1 Oral Pathological Conditions in Early Postcontact Guale, St

Home , Guale

Oral Pathological Conditions in Early Postcontact Guale, St. Catherines Island, Georgia

Thesis

Presented in Partial Fulfillment of the Requirements for the Degree Master of Arts in the

Graduate School of The Ohio State University

Kendra Susan Weinrich, B.S.

Graduate Program in Anthropology

The Ohio State University

2020

Thesis Committee

Dr. Clark Spencer Larsen, Advisor

Dr. Debra Guatelli-Steinberg

Dr. Mark Hubbe

Copyrighted by

Kendra Susan Weinrich

2020

Abstract

Situated within the broader context of oral health preceding and proceeding

Spanish contact, documentation of oral pathological conditions in Guale individuals from the pre-Spanish mission Fallen Tree site, St. Catherines Island, Georgia, provides a record of oral health during initial Spanish contact. Oral pathological conditions, including dental caries, periodontal disease (PD), antemortem tooth loss (AMTL), and abscesses, reflect the outcome of multifaceted behavioral and biological circumstances and provide insight into individual and community-based oral health trends. This thesis tests the hypotheses that age-at-death, sex, and/or tooth class correlate directly with presence and severity of dental caries, PD, AMTL, and abscesses in Guale individuals

(n=56) from Fallen Tree. Situated in the larger context of Guale oral health, this thesis predicts higher rates of oral pathological conditions in the Fallen Tree community and declining oral health during Spanish contact. Results from chi-square analyses revealed significant relationships between: 1) age and all oral pathology; 2) sex and dental caries and PD; and 3) tooth class and dental caries and AMTL. Results from a binomial logit generalized linear model revealed significant relationships between sex and dental caries by tooth (p≤0.05). These findings demonstrate variable rates and severity of oral pathological conditions among Fallen Tree individuals with overall declines in oral health compared to precontact communities that continued after Spanish mission establishment.

Dedication

I dedicate this thesis to my parents, Alan and Vicki, and my soon-to-be husband, Micky, who have always encouraged me to pursue my dreams.

iii

Acknowledgments

I would like to thank my advisor, Dr. Clark Larsen, for encouraging me to pursue this research endeavor and carry on the incredible legacy and history of St. Catherines Island.

Additionally, I would like to thank my committee members, Drs. Debra Guatelli-

Steinberg and Mark Hubbe for helping me to refine my research focus and pursue deeper meaning. Special thanks to Drs. Fabian Crespo and Kathryn Marklein for help with periodontal disease data collection and analysis. Lastly, I would like to thank Dr. Sean

Downey for never-ending statistical support.

Vita

May 2014………………Gilmanton High School

May 2018………………B.S., Anthropology Honors, summa cum laude

Beloit College

2019-2020………………Graduate Teaching Assistant

Department of Anthropology

The Ohio State University

Field of Study

Major Field: Anthropology

Table of Contents

Abstract ...... ii Dedication ...... iii Acknowledgments...... iv Vita ...... v List of Tables ...... viii List of Figures ...... ix Chapter 1. Introduction ...... 1 Biocultural Context ...... 2 Oral Pathology Etiologies ...... 9 Behavioral and Biological Contributors to Oral Health ...... 10 Research Questions and Hypotheses ...... 14 Chapter 2: Materials and Methods ...... 17 Materials ...... 17 Methods...... 18 Statistical Approach ...... 19 Chapter 3. Results ...... 22 Age and Oral Pathology Presence and Severity ...... 22 Sex and Oral Pathology Presence and Severity ...... 25 Tooth Class and Oral Pathology Presence and Severity ...... 29 Chapter 4. Discussion ...... 31 Age and Oral Pathology Presence and Severity ...... 31 Sex and Oral Pathology Presence and Severity ...... 32 Tooth Class and Oral Pathology Presence and Severity ...... 35 Temporal Trends in Oral Health ...... 36 Future Research ...... 37 vi

Chapter 5. Conclusion ...... 39 References ...... 40 Appendix A. Oral Pathology Scoring Systems ...... 46

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List of Tables

Table 1. Age Class and Sex of Fallen Tree Individuals ...... 18 Table 2. Description of Caries Severity Scoring System (Adapted from Larsen (1985)) 46 Table 3. Description of Periodontal Disease Scoring System for CEJ to Alveolar Bone Measurement (Adapted from DeWitte and Bekvalac (2009) and Fujita and Adachi (2017))...... 46 Table 4. Description of Antemortem Tooth Loss Scoring System (Adapted from Buikstra and Ubelaker (1994)) ...... 47

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List of Figures

Figure 1. Map of Georgia Coast (Adapted from Thomas, 2008) ...... 3 Figure 2. Tooth Level Age and Oral Pathology Prevalence...... 24 Figure 3. Individual Level Age and Oral Pathology Prevalence...... 25 Figure 4. Tooth Level Sex and Oral Pathology Prevalence ...... 28 Figure 5. Individual Level Sex and Oral Pathology Prevalence ...... 28 Figure 6. Tooth Class and Oral Pathology Prevalence ...... 30 Figure 7. Temporal Trends in Dental Caries Prevalence at the Tooth Level...... 37

Chapter 1. Introduction

Known as the Georgia Bight, the Atlantic coastal area from North Carolina to

Florida was home to numerous native societies, including the Guale of coastal and inland

Georgia (Larsen et al., 2002). Although Guale cultural records are sparse, Spanish missionaries documented a clear social hierarchy within Guale culture, reflected in the mortuary record by the presence and extent of grave goods (Winkler, 2017). The remarkable preservation of the mortuary record along this coast, and especially on St.

Catherines Island, Georgia, has produced comprehensive information relating to two

Guale transitionary periods—the adoption of maize agriculture in the 12th century and the beginning of Spanish colonization and later missionization in the 16th and 17th centuries. The transition from hunting and foraging to agricultural practices for the Guale people was accompanied by increasing reliance on starch carbohydrates, especially maize, and simultaneous decline in marine resource consumption, creating a dietary environment conducive to poor oral health (i.e. dental caries, periodontal disease, abscesses, and antemortem tooth loss). Together, these developments created conditions for nutritional stress in Guale individuals living at the time of initial Spanish contact.

