Michael 2016.1 32 Dental Anthropology 2016 │ Volume 29 │ Issue 01 Keywords: histology, Wilson bands, health, bioarchaeology The Classic Period Maya (A.D. 250-900) was charac- terized by social stratification, emergence and expan- sion of elite classes, and integration of large urban cen- ters (Cucina and Tiesler, 2003). During this period, the Maya incorporated multiple types of burial sites into their mortuary program across the ancient landscape. Traditionally, archaeologists have focused on the exca- vation of elite structures and tombs in the Maya area, so comparatively little research has been conducted on the bioarchaeological analysis of non-elite Maya burials from mortuary sites outside of civic-ceremonial centers. This study analyzed the dental remains of non-elite in- dividuals distinguished in death by placement in two peripheral locations, Sapodilla (SDR) and Caves Branch (CBR) rockshelters, in Central Belize. Three indicators of stress (dental caries, linear enamel hypoplasia, and Wilson bands) were collected and estimations of age at defect formation were calculated to determine if these peripheral non-elite groups exhibited stress indicators in frequencies comparable to elite groups in the area. This study makes two original contributions: 1) the analysis of an understudied Maya mortuary sample using a method (dental histology) that is not widely applied in the regional bioarchaeology literature; and 2) the assignment of age at defect estimation to better in- terpret episodic health stress events during life. Using these data, rockshelter burials can be compared to two other main mortuary site types in the area, caves and surface sites, to better understand Maya mortuary be- havior. Investigations of rockshelter burials, especially those of the highly socially stratified Classic Period, will help to close the gap between the much-studied elite class and those many communities that existed inde- pendently or semi-independently from urban site cores. Enamel defects can be seen macroscopically on the enamel crown as hypoplasias, taking furrow or pit forms. Microscopically, the defects present as Wilson bands, which are visualized as areas of disrupted ame- loblast activity when the enamel is viewed in cross- ABSTRACT Objectives: Investigations of dental health in the Maya region have frequently focused on indi- viduals buried at urban sites rather than in peripheral or intermediary zones. This study presents a dental analy- sis of a different type of mortuary sample, those persons buried in two non-elite peripheral rockshelters, in Cen- tral Belize using a combined dental micro- and macrodefect approach to interpret health experience. Materials and Methods: A total of 22 teeth (permanent mandibular canines, and mandibular and maxillary third molars) from the two sites were assessed for dental caries, enamel hypoplasias, and Wilson bands. The maximum and minimum ages of microdefect formation for each tooth was calculated. Results: Carious lesions were infrequently represented in the sample, while linear enamel hypoplasias were expressed in less than half the sample. Wilson bands, conversely, were present in nearly every tooth indicating that the rockshelter populations experienced more acute stress. Individuals interred at Caves Branch Rockshelter were affected earlier in life based on analysis of mandibular canines. Conclusion: Non-elites buried in rockshelters in Central Belize had similar dental health experiences when compared with individuals buried at elite centers. At least in terms of oral health, peripheral communities in this area were not adversely affected by their distance from urban core sites *Correspondence to: Amy Michael Department of Anthropology, Michigan State University, East Lansing, MI 48824, e-mail: michae76@msu.edu Histological Analysis of Dentition in Rockshelter Burials from Two Sites in Central Belize Amy R. Michael 1* 1Department of Anthropology, Michigan State University, East Lansing, MI 48824 This paper is one of the winners of the Albert A. Dahlberg Prize awarded at the Annual meeting of the American Association of Physical Anthropologists in 2016. 