Cementum Annulations, Age Estimation, and Demographic Dynamics of mid-Holocene Forgers from North...

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HOMO - Journal of Comparative Human Biology 63 (2012) 94– 109

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Cementum annulations, age estimation, and demographicdynamics in Mid-Holocene foragers of North India

G. Robbins Schuga,∗, E.T. Brandtb, J.R. Lukacsc

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

a r t i c l e i n f o

Article history:Received 18 April 2008Accepted 22 January 2012

a b s t r a c t

One of the principal problems facing palaeodemography is age esti-mation in adult skeletons and the centrist tendency that affectsmany age estimation methods by artificially increasing the propor-tion of individuals in the 30–45-year age category. Several recentpublications have indicated that cementum annulations are signifi-cantly correlated with known age of extraction or death. This studyaddresses the question of how demographic dynamics are alteredfor an archaeological sample when cementum-based age estimatesare used as opposed to those obtained via conventional macro-scopic methods. Age pyramids were constructed and demographicprofiles were compared for the early Holocene skeletal populationfrom Damdama (India). The results demonstrate that the use ofcementum annulations for age estimation in only a subset of theskeletal sample has a significant impact on the demographic pro-file with regard to specific parameters such as mean age at deathand life expectancy at birth. This confirms the importance of usingcementum annulations to refine age estimates in archaeologicalsamples, which, when combined with a fertility-centred approachto demography, can provide new insights into population dynamicsin the past.

© 2012 Elsevier GmbH. All rights reserved.

Introduction

Palaeodemographic profiles are the foundation of bioarchaeological research, used primarily tocompare prehistoric populations (Waldron, 1994) and address bioarchaeological hypotheses (Cohen

∗ Corresponding author. Tel.: +1 828 262 7505; fax: +1 828 262 2982.E-mail address: [email protected] (G. Robbins Schug).

0018-442X/$ – see front matter © 2012 Elsevier GmbH. All rights reserved.doi:10.1016/j.jchb.2012.01.002


G. Robbins Schug et al. / HOMO - Journal of Comparative Human Biology 63 (2012) 94– 109 95

and Armelagos, 1984; Cohen and Krane-Cramer, 2007; Steckel and Rose, 2002). However, there areflaws that severely limit the usefulness of these profiles. The most significant issue may be that theconstruction of these profiles depends upon accurate and precise age and sex estimates for adultskeletons (Bocquet-Appel and Masset, 1982). However, many techniques currently used for ageingadults are imprecise and need improvement (Lovejoy et al., 1985; Mays et al., 1998: 50; Hoppa andVaupel, 2002). Counts of tooth cementum annulations have demonstrated promise as an accuratemethod for estimating age in humans. It has been suggested that techniques that provide increasedaccuracy in age estimation could significantly alter demographic profiles for archaeological samples,but the degree to which that is true has yet to be tested. It is also possible that the broad age categoriesoften in use for demography in archaeological samples could mute the effect of more accurate ageestimates. This paper examines the usefulness of cementum annulations for refining age estimatesand demographic profiles in archaeological samples.

Macroscopic methods for estimating age at death in the adult human skeleton are based on degen-erative changes. The majority of the methods available rely on observations of a suite of complexmorphological characteristics that are not always highly correlated. These traits are scored using dis-crete categories that limit statistical evaluation and increase rates of intraobserver and interobservererror (Bocquet-Appel and Masset, 1982; Lovejoy et al., 1985; Meindl and Russell, 1998). Age can also beestimated from the dentition, based on macroscopic evaluation of attrition, secondary dentine forma-tion, and hypercementosis. These changes are clearly influenced 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 andinaccuracy. 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 reachesan 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 pro-cesses in the dentition, including root dentine translucency and sclerosis, the formation of secondarydentine, 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; Lopez-Nicolas 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; forexample, dentine translucency age estimates can be exaggerated by post-depositional mineralizationinfiltrating the dentine tubules, causing substantially inflated age estimates (Robbins et al., 2004).