Guale life prior to and after contact has been well-documented ethnohistorically, archaeologically, and bioarchaeologically over the last 40 years from preserved material culture and human remains. However, a distinct gap in the bioarchaeological record in

1 the century between the end of the precontact period and the establishment of the Santa

Catalina de Guale mission (ca. 1600) exists due to the absence of material culture and skeletal remains from this period. The 2013 discovery of the cemetery at the Fallen Tree site on St. Catherines Island holds promise in bridging the gap between the precontact and postcontact periods and increasing knowledge of Guale life during initial Spanish contact. The study of oral pathology prevalence and severity at the Fallen Tree site may provide insight into the direct impacts of Spanish colonial policies on Guale diet, oral health, and ultimately, Guale everyday life.

Biocultural Context

St. Catherines Island is one of several Georgia coastal islands (See Figure 1) home to the Indigenous Guale people for thousands of years that reveals a rich archaeological and bioarchaeological narrative of Guale adaptation and subsistence transitions from foraging and hunting to agriculture (Larsen et al., 2002). Based on varying subsistence strategies, Guale history may be separated into four general periods, including the precontact preagricultural (BC1100-1150AD), precontact agricultural

(AD1150-1550), early contact (AD1550-1680), and late contact (AD1680-1702) periods.

Figure 1. Map of Georgia Coast (Adapted from Thomas, 2008)

Precontact Preagricultural Period

The Guale are likely the descendants of Native American groups who migrated to

St. Catherines Island around 4,500 years ago during the Late Archaic period (Thomas,

2008; Reitz et al., 2010). Beginning during the initial Guale settlement of St. Catherines

Island and ending during the mid-12th century after the adoption of agricultural practices, the precontact preagricultural period encompasses the longest timespan of the four periods. Based on the presence of shellfish middens and highly dispersed surface deposits, Guale individuals likely lived in small temporary communities along the coast 3 of the island and subsisted via foraging, hunting, and fishing (Larsen, 1982; Larsen et al.,

2002). The unique intersection of marine, freshwater, and terrestrial ecosystems along the

Georgia coast enabled Guale communities to make use of a wide variety of marine, estuarine, and terrestrial resources (Reitz et al., 2010). The zooarchaeological record suggests marine resources were the most abundant dietary staple, as evidenced by the plethora of remains from oysters, molluscs, and other crustaceans, as well as sharks, fish, and other marine vertebrates and invertebrates in midden deposits. Although marine resources provided the most consistent protein source for Guale individuals during this period, diet was also supplemented by occasional terrestrial fauna, with deer accounting for the highest biomass of any terrestrial animal (Reitz et al., 2010). Although the zooarchaeological record provides a detailed history of faunal marine and terrestrial resources used by Guale communities, poor preservation and limited recovery of plant remains have obscured the ethnobotanical record for this period. However, foraged plant material, including nuts, berries, tubers, and roots, likely contributed a significant portion of dietary nutrients (Reitz et al., 2010; Larsen, 1990; Reitz and Scarry, 1985).

In the cultural context of small, isolated communities and subsistence via foraging, hunting, and fishing, the bioarchaeological record provides additional evidence of activity patterns, pathology, nutrition, and stress experienced by Guale individuals living during the precontact preagricultural period. As expected by frequent foraging, hunting, and fishing, Guale individuals had high rates of degenerative joint disease and high long bone robusticity (Ruff and Larsen, 1990). Consistent with zooarchaeological and paleoethnobotanical evidence, low prevalence of dental caries suggests Guale

4 individuals in the pre-farming context consumed limited amounts of carbohydrate-rich foods (Larsen et al., 2002; Reitz et al., 2010). Individuals likely experienced frequent stress events, as evidenced by high linear enamel hypoplasia prevalence (Hutchinson and

Larsen, 1990). Because low prevalence of cribra orbitalia, porotic hyperostosis, and periostitis indicate low occurrence of infectious disease, consistent with low population density, these stress events were likely nutritionally based (Larsen et al., 2002).

Precontact Agricultural Period

Based on settlement remains, Guale communities began to migrate towards the island’s interior and clustered together more closely at the beginning of the precontact agricultural period (Larsen, 1982). Furthermore, settlement population density increased, likely resulting from increases in fertility and fecundity facilitated by agriculturally- produced weaning foods (Larsen et al., 2002). Based on paleoethnobotanical evidence,

Guale communities on St. Catherines Island began to cultivate native plants around

AD1000, including varieties of melon, squash, beans, and maize (Thomas, 2008; Reitz et al., 2010). Guale individuals used agriculturally-produced resources to supplement their diet, as paleoethnobotanical evidence suggests foraged resources, such as acorns and hickory nuts, continued to be eaten (Thomas, 2008). Although agriculturally-sourced plants began to supplement foraged plants, stable isotopic evidence suggests local terrestrial, estuarine, and marine resources remained important dietary staples, most likely as consistent protein sources (Hutchinson et al., 1998). Furthermore, such resources served as fallback foods in case of crop failure or shortage (Thomas, 2008).

Such societal changes are also reflected in the bioarchaeological record, as individuals from the precontact agricultural period exhibit higher rates of periostitis and porotic hyperostosis—albeit lower rates of cribra orbitalia—than precontact agricultural individuals, likely indicative of higher rates of infectious disease facilitated by higher population density (Schultz, Larsen, and Kreutz, 2001). Additionally, the introduction of agriculturally-sourced plants likely increased carbohydrate consumption, contributing to a distinctive spike in dental caries prevalence during the precontact agricultural period

(Larsen et al., 2002). Combined with noted increases in fertility, this dietary shift especially increased dental caries prevalence in female individuals (Larsen et al., 2002;

Russell, Choi, and Larsen, 1990). However, Guale individuals living during the precontact agricultural period have lower linear enamel hypoplasia prevalence than individuals from earlier periods suggesting fewer general stress events (Hutchinson and

Larsen, 1990), as well as lower rates of degenerative joint disease and reduced long bone robusticity, indicative of changing activity patterns (Ruff and Larsen, 1990).

Early Contact

According to 16th century Spanish texts written upon Spanish arrival on St.