33 Dental Anthropology 2016 │ Volume 29 │ Issue 01 section. Hypoplasias, observed externally, and Wil- son bands, viewed internally, are indelible markers that indicate a generalized systemic response to stress. The etiology of both defect types is not conclu- sively understood (Hillson 2014), but a combined study of hypoplasias, carious lesions, and Wilson bands allows for more nuanced conclusions about health experience to be drawn from a particular sam- ple. If only macroscopic indicators of health stress were observed, the burial populations may appear to have had a more positive health experience than was their reality. However, by analyzing the teeth for three defect types, it is possible to observe more de- fects at more age ranges, resulting in a more com- plete picture of episodic health stress in the individu- al. Health experience has historically been measured in human skeletal remains as a physiological disrup- tion resulting in some kind of osteological or dental manifestation of pathology. The physical embodi- ment of poor health has often been read in the assess- ment of dental pathology in particular, but a confla- tion of “health” and “stress” is not intended in this study, but the limitations of interpreting dental pathological conditions are acknowledged here. Re- cently, bioarchaeologists have made increasing ef- forts to critically evaluate health studies and stress markers, drawing more conclusions from the incor- poration of theoretical models and data from fields of epidemiology, primatology, and clinical biology (Gowland 2015; Temple and Goodman 2014). For this sample, dental pathological conditions are still used as a proxy of health experience, but the complexities of individual well-being cannot possibly be totalized by observable defects. It is recognized that intangible processes and states of emotional being that leave no inscription on the archaeological record and are diffi- cult to measure on the physical body contribute to overall health experience. To mitigate this fact, multi- ple dental pathological conditions are combined to determine the extent of expression between and among individuals at these rockshelter sites. These baseline data then allow for the proposition of larger anthropological questions of health experience as more archaeological data becomes available. Specifically, five research questions were asked of this sample: 1) What is the age at defect formation range and mean age at defect formation for each tooth?; 2) How many Wilson bands were observed for each tooth?; 3) How many enamel hypoplasias were observed for each tooth?; 4) How many carious lesions were observed for each tooth?; and 5) What is the relationship, if any, between dental macro- and microdefects? BIOCULTURAL CONTEXT The assessment of rockshelter burials in the Maya region is still a relatively new endeavor and certainly part of an ongoing focus on analyzing non-elite buri- al populations (Dunham et al., 1998; Glassman and Villarejo, 2005; Goldstein and Prufer, 1999; Saul et al., 2005; Scott and Brady, 2005; Wrobel, 2008). The data presented in this paper contribute to the continuing collection of information about the lifeways and so- cial identities of individuals from rural communities. Frequently, archaeologists have posited that cave access was governed in part by social status, whereby elites controlled large darkzone caves for the purpos- es of ritual activity while non-elite activity was rele- gated to the smaller, less visually impressive caves and rockshelters (Awe et al., 1998; Graham, 1980; Peterson, 2006; Reents, 1980; Wrobel et al., 2009). The debates over cave and rockshelter use continue in the literature, so the biological data presented here can usefully augment current and future archaeological data collected from these mortuary spaces. Sapodilla Rockshelter (SDR) is situated near a small tributary of the Caves Branch River system within the northern portion of the Caves Branch Riv- er Valley in Central Belize. Over the course of two field seasons of the Central Belize Archaeological Survey (CBAS), the presence of 40 – 50 