Cementum is an extracellular matrix composed of calcified collagenous Sharpey’s fibrils, colla-gen, glycosaminoglycans, proteoglycans, and inorganic hydroxyapatite (Ten Cate, 1998). As the tootherupts, acellular cementum develops slowly, eventually covering the coronal two-thirds of the rootsurface. Once the tooth has reached occlusion, subsequent layers of cementum are formed by thecementoblasts residing in the periodontal ligament. The cellular cementum, in the apical portion ofthe root, is formed more rapidly, being a dynamic tissue that responds to attrition and tooth move-ment, 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 theannulations is unclear, their number appears to correlate with the age (in years) of the root of thetooth. Correlation between root age and apposition is considered strongest in the acellular cementum1/3 of the distance from the root’s apex. The annulations of acellular cementum are better predictorsof age than cellular cementum because they are less influenced by environmental factors and stresses.They are also more easily microscopically resolved (Lieberman and Meadow, 1992) because in thisregion, the cementum is less compressed than the cementum near the cemento-enamel junction andcontains 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 ordestroyed in vivo in the absence of other pathological conditions (Lieberman and Meadow, 1992; Steinand Corcoran, 1994); (2) cementum occupies a protected location within the alveolus, which makesthis delicate material less sensitive to the oral environment and to the effects of diagenesis; (3) since


96 G. Robbins Schug et al. / HOMO - Journal of Comparative Human Biology 63 (2012) 94– 109

teeth are the most taphonomic-resistant material of the human skeleton, cementum analysis allowsfragmentary, incomplete, and poorly preserved individuals to be included in a palaeodemographicanalysis.

The use of cementum annulations in human age estimation is a relatively recent extension of amethod used by wildlife biologists to ascertain demographic parameters of faunal assemblages (Stottet al., 1982). Tooth cementum annulations are regularly used in archaeofaunal analysis of archaeo-logical sites to determine not only age at death, but season of death (Burke, 1993; Klevezal, 1996;Lieberman, 1994; Wall-Scheffler, 2007), something rarely done with hominin material (Berryman,2001; Klevezal and Shishlina, 2001; Rau, 2007; Wall-Scheffler, 2007). Seasonality is determined byexamining band width and colour in the outer-most annulation; thicker bands appear to be depositedduring the warmer months of the year. Thinner, darker bands are apparently secreted during thecolder months. It is thought that a seasonal variation in nutritional status and growth rate may lead todifferences in cell density, collagen fibre proportions and/or degree of mineralization (Hillson, 1996;Lieberman, 1993).

While cementum annulations have become the most commonly used technique for ageing largeanimals (Calvert and Ramsay, 1995), research involving human tooth samples has produced mixedresults, 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 resultsvary so widely include: the type of tooth sample used (archaeological, forensic, chemically-preservedcadaver, or live extraction), sample size and condition (level of damage, pathology, etc.), samplepreparation method used (mineralized versus decalcified, stained versus unstained, transverse versuslongitudinal cuts), type of technology used (microscopy technique, photography package, use of imageprojection, increment-counting, and photoshopping software), the experience level of the observer(s),and finally, whether “unreadable slides” are included in or discarded from the study.

Thus far, there have been few cementum annulation studies that have focused specifically onhuman 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 findingthat microscopic analysis of archaeological teeth can present particular challenges that are not oftenencountered with modern teeth. Archaeological cementum may differ in composition due to the long-term effects of weathering and leaching of the organic matrix (Geusa et al., 1999). Stutz (2002) advisesthat band-like diagenic artefacts may be mistaken for cementum growth layers, and suggests a mod-ified 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 five Bronze Age sam-ples from Russia (circa 2000 BCE). They experienced difficulty finding adequate areas of undamagedcementum for analysis, and attributed this to the occurrence of postmortem taphonomic changes. Inanother study, Roksandic et al. (2009) began with a sample of 116 Serbian teeth dating from 8500 to5500 BCE. Of this sample, 93 teeth were discarded (80%) because advanced diagenesis prevented ade-quate readability of the cementum annulations. Alternatively, in a German sample aged 450–680 CE,104 of 121 individuals yielded countable teeth (Wittwer-Backofen et al., 2004), and an overall correla-tion of 0.72 was obtained between estimated age using the “complex method” (a suite of macroscopicmethods) and ageing via cementum annulations.