Catherines Island, agricultural production facilitated expansion of small Guale communities into stratified chiefdoms at the time of Spanish contact (Thomas, 2008;

Jones, 1978). Consisting of two towns that alternated leadership roles, chiefdoms were the social mechanism for amassing and redistributing resources among different communities. Exploiting this social system, Spanish missionaries and military members

6 established the mission outpost Santa Catalina de Guale on St. Catherines Island in the mid-16th century after having built several missions along the Atlantic coast in the early

16th century (McEwan, 2001; Hann, 1990).

Upon the creation of the mission Santa Catalina de Guale, most Guale individuals on St. Catherines Island, as well as neighboring Apalachee and Timucua groups on the mainland coast, were forcibly missionized and congregated together in or near the mission (Stojanowski, 2005; Larsen, 1990). Known as reducción, this system was an attempt to establish Spanish control and promote Spanish acculturation and indoctrination of Guale individuals (Larsen, 1990). Additionally, reducción aggregated a Guale labor force, resulting in a draft system known as repartimiento in which Guale males were forced to cultivate maize as a monocrop for food tribute to local missions as well as for

Spanish export (Worth, 2001). As a result, maize surpassed other agriculturally-produced foods as a dietary staple for Guale individuals living in the island mission, Santa Catalina de Guale (Larsen et al., 2002; Larsen, 1990).

Despite dramatically reduced dietary diversity and increasing stress from the other effects of Spanish colonialism, Guale individuals living during early Spanish contact have rates of pathology similar to those of precontact agricultural individuals, including nearly identical rates of dental caries, linear enamel hypoplasia, and periostitis (Larsen et al., 2002; Hutchinson and Larsen, 1990). Unlike precontact agricultural individuals,

Guale individuals from Santa Catalina de Guale exhibit increases in cribra orbitalia and porotic hyperostosis, likely a result of increased rates of infectious disease due to poor sanitary conditions and overcrowding in the mission (Schultz, Larsen, and Kreutz, 2001).

Although not pathological, there is also a distinct increase in long bone robusticity, especially in males, during this period presumably resulting from increases in labor demand required by the Spanish (Ruff and Larsen, 1990).

Late Contact

In response to British attacks in the area, all occupants of the Santa Catalina de

Guale mission on St. Catherines Island relocated to the abandoned mission Santa Maria on Amelia Island on the Atlantic coast of Florida (Worth, 1995; Hann, 1990). Consistent with historical narratives of unprecedented stress, Guale individuals living during the late contact period exhibited the highest pathology rates of all groups, including dramatic increases in dental caries prevalence, cribra orbitalia, porotic hyperostosis, and periostitis

(Larsen et al., 2002; Schultz, Larsen, and Kreutz, 2001). Additionally, Guale individuals have higher rates of degenerative joint disease and increases in long bone robusticity, a result of prolonged subjugation to the reducción and repartimiento forced labor systems

(Reitz et al., 2010; Ruff and Larsen, 1990). Contrary to other health trends, late contact individuals have the lowest prevalence of linear enamel hypoplasia (Hutchinson and

Larsen, 2001). Although individuals from this period may reflect a heavily biased sample of Guale communities living on Amelia Island, this finding may also reflect Guale adaptation to stress or fewer stress events overall (Larsen et al., 2002).

Oral Pathology Etiologies

Oral pathological conditions, including dental caries, periodontal disease, antemortem tooth loss, and abscesses, reflect the outcome of multifaceted etiological circumstances and thus are highly interconnected (Roberts and Manchester, 2005;

Selwitz et al., 2007). Dental caries is a process by which carbohydrate-consuming

Streptococcus mutans bacteria produce acid byproducts, lowering the pH of the oral environment surrounding a tooth (Larsen, 2015; Selwitz et al., 2007). Because strains of

Streptococcus mutans outcompete other oral bacterial species, there is a drastic decline in oral microbiome diversity, thus increasing risk of future dental caries (Mukherjee et al.,

2018). This acidic imbalance slowly demineralizes tooth enamel, thus weakening the outer structural integrity of the tooth and promoting the future decay of the inner tissue layers. If left untreated, the pulp chamber—the innermost layer of soft tissue containing nerves and blood vessels—becomes exposed to pathogens and subsequent infection

(Roberts and Manchester, 2005). Severe infection of the pulp chamber produces excessive pus that permeates the surrounding alveolar bone, creating characteristic drainage channels identified as abscesses (Buikstra and Ubelaker, 1994). Over time, a severe infection cuts off blood supply from a tooth and the nerve tissue dies, ultimately resulting in antemortem tooth loss and alveolar resorption (Roberts and Manchester,

2005). Therefore, dental caries, abscesses, and antemortem tooth loss have highly interconnected etiologies.

Periodontal disease has a slightly differently connection to infection and antemortem tooth loss than dental caries (Larsen, 2015). The process leading to

9 periodontal disease begins when dental plaque accumulates in small crevices between teeth and surrounding soft tissue (Roberts and Manchester, 2005). Over time, dental plaque accumulation forms gaps between the tooth and gingiva and generates visible gingival inflammation that leads to collagen degeneration (Larsen, 2015). If left untreated, gingival inflammation permeates the surrounding alveolar bone, often causing alveolar bone resorption and eventual antemortem tooth loss (American Academy of

Periodontology, 2003). Therefore, due to the mix of overlapping and distinct etiologies of oral pathologies, it is crucial to include multiple indicators of oral pathology in documenting oral health.

Behavioral and Biological Contributors to Oral Health

As teeth are often the only remains to preserve in the bioarchaeological record, dental remains offer a wealth of information about an individual’s life that may not preserve in other postcranial remains (Kendall et al., 2018). Therefore, dental remains from the Fallen Tree community hold information about individual oral health, which may yield information about general health in the narrative of Guale resistance and adaptation to Spanish colonialism.

In the behavioral realm of oral health, dietary composition and texture and oral hygienic practices are important factors to consider in understanding oral health and risk of oral pathology development (Larsen et al., 1990; Kumar et al., 2016). Oral bacteria feed on the carbohydrates consumed by an individual, in turn, producing acid byproducts that lower the pH of the mouth, leading to enamel demineralization and dental caries

(Selwitz et al., 2007). Therefore, consumption of a carbohydrate-rich diet may increase risk of developing oral pathology and create additional risk for socially-marginalized individuals with reduced access to non-carbohydrate-based foods, such as females

(Larsen, 1998). Furthermore, such diets are often texturally soft, allowing food particles to coat the tooth surface and accumulate in interproximal surfaces and crevices on the occlusal surface, thus encouraging bacterial growth, acid byproduct production, and dental plaque (Organ et al., 2005; Roberts and Manchester, 2005). If not removed via oral hygienic practices, such as brushing or flossing, dental plaque accumulation may increase dental caries and periodontal disease risk, thereby increasing risk of abscess formation and antemortem tooth loss (Kumar et al., 2016). Therefore, such behavioral factors place additional oral pathology risk on older individuals with extensive exposure to oral bacteria and dental plaque (Hillson, 1996). However, several biological factors may mitigate or amplify such behavioral risks.