primary buri- als, commingled human and faunal bones, ceramics, and lithics was confirmed (Michael and Burbank 2013). Ceramic types recovered at SDR reveal that the use of the site was predominantly limited to the Protoclassic and Early Classic periods (Michael and Burbank 2013). The mortuary patterns and artifact assemblages appear similar to another peripheral site, Caves Branch Rockshelter (CBR), located ap- proximately 1 kilometer away (Glassman and Bonor, 2005; Wrobel, 2008). Some evidence for post-mortem secondary manipulation of remains was noted, as one nearly complete burial was absent the cranium yet the mandible was present, and two isolated skulls were discovered. Burials at SDR were easily individ- ualized since interments were either undisturbed or only slightly commingled. Teeth were mostly re- tained in the skull or were scattered in close proximi- ty to the body. Caves Branch Rockshelter, situated in the Caves Branch River Valley east of the present-day Belizean capital of Belmopan, was first excavated by Juan Luis Bonor in the mid-1990s after a number of looting events (Glassman and Bonor, 2005). This salvage op- eration yielded 32 primary burials, but dense com- mingling contributed to the countless bone fragments mixed throughout the burial matrix. Following Bonor’s work, Wrobel continued excavations at CBR during 2005-2007 with the Belize Valley Archaeologi- cal Reconnaissance (BVAR) Project and again in 2015 with CBAS. All CBR excavations demonstrated that the site was unrestricted by sex or age, further sug- 34 Dental Anthropology 2016 │ Volume 29 │ Issue 01 gesting that the mortuary regulations that governed interment in rockshelters were inclusive. Attendant grave artifacts were sparse and utilitarian reflecting use consistent with a rural farming population (Wrobel, 2008). Based on excavations to date, an esti- mated 400-500 individuals may be interred at the site (Wrobel et al., 2009). The ceramic assemblage spanned a large period of time from the Formative Period to the Terminal Classic (Bonor, 2002; Wrobel et al., 2009; Wrobel, 2008). However, the diagnostic ceramics interred as grave goods were largely from the Late Preclassic, indicating that the most intensive use of the rockshelter likely occurred during this pe- riod (Wrobel, 2008a). Two burials were subjected to AMS dating and returned Late Preclassic and Late Classic dates suggesting that use may not have been punctuated, but rather persistent (to varying degrees) through time. During the Late Preclassic, there is no evidence for urban centers in the valley, further un- derscoring the use of the rockshelter by agrarian communities (Wrobel, 2008a). MATERIALS AND METHODS For this study, two tooth classes (permanent and deciduous) mandibular canines, and maxillary and mandibular third molars were selected for analysis following Danforth’s (1989) study on pre-Hispanic Maya burials (Tables 1 and 2). Left teeth were prefer- entially selected, but when not available the antimere was collected for analysis. Due to similarity in enam- el formation rates, both the mandibular and maxil- lary third molars were selected when either was pre- sent. Prior to thin sectioning for microscopic analysis, data were collected on dental caries and enamel hy- poplasias following Buikstra and Ubelaker (1994). Enamel hypoplasias were scored using a combination of the “thumbnail” test and taking an impression in putty. The putty records both slighter expressions of horizontal grooves, as well as pit defects that may not be as immediately visible during the thumbnail test. Color and width of the hypoplasias were not record- ed, as these data have not been demonstrated to pro- vide any useful biological information (Buikstra and Ubelaker 1994). Fitzgerald and Saunders (2005) also stated that variables other than defect presence do not factor into the threshold level or denote severity of the defect. To identify Wilson bands, the parameters ad- vocated by Hillson (2014:174-175) ,which were adapted from Rose et al. (1978:513), and Goodman and Rose (1990:93) were first considered as the stand- ard observation and identification