Materials and methods

The sample for this study was excavated from the Indian Mesolithic site of Damdama, which islocated near the confluence of the Ganga and Jumna Rivers in the modern state of Uttar Pradesh(Fig. 1). This site, dated to 5250–5550 BCE (uncalibrated ASM dates), is remarkable because it is oneof three Mesolithic sites that is not simply a lithic scatter; it provides deep stratified deposits fromwhich faunal and floral 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 wereselected for the present study. Based upon macroscopic age estimates made during the 1992 fieldseason, these 18 adults were assigned to the following age categories: 9 young adults (16–29 years), 7middle adults (30–45 years), and 2 older adults (46–60 years) (Lukacs and Pal, in preparation; Robbinset al., 2004; Table 4). Lukacs used the following methods to estimate age at death: dental eruption



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 (Lovejoyet 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 shapeof 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 obser-vations of the postcranial skeleton (Steele and Bramblett, 1988). Details on macroscopic age and sex



Table 1Dental sample collected from Damdama assemblage.

Individual Teeth available

6b LRP3, URM27 LLtC, URM2, URM38 LLtI2, ULtM311 URM212 ULtM313 LRC, LRM315 URM316a ULtI217 ULtM1, ULtM318a ULtM220a ULtM128 URI1, LRM330a URM330b ULtC, LRM132 ULtI1, ULtP4, LRM334 ULtM2, ULtM336a LRP437 URM3

Total18

Total29

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: first molar; M2: second molar; M3: third molar.

estimates are given in the forthcoming monograph on the Damdama skeletal collection (Lukacs andPal, in preparation).

Due to the heavy encrustation of the Damdama skeletons with a calcareous matrix, the selectionof teeth for this analysis was largely influenced by which teeth could be harvested without inflictingdamage to either the tooth or the supporting alveolar bone. The selection of teeth for histologicalageing 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, andthus 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 positionand section, though there may be few differences in accuracy between anterior and posterior teeth inevaluating root dentine translucency (Drusini et al., 1989) and cementum annulations (Klevezal andShishlina, 2001; Maples, 1978). In archaeological specimens, where preservation, recovery, taphon-omy and diagenesis are issues, single rooted teeth are not always available and posterior teeth may besubstituted (Klevezal and Shishlina, 2001). The tooth sample used in this study was initially selectedfor 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 ofAncient History, Culture, and Archaeology, University of Allahabad (Table 1). Eight of the teeth wereextracted along with portions of the surrounding alveolar bone (30%), while the remainder are isolatedteeth. The sample consists of 4 incisors, 3 canines, 3 premolars and 19 molars (of which 11 are thirdmolars). 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 fixation due to their archaeological derivation.

Prior to sectioning, all teeth were analysed for the presence of pathological conditions that mayhave a significantly 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 tothe oral environment. Table 2 provides details on the condition and pathological profile of the teethused in this sample. Attrition had reached the level of dentine exposure in 17 teeth (67%) and pulpexposure in 7 teeth (26%). There were 13 teeth (48%) which had large interproximal wear facets, and 1tooth (3.7%) exhibited an antemortem interproximal groove at the cemento-enamel junction, whichprobably resulted from some habitual idiosyncratic behaviour such as tooth picking (not included



Table 2Pathological profile.

Ind. # Alvoelar bone Caries Dentine exp. Pulp exp. Interproxwear p.m. damage

6b URM2 LRP3, URM2 LRP3,URM2 URM27 URM2 URM2 LLtC8 URM3 LLt12, ULtM3 LLtI2, ULM3 LLtI2, ULtM311 URM2 URM212 ULtM313 ULtI1, LRM3 ULtI1, LRM3 LRM3 ULtI115 URM317 ULtI1, ULtM318a ULtM2 ULtM228 LRM3 UR1I, LRM3 LRM3 URI130a URM3 LRM330b ULtC, LRM1 ULtC, LRM1 LRM132 ULtP4, LRM3 ULtI1 ULtI1 ULtI134 ULtM2, ULtM3 ULtM2, ULtM336a LRP4 LRP4 LRP437 URM3 URM3