In the biological realm of oral health, salivary composition and flow are two of the most important factors in assessing oral pathology risk (Ferraro and Vieira, 2010).

Salivary composition includes multiple buffering systems that offer protection against oral pathology, including protein and bicarbonate buffering systems (Humphrey and

Williamson, 2001). Numerous salivary proteins, including immunoglobin A (IgA) and amylase, accept hydrogen ions and function as basic buffers in highly acidic oral environments with a pH less than 5, thereby neutralizing acids and limiting enamel demineralization (Cheaib and Lussi, 2013; Lukacs and Largaespada, 2006). However, the bicarbonate buffering system provides the most protection against oral pathology

11 formation because this system operates within the average pH range of 7.2-6.3 for human saliva, unlike the highly acidic range for protein buffering (Baliga et al., 2013; Cheaib and Lussi, 2013). The bicarbonate buffering system stabilizes salivary pH by neutralizing free floating hydrogen ions produced by bacteria in the mouth, in turn producing carbon dioxide and water (Bardow et al., 2000).

In addition to buffering capacity, the antimicrobial and mechanical properties of saliva and the composition of the oral microbiome offer protective qualities against oral pathology (Mukherjee et al., 2018; Lukacs and Largaespada, 2006). Several salivary proteins possess antimicrobial properties that regulate the oral microbiome, limiting the ability of pathogenic bacteria to reproduce (Lukacs and Largaespada, 2006). For example, lactoferrin binds iron and promotes iron withholding from pathogenic bacteria

(Fine et al., 2002). Although individual oral microbiomes vary drastically in composition in relation to environmental factors, oral microbiomes remain consistent over time within a single individual (Mukherjee et al., 2018). Saliva also acts mechanically as a tooth cleanser, removing food particles, dental plaque, and bacteria from tooth crevices and interproximal areas (Dodds et al., 2015). Therefore, it is the combination of high salivary flow and the antimicrobial and buffering properties of saliva that provides the most protection against oral pathologies (Lukacs and Largaespada, 2006).

There are well-documented differences in salivary flow and composition related to age (Nassar et al., 2014; Smith et al., 2013; Eliasson et al., 2006). For example, Nassar and colleagues (2014) and Smith and colleagues (2013) found significantly lower rates of salivary flow in older adults than younger age groups, suggesting age-related salivary

12 gland degeneration may heighten risk of oral pathology. Concerning salivary composition, Eliasson and colleagues (2006) documented higher IgA protein levels in the minor gland salivary secretions of older adults, likely resulting from extended exposure to oral pathogens. Therefore, acid neutralization via the protein buffering system may offer additional protection against enamel erosion and oral pathology in older adults, although as Xu and colleagues (2019) note, higher salivary protein levels may be a function of reduced salivary flow. Therefore, in conjunction with low salivary flow, the beneficial properties of heightened IgA levels do not provide adequate protection from oral pathology, resulting in generally higher rates of oral pathology in adults and older individuals (Lukacs and Largaespada, 2006).

It has been suggested that sex-based differences in salivary flow rate and composition may also play a role in differential risk of oral pathology development

(Ferraro and Viera, 2010). For example, Eliasson and colleagues (2006) found females have lower salivary flow rates than males, which is typically associated with smaller salivary glands and hormonal fluctuations in females. Additionally, Eliasson and colleagues (2006) documented higher concentrations of IgA in the minor gland secretions of males than females, thus providing greater protection against enamel demineralization in males. However, it is unclear how pregnancy may heighten some of these sex-based differences (Laine, 2002). Although Eliasson and colleagues (1988) and Salvolini and colleagues (1998) found higher protein levels in the salivary composition of pregnant women, suggested enhanced protein buffering against enamel erosion, Laine and colleagues (1988) and Rockenbach and colleagues (2006) found no significant difference

13 in salivary composition or salivary flow rate in pregnant females. Contrastingly, Karnik and colleagues (2015) found lower salivary rates in pregnant females. Because biological contributors to oral health are highly interconnected and often influence one another, it is often difficult to tease apart exact connections and there is not a clear-cut biological association between sex and oral health (Laine, 2002; Streckfus et al., 1998).

Hypotheses and Research Questions

Although Guale subsistence and diet before and after contact is well-known, there is relatively little known about Guale life during initial Spanish. Guale individuals interred at the Fallen Tree Mortuary site lived through the beginnings of Spanish colonization, mission establishment, and forced agricultural production in North America and endured drastic assaults by the Spanish to usurp cultural lifeways and norms

(Garland and Reitsema, 2018). Therefore, in documenting oral health in the Fallen Tree community, this thesis seeks to understand how Guale diet changed after Spanish contact, and in turn, how subgroups within the Fallen Tree community were differentially impacted by dietary change. To understand Guale cultural change, resistance, and adaptation in the face of colonization in the bioarchaeological context of St. Catherines

Island, I aim to address the following hypotheses.

Hypothesis 1: Previous Guale oral health studies found similar oral pathology rates for juveniles and adults after the adoption of agriculture, a decline in juvenile oral pathology rates after Spanish missionization, and a marked increase for juveniles and adults during

14 the late contact period (Larsen et al., 1991; Larsen, 1982). I hypothesize that due to prolonged exposure to oral pathogens as the Fallen Tree community shifts towards intensive maize farming, adults will exhibit higher rates and greater severity of oral pathology than juveniles.

Hypothesis 2: Although oral pathology rate and severity increased over time for most

Guale communities living on St. Catherines Island, Larsen (1998) notes a marked increase for females compared to males after the adoption of agriculture. This has been attributed, at least in part, to behavioral risk factors, including greater maize carbohydrate consumption for females than males (Larsen et al., 1990). Therefore, I hypothesize due to differences in subsistence behavior, females in the Fallen Tree community consumed more maize than males, resulting in greater risk and higher prevalence of oral pathology for females compared to males.