method for this project. Because there is largely no congruent defini- tion in the literature as every author chooses biologi- cal criteria and visual representation to prioritize, it was determined that enamel disruptions would be recorded as Wilson bands if two of the three criteria were met, following a recent Wilson band study done by Reeves (2013:42): 1. the stria appears darker and wider than sur- rounding striae, extending clearly from the dento -enamel junction to the enamel surface 2. the stria exhibits rod disorganization on exami- nation at 1000x magnification 3. the stria has a corresponding darkened stria in the lingual enamel Other criteria could potentially be added to this list, but the general presentation of disorganized enamel prisms, darkened striae, and bilateral expression are repeated most frequently throughout the literature. Each sample was impregnated with a resin/ hardener mixture and cut in midline. One thin sec- tion of approximately 80-100µm (Fitzgerald and Rose, 2000; Hillson, 2014) was created for examina- tion of the internal surface. Thin sections were ana- lyzed using a standard light transmitted binocular LED digital compound microscope with 3D stage and 9MP camera attachment from United Scope. The AmScope 3.7 software included with the microscope was used to image the samples. The digital camera attachment provided a live feed to the computer, as well as an image capture feature. Thin sections were first magnified at 1000x to identify defects, then they were observed and photographed between 400 – Tooth Type Right Left Mandibular canine 4 6 Maxillary third molar 0 0 Mandibular third molar 0 0 Deciduous mandibular canine 3 1 TOTAL 7 7 TABLE 1. Distribution of teeth in the sample from CBR Tooth Type Right Left Mandibular canine 5 1 Maxillary third molar 1 0 Mandibular third molar 0 1 Deciduous mandibular canine 0 0 TOTAL 6 2 TABLE 2. Distribution of teeth in the sample from SDR 35 Dental Anthropology 2016 │ Volume 29 │ Issue 01 600x. Following Cook (1981), Danforth (1989) developed a population-specific age-at-defect formation sched- ule for Maya dental remains (deciduous canines, per- manent canines, third molars). Danforth (pers. comm. 2011) stated that it would be reasonable to use these standards for the rockshelter samples. Measurements of the location of the Wilson bands were taken along the DEJ with the CEJ acting at the zero point. For in- stance, if a Wilson band was recorded at 2.25mm, that means that the defect began 2.25mm from the CEJ. These measurements were matched to the ap- propriate increment for each tooth class. For exam- ple, if a defect in a third molar of a female was noted at 3.1mm from the CEJ, the associated increment would be in Danforth’s DEJ zone 5 and the associated age range would be 10.7 – 11.3 years; (Table 3; see Tables 4 and 5 for other tooth classes). In instances where the sex of the individual was estimated, then the male or female age range was used. The majority of the individuals from CBR and SDR do not have sex estimations, and the combined age ranges were employed. TABLE 3. Age at development for mandibular third molars* (adapted from Danforth 1989) DEJ Zone AmScope Measurement (mm from CEJ) Age in Years (Males) Age in Years (Females) Age in Years (Combined Sexes) 1 6.01 – 7.0 9.0 – 9.6 9.1 – 9.6 9.0 – 9.6 2 5.01 – 6.0 9.6 – 10.3 9.6 – 10.2 9.6 – 10.2 3 4.01 – 5.0 10.3 – 10.9 10.2 – 10.7 10.2 – 10.8 4 3.01 – 4.0 10.9 – 11.6 10.7 – 11.3 10.9 – 11.4 5 2.01 – 3.0 11.6 – 12.3 11.3 – 11.8 11.4 – 12.0 6 1.01 – 2.0 12.3 – 13.0 11.8 – 12.3 12.0 – 12.7 7 0.0 – 1.0 13.0 – 13.6 13.0 – 13.6 *The same chart was used for maxillary third molars based on the very similar development times (Logan and Kronfield 1933) DEJ Zone AmScope Measurement (mm from CEJ) Age in Years (Males) Age in Years (Females) Age in Years (Combined Sexes) 1 11.01 – 12.0 0.7 – 1.1 0.5 – 0.9 0.6 – 1.0 2 10.01 – 11.0 1.1 – 1.5 0.9 – 1.3 1.0 – 1.4 3 9.01 – 10.0 1.5 – 1.8 1.3 – 1.7 1.4 – 1.8 4 8.01 – 9.0 1.8 – 2.2 1.7 – 2.1 1.8 – 2.1 5 7.01 – 8.0 2.2 – 2.6 2.1 – 2.5 2.1 – 2.5 6 6.01 – 7.0 2.6 – 3.0 2.5 – 2.9 2.5 – 2.9 7 5.01 – 6.0 3.0 – 3.4 2.9 – 