum annulations were used for age estimation and we used a fertility-centred approach to demography, or aggregated age ratio analysis following Bocquet-Appel and Paz de Miguel Ibanez (2002). Based on the original macroscopic age estimates, we obtained the following ratio of adults to subadults: 15 P5 = d(5 i) /d(5+) = 0.1951. The age pyramid constructed from a combination of macroscopic methods and counting of cementum annulations produces an estimate of 0.1220 (Fig. 4). With the inclusion of both macroscopic and histological indicators of age at death, the Damdama sample appears to have been in the intermediate range of population fertility (10 < ratio < 35) according to standards set by the Health in the Western Hemisphere study (McCaa, 1998). The ratio of 12.21 corresponds to a calibrated Gross Reproductive Rate (GRR, or number of female offspring per woman) of roughly 2.4 if the life expectancy at birth is 3040 years (GRR is 2.7 if the life expectancy were 20 years). Assuming the higher life expectancy range is correct, this corresponds to a Total Fertility Rate (TFR) of 4.8 offspring per woman. These results demonstrate that the use of cementum annulations in a proportion of a prehistoric cemetery sample (0.30) leads to a signicant difference in the age pyramid and this difference will yield a signicant change in the demographic prole if a fertility-centred model is employed. The results of our analysis indicate support for the hypothesis that problems with age estimation methods signicantly affect the accuracy of palaeodemographic reconstructions (Bocquet-Appel and Masset, 1982). The results suggest support for the hypothesis that fertility-centred approaches to demography are more sensitive than mortality-centred approaches to shifts in the structure of the age pyramid (McCaa, 2002). Finally, our results also demonstrate that changes to the age pyramid

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resulting from the use of cementum annulations are of a magnitude signicant enough to affect the demographic prole. Based on these results, we contend that the counting of cementum annulations is a useful age estimation technique in bioarchaeology because, even when sampling and preservation are not ideal, counting of cementum annulations does signicantly improve the accuracy of adult age estimation and improve palaeodemographic reconstruction.

Acknowledgments This work would not have been possible without the help and monetary support of several organizations and individuals. Firstly, we thank Dr. J.N. Pal and the Department of Ancient History, Culture, and Archaeology at the University of Allahabad. This research was funded in part by grants from the National Geographic Society and the University of Oregon Graduate School. Thanks to Dr. Murray Marks, Jeanne Selker and Joanna Lambert for feedback on the methods and the manuscript.

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