Hypothesis 3: Previous research has demonstrated a consistent relationship between tooth class and oral pathology prevalence and severity in Guale communities over time on St.

Catherines Island (Larsen et al., 1991; Larsen, 1983). Based on these studies, posterior teeth for all individuals exhibit the highest increase in oral pathology prevalence and severity over time, although anterior teeth also exhibit heightened oral pathology rates after the adoption of maize agriculture through the late contact period. I hypothesize the combination of complex tooth morphology and high occlusal surface area for posterior

15 teeth, as well as increased maize consumption for all individuals, resulted in higher oral pathology rates and severity for posterior teeth compared to anterior teeth.

Chapter 2: Materials and Methods

Materials

Individuals comprising the basis of this study were buried at the Fallen Tree cemetery on St. Catherines Island and are currently housed at the Bioarchaeology

Research Laboratory at The Ohio State University. Situated in the larger context of the ongoing St. Catherines Island Research Project, this study contributes oral pathology prevalence and severity as an additional line of evidence related to the dietary shift towards maize consumption, complementing previous studies of ethnobotanical remains and stable isotope analysis in the Fallen Tree community.

Demographic profiles of the Fallen Tree community demonstrate that individuals represent a wide variety of ages, ranging from neonates (0-2 months) to old age (50 years and older) (Betz and Mayus, 2019). Although at least 72 individuals were uncovered from the Fallen Tree cemetery, only 56 individuals were included in this study based on dental preservation. Of the individuals with observable sex, there are approximately equal numbers of females (n=17) and males (n=15). However, there are far more adults (n=40) than children (n=16) in this collection. A complete demographic summary of the individuals included in this study is provided in Table I. Most skeletal elements are present for most individuals and most cranial and postcranial skeletal elements are well-

17 preserved, although the remains of a small group of individuals include only the maxillary and mandibular dentition.

Table 1. Age Class and Sex of Fallen Tree Individuals

Description n Age Class 1 (Neonate-0-2 months) 2 2 (Infant-2 months-3 years) 8 3 (Child-4-12 years) 3 4 (Adolescent-13-19 years) 3 5 (Young Adult-20-34 years) 11 6 (Middle Adult-35-49 years) 9 7 (Old Adult-50+ years) 2 8 (Adult) 18 Sex 0 (Unobservable) 19 1-2 (Female) 17 3 (Indeterminate) 5 4-5 (Male) 15 Total 56

Methods

For this project, I recorded carious lesion, periodontal disease, antemortem tooth loss, and abscess prevalence in 56 individuals from St. Catherines Island, Georgia. I collected dental caries data by recording the number and severity (summarized in Table

II) of carious lesions in observable teeth using standards adopted from Larsen (1985).

Periodontal disease data were collected by scoring alveolar bone porosity (summarized in

Table III) and measuring the distance between alveolar bone and cementoenamel junction

(CEJ) with a digital calipers, based on DeWitte and Bekvalac’s (2009) and Fujita and

Adachi’s (2017) methods. Abscesses were recorded on an absence/presence basis based on the presence of a drainage channel in alveolar bone near the root apex as recommended by Buikstra and Ubelaker (1994). Antemortem tooth loss was recorded based on the extent of alveolar resorption as outlined in Buikstra and Ubelaker (1994) in observable maxillary and mandibular alveolar bone (summarized in Table IV). Unerupted teeth, teeth with postmortem damage or breakage, and severely worn teeth were scored as unobservable. Teeth without associated alveolar bone and teeth not in occlusion were also scored as unobservable for periodontal disease and abscesses. Due to the low count of observable teeth for abscesses and antemortem tooth loss, limited statistical analysis of these pathologies was included in this study.

Statistical Analysis

Age and Oral Pathology Presence and Severity

I incorporate several statistical methods to address the different facets of

Hypothesis 1. For all statistical tests, juveniles were defined as individuals with an age class of 1 (neonate or 0-2 months), 2 (infant or 2 months-3 years), 3 (child or 4-12 years), or 4 (adolescent or 13-19 years). Adults included individuals with an age class of 5

(young adult or 20-34 years), 6 (middle adult or 35-49 years), 7 (old adult or 50 years or older), and 8 (adult with indeterminable age range). Chi-square is an appropriate statistical test for comparing differences in oral pathology absence and presence by age at the tooth and individual levels (Meng et al., 2011; Wasterlain et al., 2009). For dental caries, I also created a Generalized Linear Model (GLM), which allows age class to be 19 used as a factor to predict binary oral pathology absence or presence, providing a means to quantify risk of oral pathology with every increase in age class. I did not create a GLM for antemortem tooth loss or abscesses due to the limited data for these pathologies, nor periodontal disease, as these data are not binary. To assess age-based differences in dental caries severity, I used a Kruskal-Wallis test, which allows dental caries to be ranked by severity and age. A Shapiro-Wilk test for normality suggests periodontal disease measurement data were not normally distributed (W (152)=0.97, p=0.001).

Therefore, to assess periodontal disease severity by age and sex, I ran a nonparametric

Wilcoxon test with age class as a categorical variable.

Sex and Oral Pathology Presence and Severity

Similar statistical approaches were used to test Hypothesis 2. For all statistical tests, females were defined as individuals with a sex score of 1 (female) or 2 (likely female) and males included individuals with a sex score of 4 (likely male) or 5 (male). I used a chi-square test to evaluate the hypothesis that females have higher rate and severity of oral pathology at the individual and tooth levels than males (Meng et al.,

2011; Wasterlain et al., 2009). I also created a GLM using sex as a predictor of binary dental caries absence and presence, as well as testing possible interactions between age and sex. Additionally, I incorporated a Kruskal-Wallis test to assess sex-based differences in dental caries severity. Periodontal disease measurement data were not normally distributed, and I therefore ran a nonparametric Wilcoxon test to assess sex- based differences in periodontal disease severity (Abanto et al., 2014).

Tooth Class and Oral Pathology Presence and Severity

To evaluate Hypothesis 3, I use chi-square to evaluate the hypothesis that there is a relationship between tooth class and oral pathology at the tooth and individual levels.

Statistical testing was not possible for tooth class and periodontal disease due to limited data.