3.3 2.9 – 3.3 8 4.01 – 5.0 3.4 – 3.8 3.3 – 3.7 3.3 – 3.7 9 3.01 – 4.0 3.8 – 4.1 3.7 – 4.0 3.7 – 4.1 10 2.01 – 3.0 4.1 – 4.5 4.0 – 4.4 4.1 – 4.5 11 1.01 – 2.0 4.5 – 4.9 4.4 – 4.8 4.5 – 4.9 12 0.0 – 1.0 4.9 – 5.3 4.9 – 5.3 TABLE 4. Age at development for mandibular canines (adapted from Danforth 1989) 36 Dental Anthropology 2016 │ Volume 29 │ Issue 01 RESULTS Table 6 summarizes the data collected for the SDR sample. Six individuals were represented in the sam- ple, but only two of these burials retained more than one desired tooth. Of the eight teeth available, none exhibited carious lesions, three showed linear enamel hypoplasias, and all but one tooth had Wilson bands. The number of Wilson bands in each tooth class and the average age point estimate for each tooth class is summarized in Table 7. With one exception, the teeth were all mandibular canines with Wilson band for- mation occurring as early as 2.1 years (Burial 17) and as late as 4.9 years (Burials 9 and 13). The number of Wilson bands per tooth was five or under for six of the seven teeth with microdefects. Burial 17, exhibit- ing 12 Wilson bands, was an outlier. Interestingly, Burial 17 was the only individual in this sample in- terred in the liminal zone between the rockshelter overhang (where most skeletal remains were found) and the small dark zone cave. This burial was also distinguished by the hundreds of shell tinklers form- ing a belt and bracelet on the body; no other burial at SDR was as decorated. Table 8 summarizes the data collected for CBR. Of the fourteen teeth available, all were from different individuals. Only two of these teeth exhibited carious lesions, while six teeth demonstrated at least one lin- ear enamel hypoplasia. Every tooth in the sample expressed at least one Wilson band. The earliest age of Wilson band formation in the deciduous teeth oc- curred at 0.33 – 0.42 years (Burial 19) and the latest DEJ Zone AmScope Measurement (mm from CEJ) Age in Months (Combined Sexes) 1 7.01 – 8.0 5 – 6 (in utero) 2 6.01 – 7.0 7 - 8 (in utero) 3 5.01 – 6.0 9 (in utero) – 1 (post -birth) 4 4.01 – 5.0 2 – 3 5 3.01 – 4.0 4 – 5 6 2.01 – 3.0 6 – 7 7 1.01 – 2.0 8 – 9 8 0.0 – 1.0 10 – 11 TABLE 5. Age at development for deciduous man- dibular canines (adapted from Danforth 1989) Burial Tooth No. of Caries No. of Hypplasias No. of Wil- son bands Min. Age Max. Age Point Esti- mate Age Burial 6 RC_ 0 2 4 3.7 4.5 4.1 Burial 7 RC_ 0 0 5 2.5 3.7 3.1 Burial 9 RC_ 0 0 4 3.3 4.9 4.1 Burial 10 RM3 0 0 0 N/A N/A N/A Burial 10 RC_ 0 1 2 3.3 4.1 3.7 Burial 13 LM3 0 0 3 11.4 12.0 11.7 Burial 13 LC_ 0 2 4 2.5 4.9 3.7 Burial 17 RC_ 0 0 12 2.1 4.1 3.1 Tooth Class Number of Wilson Bands in Entire Tooth Class Point Estimate of Average Affected Age Mandibular canine (n=6) 31 3.63 Third Molar (mandibular and maxillary) (n=2) 3 11.7 Deciduous mandibular canine (n=0) N/A N/A TABLE 6. Summary of data collected for SDR TABLE 7. Summary of SDR sample: Wilson bands and ages 37 Dental Anthropology 2016 │ Volume 29 │ Issue 01 age at defect formation in the deciduous teeth oc- curred at 0.5 – 0.75 years (Burial 71). For the perma- nent teeth, the earliest age at defect formation was 1.3 years (Burial 51) and the latest was 4.5 years (Burial 46C/42). Table 9 summarizes the number of Wilson bands in each tooth class and the average age at de- fect formation for each tooth class. DISCUSSION AND CONCLUSIONS The rockshelter burials investigated in this paper provide baseline data for understanding rural burial populations in Central Belize. Unfortunately, the samples are too small to present statistically signifi- cant results, but the descriptive data do demonstrate some patterns. Carious lesions were rarely present at either site, a trend that follows other sites in Central Belize (Slon and Michael, 2013). Hypoplasias were always linear in formation, but at both sites the total frequency of hypoplastic defects was under 50% for the sample. Nearly all teeth expressed at least one Wilson band indicating that these defects, signaling more acute stress events (Wright, 1990), were the norm in these rockshelter burials. The etiology of these microdefects