TT# 8 2 17 7 13 8Prop. 0.3 0.07 0.67 0.26 0.48 0.3

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: first 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%). Twoteeth had occlusal caries (7.4%), but the root was unaffected. In this sample most of the teeth weremissing portions of the periodontal ligament, making periodontitis impossible to judge with completeaccuracy. 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 histologicalanalysis. Some authors have obtained good resolution of cementum annulations when teeth weredecalcified, microtome sectioned, and stained with haematoxylin (Charles et al., 1986, 1989; Condonet al., 1986; Hillson, 1996). However this procedure is often too harsh for archaeological specimensand 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 wasembedded in Spurr’s resin (mixed for medium hardness) and polymerized for 24 h at 60◦, using aprotocol similar to that of Stein and Corcoran (1994). The teeth were sectioned in the bucco-lingualplane near the centre of the root, using a Buehler Isomet low speed saw with a diamond-impregnatedblade. Longitudinal sections were used because they are considered more conservative for archaeo-logical samples. Due to post-depositional mineralization, the teeth had to be sectioned at 200 �m andthen ground to a final thickness of approximately 100 �m using a series of sandpapers (grit 200–600)and 9 �m diamond paste on a Buehler Minimet automatic polisher. (A trial run had indicated that the

Table 3Dental emergence timing for children in Western Australia (years).

Maxilla Males Females Mandible Males Females

Central incisor 7.10 6.20 Central incisor 7.60 6.40Lateral incisor 8.00 7.20 Lateral incisor 8.30 7.70Canine 10.80 9.80 Canine 11.60 10.70Third premolar 10.00 10.50 Third premolar 10.40 11.30Fourth premolar 10.92 11.50 Fourth premolar 11.20 12.30First molar 6.30 6.10 First molar 6.40 6.30Second molar 11.50 11.10 Second molar 11.10 11.70

Adapted from Halikis (1961).



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 teethwere 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 theroot, 1/3 of the distance from the root apex to the cemento-enamel junction, to avoid encounterswith 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 werephotographed using a Polaroid digital camera (DMC1) at 1600 × 1200 resolution through a Zeiss 2000Cstereomicroscope at 6.5× magnification (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 beenassigned a random number. Each observation was repeated by Robbins after one week to evaluateintra-observer error. Age estimates were obtained by adding the number of annulations to the ageat which the tooth generally completes eruption and reaches the occlusal plane among Aboriginalchildren in western Australia (Halikis, 1961). This source was chosen based upon having a similarsubsistence pattern to the study sample (Table 3).

A Pearson’s correlation was employed to examine the relationship between the macroscopic ageestimations and those of the cementum annulations. The sample was divided into younger adults(16–29) and middle adults (30–55), and a Student’s t-test for paired samples was used to test for sig-nificant differences between the sets of microscopic and macroscopic age estimates. These estimateswere 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. Afterhistological processing, 5 teeth (17%) belonging to 4 individuals were unscorable and had to be elimi-nated from the analysis, decreasing the sample size to 14 individuals and 24 teeth (Table 5). The teethwere not scorable because there were no annulations visible in the cementum. In a Student’s t-testfor paired samples, the level of intra-observer error was not significant (P = 0.307). The Pearson’s cor-relation between the macroscopic and cementum-based age estimates is r = 0.642, significant at the∝ = 0.01 level (P = 0.013). The mean difference of 5 years (S.D. = 3.77) between the two sets of estimateswas not significant (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 significantly differentfrom that of the macroscopic estimates; and the estimates for males were not significantly differentfrom those of the females. However, five females (35.7% of the total sample) produced estimatesdiffering from the mean macroscopic estimate by 9–10 years. The cementum annulations providedestimates 9–10 years below the macroscopic estimates for three of the females (12, 13, 30a), based on



Table 4Estimates for age at death in histological sample.