Chapter 3. Results

Age and Oral Pathology Presence and Severity

Dental Caries

In support of the proposed hypothesis, a chi-square test comparing dental caries absence and presence in juveniles and adults at the tooth level suggests teeth from adults have significantly more carious lesions than those of juveniles (χ2 (1, N=889) 24.27, p=8.37x10-7). While 23.7% (163/689) of adult teeth had dental caries, only 7.5% (15/200) of juvenile teeth exhibited the same oral pathology (See Figure 2). At the individual level, there is also a significant difference in dental caries absence and presence between juveniles and adults (See Figure 3). 80% (32/40) of adults and 28.6% (4/14) of juveniles have dental caries (χ2 (1, N=54) 10.14, p=0.0015). A GLM (family= “binomial”, link=

“logit”) was used to determine if age class could predict binary dental caries absence or presence, as well as test potential interactions between sex and age. Although the model was not significant for age or an age and sex interaction, this is likely due to limited observations for individuals with known sex and age. Upon removing sex as a limiting factor in the model, there are marked increases in the odds of having dental caries for middle adults and old adults compared to adolescents and young adults. The results of the

Kruskal-Wallis test ranking dental caries severity in juveniles and adults also suggests the

22 severity of dental caries increases with age, as adults (n=689, mu=0.49) have more severe carious lesions than do juveniles (n=200, mu=0.12) (K(1)=26.15, p=3.22x10-7).

Periodontal Disease

The results of a chi-square analysis suggest there is a significant difference in periodontal disease prevalence between juveniles and adults at the tooth level (χ2 (2,

N=152)=21.69, p=1.95x10-5). Although 27.6% (35/127) of adult teeth had moderate to severe periodontal disease, no juvenile teeth exhibited evidence of periodontal disease

(See Figure 2). At the individual level, there were an insufficient number of individuals to statistically compare periodontal disease prevalence. 68% (17/25) adults had at least one tooth or alveolus with moderate to severe periodontal disease, compared to no juveniles

(See Figure 3).

A Wilcoxon test was performed to examine the average extent of alveolar resorption between juveniles and adults. The relation is significant, W (m1=1.15, m2=2.61, N=152)=2772.5, p=3.88x10-9. Therefore, there is a relationship between periodontal disease severity and age, and adults have greater periodontal disease severity than juveniles.

Antemortem Tooth Loss

A chi-square test comparing antemortem tooth loss in juveniles and adults at the tooth level suggests adults lose significantly more teeth before death than juveniles (χ2 (1,

N=1,046)=30.75, p=2.93x10-8). While 14.1% (119/844) of adult observable alveolar bone or tooth sockets exhibited evidence of resorption, no juvenile teeth did so (See Figure 2). 23

This significance also holds at the individual level, as 67.5% (27/40) of adult individuals exhibited evidence of antemortem tooth loss, compared to no (0/13) juvenile individuals

(χ2 (1, N=53) 15.29, p=9.23x10-5; See Figure 3).

Abscesses

Although there were an insufficient number of observable teeth to statistically compare abscess prevalence between adults and juveniles, abscess prevalence at the tooth and individual levels is informative. On the tooth level, 6.8% (33/484) of adult teeth had abscesses compared to 4.8% (3/63) of juvenile teeth (See Figure 2). At the individual level, 51.6% (17/33) of adults had least one abscess compared to 37.6% (3/8) of juveniles

(See Figure 3).

10 Teeth Teeth Affected (%) 5

0 Dental Caries Periodontal Disease Antemortem Tooth Abscesses Loss

Juveniles Adults

Figure 2. Tooth Level Age and Oral Pathology Prevalence.

There are marked age-based differences for oral pathology at the tooth level, most notably for dental caries, periodontal disease, and antemortem tooth loss.

90 80 70 60 50 40 30

20 Individuals Affected IndividualsAffected (%) 10 0 Dental Caries Periodontal Disease Antemortem Tooth Abscesses Loss

Juveniles Adults

Figure 3. Individual Level Age and Oral Pathology Prevalence. Although juvenile and adult individuals have relatively similar rates of abscesses, adult individuals exhibit markedly higher rates of dental caries, periodontal disease, and antemortem tooth loss than juvenile individuals.

Sex and Oral Pathology Presence and Severity

Dental Caries

A chi-square test comparing dental caries absence and presence in females and males at the tooth level suggests teeth from females have significantly more carious lesions than those of males (χ2 (1, N=643)=11.2, p=0.0008). Comparatively, 31.6%

(104/329) of female teeth versus 19.7% (62/314) of male teeth had at least one carious lesion (See Figure 4). However, this sex-based significance does not hold at the individual level, as female (94.1% (16/17)) and male (80% (12/15)) individuals had roughly equal levels of dental caries absence and presence (χ2 (1, N=32)=0.448, p=0.503;

See Figure 5). Females (n=334, mu=0.70) also have significantly more severe dental 25 caries than males (n=316, mu=0.35) at the tooth level, based on the results of the

Kruskal-Wallis test (K(1)=13.4, p=0.00025). A GLM (family= “binomial”, link= “logit”) was used to determine if sex could predict binary dental caries absence or presence. The model was found to be significant (Est. 0.616; std. err. 0.186; null deviance of 742 of 649 df; p=0.0009). The results of the GLM suggest that the effect of being female markedly increases the odds of having dental caries 6.15:1, meaning females are more likely to have dental caries than males.

Periodontal Disease

A chi-square test comparing periodontal disease absence and presence in females and males at the tooth level suggests male teeth (40% (22/55)) are significantly more affected by periodontal disease than female teeth (19.7% (13/66)) (χ2 (2, N=121)=16.74, p=0.0002; See Figure 4). Although there are an insufficient number of individuals to statistically compare periodontal disease absence and presence by sex at the individual level, male individuals (72.7% (8/11)) had higher rates of periodontal disease than female individuals (64.3% (9/14); See Figure 5).

A Wilcoxon test was performed to examine the average extent of alveolar resorption between females and males. There is a significant relationship between periodontal disease severity and sex, as males have more severe periodontal disease than females (W (m1=2.28, m2=3.05, N=121)=1029, p=4.29x10-5).