is still not conclusively known, but the disruption of the enamel prisms can, at mini- mum, be understood to be reflective of some stress event. Of the 22 teeth in the sample, 12 exhibited Wil- son bands without presentation of enamel hypo- plasias demonstrating that the presence of one defect does not necessarily predict the presence of the other. In fact, the burial with the most microdefects (Burial 17 from SDR) showed no linear enamel hypoplasias. The average ages at defect formation for each tooth type demonstrate that individuals at CBR were affected by health stress somewhat earlier (2.96 years Burial Tooth No. of Caries No. of Hypoplasias No. of Wil- son bands Min. Age Max. Age Point Estimate Age Burial 2 LC_ 0 2 1 2.5 2.9 2.7 Burial 9 RC_ 0 5 3 2.6 3.4 3.0 Burial 10 LC_ 0 3 1 2.1 2.5 2.3 Burial 11 LC_ 0 0 3 3.0 3.8 3.4 Burial 19 rc_ 1 0 1 0.33 0.42 0.38 Burial 23b rc_ 0 0 1 0.5 0.58 0.54 Burial 36A rc_ 0 0 1 0.5 0.58 0.54 Burial 38 RC_ 0 0 3 2.9 3.3 3.1 Burial 46C/42 LC_ 0 1 9 1.8 4.5 3.15 Burial 51 RC_ 0 2 7 1.3 4.0 2.65 Burial 63 LC_ 0 0 3 2.9 3.3 3.1 Burial 71 lc_ 0 0 3 0.5 0.75 0.63 Burial 86 RC_ 0 1 7 2.2 3.8 3.0 Burial 246 LC_ 1 0 6 2.0 4.4 3.2 TABLE 8. Summary of data collected for CBR Tooth Class N Number of Wilson Bands in Entire Tooth Class Point Esti- mate of Av- erage Affect- ed Age Mandibular canine 10 43 2.96 Third Molar (mandibular and maxil- lary) 0 N/A N/A Deciduous mandibular canine 4 6 0.52 TABLE 9. Summary of CBR sample: Wilson bands and ages 38 Dental Anthropology 2016 │ Volume 29 │ Issue 01 for the mandibular canine), while health stress oc- curred later at SDR (3.63 years for the mandibular canine). Deciduous canines, only present at CBR, ex- pressed an average age at defect formation at 0.52 years. Third molars, only present at SDR, expressed an average age of defect formation at 11.7 years. What can be gleaned from this project is that, largely, the individuals buried at these two rockshel- ter sites did not experience overwhelming dental health disturbances. These rural communities may once have been assumed to have suffered greater health disparities due to their lower social status, but that hypothesis is proven incorrect here. Previous bioarchaeological research has focused on health ex- perience leading up to (or at the time of) “collapse” when the Maya were re-organizing their political alliances and social structure (Cucina and Tiesler, 2003, 2005; Danforth, 1989, 1997; Gerry, 1997; Storey, 1997; White et al., 2001; White, 1997, 2005; Wright, 1997; 2006), or during the Contact period when the Maya were introduced to new biological and social stresses brought on by the arrival of the Spanish (Danforth, 1989; Wright, 1990). Neither of these peri- ods, in spite of extraordinary cultural change, has been shown to have a significant effect on the devel- opment of Wilson bands and/or enamel hypoplasias. While this study does not focus on periods of so- cial change, the similar results cautiously suggest that residence in peripheral communities (rather than ur- ban centers) did not result in negative biological con- sequences. Non-elites living in rural settlements both adapted to and acted in concert with their surround- ings, responding to both environmental and social pressures. The addition of more health data from rockshelter sites in Central Belize, as well as other Classic Period rockshelters throughout the country, are necessary to determine the extent to which resi- dence in peripheral zones affected health experience. These data indicate that a binary model of health stress (e.g. elites did not suffer, while commoners suffered greatly) likely does not encapsulate the ex- perience of the Classic Period Maya. ACKNOWLEDGEMENTS Dr. Gabriel Wrobel, Dr. Kip Andres, and Dr. Shawn Morton, of the Central Belize Archaeological Survey, allowed me to work on the CBAS project and collect these dental samples. Michigan State University De- partment of Anthropology and the Caves Research Foundation provided partial funding for sample preparation. the samples to be sent to Spectrum Pet- rographics. 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