Ind. Sex Macroscopicmethods (range)

Macroscopicmethods (mean)

Estimate fromcementum annulations

Absolutedifference (yrs)

6b M 30–35 32.5 34.0 1.57 M 18–20 19.0 24.0 5.08 M 20–23 21.5 27.0 5.511 M 35–45 40.0 Not scorable12 F 35–45 40.0 30.0 10.013 F 35–45 40.0 30.0 10.015 F 25–35 30.0 27.0 3.016a M 18–24 21.0 Not scorable17 F 16–18 19.0 28.0 9.018a M 20–25 22.5 23.0 0.520a F 17–20 18.5 28.0 9.528 M 45–50 47.5 Not scorable30a F 35–39 37.0 28.0 9.030b M 27–33 30.0 26.0 4.032 M 16–20 18.0 18.0 0.034 M 25–29 27.0 24.0 3.036a F 16–20 18.0 19.0 1.037 F 45–55 50.0 Not scorable

n 14.0 14.0Mean 26.6 26.1 5.1S.D. 8.3 4.3 3.8min. 18.0 18.0 0.0max. 40.0 34.0 10.0

Absolute difference = estimate from cementum annulations − mean estimate from macroscopic methods; highest differencesin 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 estimates9 years above the macroscopic estimates for two females (17 and 20a), based on the eruption of thethird molar and amount of dental attrition.

In the young adult category (16–29) the estimates from the cementum annulations were not sig-nificantly different from the macroscopic estimates (P = 0.084) and the correlation between the two

Fig. 3. Macroscopic versus histological age estimates.



Table 5Comparison of age estimates by sex.

n Mean age(macroscopic)

Mean age(cementum)

Difference(P-value)

Correlation(P-value)

Females 7 28.93 27.14 1.79 (0.601) 0.608 (0.147)Males 7 24.36 25.14 −0.79 (0.586) 0.767 (0.044)

n: sample size.

Table 6Age estimates for individuals 16–29 years old.

Individual Macroscopic range Macroscopic mean Cementum annulations

7 18–20 19.0 24.008 20–23 21.5 27.0017 16–18 19.0 28.0018a 20–25 22.5 23.0020a 17–20 18.5 28.0032 16–20 18.0 24.0034 25–29 27.5 19.0036a 16–20 18.0 24.00

Mean 20.50 24.63Standard deviation 3.27 3.02t (P-value) 2.014 (0.084)Correlation (P-value) 0.693 (0.056)

sets of estimates was statistically significant (r = 0.693, P = 0.05). However, in the middle adult cate-gory (30–55) the cementum estimates were significantly different from the macroscopic estimates(P = 0.02) and the two were not significantly 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 tothe 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 7Test for significant differences in individuals 30–55 years old.

Individual Macroscopic range Macroscopic mean Cementum annulations

6b 30–35 32.5 34.0012 35–45 40.0 30.0013 35–45 40.0 30.0015 25–35 30.0 27.0030a 35–39 37.5 31.0030b 27–33 30.0 18.00

Mean 35.00 28.33Standard deviation 4.74 5.54t (P-value) 3.192 (0.02)Correlation (P-value) 0.514 (0.30)



(2009) reported clear evidence of advanced diagenesis, which prevented line counts in the major-ity 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 annu-lations, usually finding undamaged portions in the interroot pads. Additionally, each of the aboveauthors, including those of this study, used slightly different methods of slide preparation in theirsamples.

In this study, the mean difference between the age estimates based on cementum annulationsand those based on macroscopic indicators was 4.89 years with a range of ±9 years, well within themargin of error for those methods. This mean difference was similar to that obtained (6.5–10 years)in a previous test of macroscopic versus cementum estimates (Jankauskas et al., 2001). The Pearson’scorrelation between the two sets of age estimates (r = 0.642), significant 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 giventhat (1) the discrepancy is within the margin of error for the macroscopic methods, (2) macroscopicmethods used for younger individuals are based on development and methods for middle adult indi-viduals are based on senescence, (3) idiosyncratic and cultural factors may be responsible for differentlevels 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-observererror rates in the application of the macroscopic methods, (6) the use of discrete categories to describeseveral complex morphological changes may have led to falsely enhanced dichotomies between “old”and “young” individuals, and (7) the expected difference between chronological (cementum annula-tions) and biological (macroscopic methods) age.

Points #3 through #5 (above) could potentially provide explanations for the large age estimationdiscrepancies between the two methods for five females in this study. Pelvic indicators of the threemiddle age adult females aged them 9–10 years older than the cementum estimates did. Alternatively,dental characteristics of the younger two females aged them 9 years younger than the cementumestimates.