Antemortem Tooth Loss

The results of a chi-square test suggest there is no significant difference in antemortem tooth loss at the tooth level between females (15.1% (62/411)) and males

(14.5% (56/385)) (χ2 (1, N=796)=0.013, p=0.91; See Figure 4). This trend holds at the individual level as well, as 76.5% (13/17) of female individuals and 86.7% (13/15) of male individuals lost at least one tooth well before death (χ2 (1, N=32)=0.08, p=0.78; See

Figure 5).

Abscesses

There is no significant difference in abscess presence and absence in female

(7.3% (19/261) and male (6.5% (16/248)) teeth at tooth level (χ2 (1, N=509)=0.038, p=0.85; See Figure 4). This trend also holds at the individual level, as there is no significant difference in abscess presence and absence in female (64.7% (11/17)) and male (53.3% (8/15)) individuals (χ2 (1, N=32)=0.086, p=0.77; See Figure 5).

45 40

35 30 25

15 Teeth Teeth Affected (%) 10 5 0 Dental Caries Periodontal Disease Antemortem Tooth Abscess Loss Female Male Figure 4. Tooth Level Sex and Oral Pathology Prevalence Although teeth from females have significantly higher rates of dental caries, teeth from males have significantly higher rates of periodontal disease. Teeth from both sexes have approximately equal rates of antemortem tooth loss and abscesses.

100 90 80 70 60 50 40

IndividualsAffected (%) 20 10 0 Dental Caries Periodontal Disease Antemortem Tooth Abscesses Loss

Female Male Figure 5. Individual Level Sex and Oral Pathology Prevalence Female and male individuals have similar rates of dental caries, periodontal disease, antemortem tooth loss, and abscesses.

Tooth Class and Oral Pathology Presence and Severity

Dental Caries

A chi-square test comparing dental caries absence and presence by tooth class suggests that molars (26.7% (69/258)) have significantly more dental caries than canines

(21% (25/119)), premolars (19.7% (46/233)), and incisors (13.2% (36/272)) (χ2 (3,

N=882)=15.22, p=0.002; See Figure 6). Furthermore, molars have the most severe dental caries (n=258, mu=0.63), while canines (n=119, mu=0.34), and premolars (n=233, mu=0.36) have equally severe caries, followed by incisors (n=272, mu=0.25)

(K(3)=18.58, p=0.00033).

Periodontal Disease

Due to a low number of observable teeth for periodontal disease, statistical testing was not possible for tooth type (See Figure 6). Based on prevalence, canines (38.5%

(5/13)) and incisors (36.4% (4/11)) had the highest rates of periodontal disease, followed by premolars (25.8% (16/62)) and molars (15.2% (10/66)).

Antemortem Tooth Loss

A chi-square test suggests there are significant differences in antemortem tooth loss by tooth class as molars (21.2% (71/335)) are the most common tooth lost before death (X(3, N=1,036)=48.16, p=1.97x10-10; See Figure 6). Premolars (9.1% (24/264) are

29 the second most common tooth lost before death, followed by canines (6.4% (9/140)) and incisors (5.1% (15/297)).

Abscesses

There are no significant differences in abscess absence and presence based on tooth type (χ2 (3, N=582)=0.86, p=0.84; See Figure 6). Although incisors (7.5% (8/107)) exhibited the highest abscess prevalence, this was closely followed by premolars (6.6%

(12/183)), molars (5.9% (13/221)), and canines (4.2% (3/71)).

45 40 35 30 25 20 15

Teeth Teeth Affected (%) 10 5 0 Dental Caries Periodontal Disease Antemortem Tooth Abscess Loss

Incisor Canine Premolar Molar

Figure 6. Tooth Class and Oral Pathology Prevalence Molars exhibited significantly higher rates of dental caries and antemortem tooth loss than incisors, canines, and premolars. Incisors and canines have the highest rates of periodontal disease, albeit the smallest sample size. Rates of abscesses were approximately equal for all tooth classes.

Chapter 4. Discussion

Age and Oral Pathology Presence and Severity

Based on the results of this analysis, Guale adults have higher rates of dental caries, periodontal disease, antemortem tooth loss and abscesses than Guale juveniles at the tooth and individual levels. On average, adults have higher rates of oral pathology than juveniles due to both extended exposure to behavioral and biological etiologies of oral pathologies and gradual oral tissue degeneration due to aging (Benatti et al., 2009;

Hillson, 1996). Hillson (1996) notes that as adult teeth are exposed to more pathogens and dental plaque buildup than juvenile teeth simply due to their longer existence, adults commonly have more oral pathology than juveniles, as well as more severe pathology.

Furthermore, tissue degeneration associated with senescence may also increase the risk of oral pathology in adults, as the number of periodontal ligament cells anchoring teeth in occlusion decrease over time and the distance between the CEJ and AC increases with age, while immune response declines and chronic inflammation increases (Persson, 2017;

DeWitte, 2012; Benatti et al., 2009). Together, these four factors dramatically increase the risk of developing oral pathologies over time.

As there are far more adults (n=40) than juveniles (n=16) in the Fallen Tree community, demographic representation skews to the left, meaning there is an overrepresentation of Guale adults. Due to differential preservation of juvenile and adult 31 remains, it is likely juveniles are especially underrepresented from the living Fallen Tree community. However, statistical testing accounts for such differences in sample sizes, thereby validating the higher rates and severity of oral pathology in adults than juveniles.

Sex and Oral Pathology Presence and Severity

On the tooth level, females from Fallen Tree had significantly higher rates and severity of dental caries than males, while males had significantly higher rates and severity of periodontal disease. However, on the tooth level, both sexes exhibited roughly equal rates of antemortem tooth loss and abscesses. On the individual level, females and males had similar rates and severity of all oral pathologies.

Dental Caries

Across time and space, females generally exhibit higher rates and severity of dental caries than males (Larsen, 2015). This phenomenon has been associated with two explanatory paradigms, including sexual division of diet and agriculturally-enhanced fertility (Larsen, 1998; Lukacs, 2008). The sexual division of diet hypothesis attributes the higher dental caries rate in females as the result of females consuming more carbohydrate-rich foods than males (Larsen, 1998; Larsen et al., 1990). The agriculturally-enhanced fertility hypothesis attributes higher caries rates in females to increases in fertility via greater access to weaning foods, enabling females to have more children in shorter timespans (Lukacs, 2008). Lukacs (2008) claims that pregnancy highly impacts the oral microbiome, thereby lowering salivary pH and reducing salivary

32 flow, and more frequent pregnancies encourages bacterial growth and increases the risk of developing oral pathologies in females of childbearing age.