Concerning the middle age adult females, the results could be interpreted in light of mobility. Couldhigher 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 documentedarchaeological sample, underaged 80% of middle age adult females by at least 10 years. A large numberof researchers studying cementum annulations have reported a significant decrease in accuracy of themethod 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 ageadult individuals. While the true reason for the age estimation discrepancy remains uncertain, themost parsimonious explanation is that cementum annulation analysis may have underestimated theage of the three middle age adult females.

Two younger females were originally placed in the 16–20 age group based upon the estimatederuption time and attrition level of their maxillary molars. These qualities can be difficult to measureaccurately in ancient populations due to the level of temporal and cultural divergence from modernpopulations. Homo sapiens has experienced a general trend towards an overall reduced size of thedentition and its supporting structures (musculature and alveolar bone), reflecting changes in diet,food preparation and technology. The most rapid dental reduction occurred between the early andlate 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 methodsvia cementum annulations. In a plethora of studies, estimates of age using cementum annulationscorrelate significantly with known age in individuals under the age of 35 (Aggarwal et al., 2008; Diaset 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 themacroscopic dental observations.



Table 8Revised age and sex estimates for the Damdama sample.

Age Male Female Indeterminate Sex Total Macro.

n prop. n prop. n prop. n prop. n prop.

0–4 0 – 0 – 2 1.00 2 0.04 2 0.045–9 0 – 0 – – – 0 – 0 –10–14 0 – 0 – – – 0 – 0 –15–19 4 0.17 1 0.06 – – 5 0.11 8 0.1720–24 7 0.29 1 0.06 – – 8 0.17 5 0.1125–29 1 0.04 3 0.17 – – 4 0.09 1 0.0230–34 2 0.08 5 0.28 – – 7 0.15 5 0.1135–39 1 0.04 1 0.06 – – 2 0.04 3 0.0740–44 3 0.13 1 0.06 – – 4 0.09 6 0.1345–49 2 0.08 1 0.06 – – 3 0.07 3 0.0750–54 0 – 2 0.11 – – 2 0.04 1 0.0255–59 1 0.04 1 0.06 – – 2 0.04 2 0.0460–64 0 – 0 – – – 0 – 1 0.02Adult 3 0.13 1 0.06 – – 4 0.09 6 0.13Indet. 0 – 1 0.06 2 – 3 0.07 3 0.07

Total 24 1.00 18 1.00 4 1.00 46 1.00 46 1.00

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

This study reflects a low intra-observer error rate (2.5 years), although the average error rate fromseveral other studies indicates a precision of ±6.75 years (Alt et al., 1998; Jankauskas et al., 2001;Pilloud, 2004; Rösing and Kvaal, 1998), a level that is still more precise than most macroscopic methods.

Final results and discussion

Demographic dynamics

Mortality profiles from archaeological samples are often biased towards compression of the adultage distribution within the 30–45-year age category and underrepresentation of older individuals(60+). Recent approaches to this problem have focused on using Bayesian statistics or other multi-variate approaches to increase precision and accuracy of existing macroscopic adult age estimationmethods. Counts of cementum annulations have also been proposed as a potential solution for increas-ing 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 withthe age estimates for the entire skeletal series (n = 48) in Table 8. We evaluated the demographicdynamics in our analysis and examined whether or not the profile supports our hypothesis that theuse of cementum annulations will significantly change the age pyramid in both the range of agesrepresented and the proportion of middle adult individuals. This in turn will lead to significantlydifferent demographic parameters such as mean age at death, life expectancy at birth, and measuresof fertility.

The age pyramids for the males and females (Fig. 4) demonstrate that the cementum annulationsdid allow 2 individuals out of 42 (4.7%) to be moved from the “adult” age category to a more specific cat-egory, increasing the representativeness of the demographic profile. The use of cementum annulationsalso significantly altered the age pyramid for males, increasing the proportion of young individuals20–24 and expanding the range of ages represented from 15–54 to 15–59 years of age. The femaleage pyramid is also significantly different, with more females in the 20–34-year age range and fewerfemales in the 30–49-year age range. In addition, some female individuals that had previously beenaged 55–64 years moved into the 50–59-year age category. Taking the pooled sample into account,



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



Table 9Life table for Damdama (with age estimated using macroscopic methods).

Age Dx dx lx qx Lx Tx ex

0–4 2 5.13 100.00 0.05 487.18 3121.79 31.225–9 0 0.00 94.87 0.00 474.36 2634.62 27.7710–14 0 0.00 94.87 0.00 474.36 2160.26 22.7715–19 5 12.82 94.87 0.14 442.31 1685.90 17.7720–24 8 20.51 82.05 0.25 358.97 1243.59 15.1625–29 4 10.26 61.54 0.17 282.05 884.62 14.3830–34 7 17.95 51.28 0.35 211.54 602.56 11.7535–39 2 5.13 33.33 0.15 153.85 391.03 11.7340–44 4 10.26 28.21 0.36 115.38 237.18 8.4145–49 3 7.69 17.95 0.43 70.51 121.79 6.7950–54 2 5.13 10.26 0.50 38.46 51.28 5.0055+ 2 5.13 5.13 1.00 12.82 12.82 2.50

Total 39 100.00

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 awayfrom the 30–45-year age category and also changed the range of ages represented in the profile.

These two results support the hypothesis that the use of cementum annulations does alter theshape of the age pyramid and consequently changes some features of the demographic profile in asignificant manner, even when all of the individuals in a sample cannot be included in the analysisdue to preservation, recovery, and other taphonomic issues. When a mortality-centred approach todemography was employed (a life table approach), few significant differences were noted in the demo-graphic profiles. 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 changesignificantly. The median life expectancy (eo) was not significantly different either; life expectancy atbirth was 31 years for the pooled sex sample using age estimates from macroscopic methods and 28years according to cementum annulations (Table 9). Our prediction that mean age at death and lifeexpectancy would shift was not supported by the data. However, it has been previously demonstratedthat mortality-centred approaches to demography are not the most sensitive indicators of changesto the age pyramid (McCaa, 1998, 2002) and thus we also examined the effect of changes to the agepyramid when a fertility-centred approach to demography is employed.

Significant differences in the demographic profiles were obtained when cementum annula-tions were used for age estimation and we used a fertility-centred approach to demography, oraggregated age ratio analysis following Bocquet-Appel and Paz de Miguel Ibanez (2002). Based onthe original macroscopic age estimates, we obtained the following ratio of adults to subadults:15P5 = d(5 − i)/d(5+) = 0.1951. The age pyramid constructed from a combination of macroscopic methodsand counting of cementum annulations produces an estimate of 0.1220 (Fig. 4). With the inclusion ofboth macroscopic and histological indicators of age at death, the Damdama sample appears to havebeen in the intermediate range of population fertility (10 < ratio < 35) according to standards set by theHealth in the Western Hemisphere study (McCaa, 1998). The ratio of 12.21 corresponds to a calibratedGross Reproductive Rate (GRR, or number of female offspring per woman) of roughly 2.4 if the lifeexpectancy at birth is 30–40 years (GRR is 2.7 if the life expectancy were 20 years). Assuming thehigher life expectancy range is correct, this corresponds to a Total Fertility Rate (TFR) of 4.8 offspringper woman. These results demonstrate that the use of cementum annulations in a proportion of a pre-historic cemetery sample (0.30) leads to a significant difference in the age pyramid and this differencewill yield a significant change in the demographic profile if a fertility-centred model is employed.

The results of our analysis indicate support for the hypothesis that problems with age estimationmethods significantly affect the accuracy of palaeodemographic reconstructions (Bocquet-Appel andMasset, 1982). The results suggest support for the hypothesis that fertility-centred approaches todemography are more sensitive than mortality-centred approaches to shifts in the structure of theage pyramid (McCaa, 2002). Finally, our results also demonstrate that changes to the age pyramid



resulting from the use of cementum annulations are of a magnitude significant enough to affect thedemographic profile. Based on these results, we contend that the counting of cementum annulationsis a useful age estimation technique in bioarchaeology because, even when sampling and preservationare not ideal, counting of cementum annulations does significantly improve the accuracy of adult ageestimation and improve palaeodemographic reconstruction.

Acknowledgments

This work would not have been possible without the help and monetary support of several organi-zations 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 theNational Geographic Society and the University of Oregon Graduate School. Thanks to Dr. MurrayMarks, Jeanne Selker and Joanna Lambert for feedback on the methods and the manuscript.

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