Numerous lines of evidence suggest maize was an important dietary staple in the

Fallen Tree community. Stable isotopic data from bone carbonate suggests Guale individuals at Fallen Tree were consuming similar amounts of maize to individuals in the

Santa Catalina de Guale community (Reitsema, 2019). This interpretation is also supported by the abundant archaeobotanical evidence of maize at the cemetery (Purcell and Scarry, 2019). Therefore, a maize-rich diet likely played a crucial role in increasing oral health challenges for all Guale individuals (Larsen et al., 2001), which would explain the almost equal dental caries rates in Guale females and males at the individual level, as well as similar rates of antemortem tooth loss and abscesses. However, at the tooth level, females have higher dental caries rates and severity than males. Larsen (1998) suggests sex-based differences in diet were common in Guale communities and females traditionally consumed more carbohydrates than males. Therefore, despite the greater maize consumption for all individuals, females still likely consumed more maize, leading to higher dental caries rates and severity. Furthermore, demographic studies indicate population decline and reduced fertility overall, which contradicts the agriculturally- enhanced fertility hypothesis (Betz and Mayus, 2019). Rather, reduced fertility may have resulted from physiological stress exacerbated by Spanish colonialism (Stojanowski,

2005).

Periodontal Disease

In the Fallen Tree community, males exhibited higher rates of periodontal disease than females at both the tooth and individual levels. This trend of higher periodontal disease and severity in males has been well-documented in both modern and bioarchaeological populations (Tomczyk et al., 2018; DeWitte, 2012). Shiau and

Reynolds (2010) and Galligan and Fish (2015) attribute this trend to differential immune responses between males and females, as females generally have stronger innate and adaptive immune responses than males, resulting in stronger immune responses to pathogens than males. Although it is difficult to assess immune competence in human remains, preliminary results from skeletal inflammatory phenotype reconstructions at

Fallen Tree suggest immunological responses were highly complex and individualized and there were no clear sex-based trends (Marklein et al., 2020).

It is possible behaviorally-based factors were responsible for the higher rate and severity of periodontal disease for males (Larsen, 2015). Early Spanish historical records suggest tobacco use was common especially for men among Guale communities and several clay pipe fragments dating to the Savannah period have been uncovered on St.

Catherines Island (Francis et al., 2011; Thomas et al., 2008). Additional, modern studies of periodontal disease have found notable differences in oral hygiene between women and men, which may account for such sex-based differences (Furuta et al., 2011; Shiau and Reynolds, 2010).

Tooth Class and Oral Pathology Presence and Severity

Antemortem tooth loss exhibited the greatest difference in oral pathology prevalence by tooth type, as twice as many molars than any other tooth class were lost before death. Additionally, incisors and canines had the highest rates of periodontal disease, although these two tooth classes also had the smallest sample sizes, and molars had the greatest number and highest severity of dental caries. Rates of abscesses were nearly identical among all tooth classes.

As bacteria and plaque commonly accumulate in small occlusal crevices, dental caries often develop on the occlusal surface of premolars and molars, as well as the interproximal surface between teeth (Roberts and Manchester, 2005). Therefore, the high rate and severity of dental caries in molars fit the prediction. However, the high rate and severity of dental caries in incisors in the Fallen Tree community compared to individuals living during precontact and early postcontact periods was unpredicted and may be related to diet (Larsen et al., 2002).

Due to the recent discovery of the Fallen Tree site, dental microwear has not been evaluated to date. However, dental microwear and stable isotope data from Guale individuals living at the Santa Catalina de Guale mission have been well-documented and indicate most individuals consumed a soft, maize-rich diet (Hutchinson et al., 1998;

Larsen et al., 2001). Stable isotopic data from Fallen Tree individuals are similar to Santa

Catalina de Guale individuals, suggesting maize was already a dietary staple at the beginning of Spanish contact (Reitsema, 2019). As individuals from early and late contact periods likely consumed similar diets and prepared maize in a similar fashion, it 35 is likely such soft, carbohydrate-rich foods may have created microfilm on their anterior teeth, thus increasing the susceptibility of anterior teeth—especially incisors—to enamel demineralization via bacterial acid byproducts (Larsen et al., 2001).

Temporal Trends in Oral Health

This study documented dental caries, periodontal disease, antemortem tooth loss, and abscesses in the Fallen Tree community. However, dental caries are generally considered the most sensitive oral pathology measure of dietary shifts toward maize consumption (Larsen, 2015). On St. Catherines Island, there is a clear temporal trend of increasing dental caries prevalence and severity in Guale communities, most notably after the adoption of maize agriculture and the establishment of Santa Catalina de Guale

(Larsen et al., 2002). Based on data from Larsen and colleagues (2002), although low during the precontact preagricultural period for all Guale individuals (1.2% (2479)), dental caries prevalence markedly increased to 9.6% (5984) during the precontact agricultural period (See Figure 7). This notable shift is likely the result of increased starchy carbohydrate (maize) consumption for all individuals.

Although dental caries prevalence remains relatively constant during the early contact period (7.6% (4466)), there was a second spike in dental caries during the late contact period after the movement of all members of the Santa Catalina de Guale mission to Amelia Island (19.6% (1548)) (Larsen et al., 2002). In combination with an increasingly stressful environment, Guale individuals likely consumed more maize than previous periods, resulting in higher prevalence of dental caries. Interestingly, dental

36 caries prevalence in the Fallen Tree community (20% (889)) almost mirrors the late contact period, which is consistent with other evidence suggesting maize was a well- established dietary staple even at the time of Spanish contact.

20 20 19.6

10 9.6

7.6

5 TEETH WITH CARIOUS LESIONS (%) LESIONS CARIOUS WITHTEETH 1.2 0 PP PA EC LC TIME PERIOD

Figure 7. Temporal Trends in Dental Caries Prevalence at the Tooth Level. Dental caries prevalence steadily increases from the precontact preagricultural (PP) to precontact agricultural (PA) periods, followed by a slight decline in the early contact (EC) period and drastic spike during the late contact (LC) period. Situated during initial contact, the Fallen Tree community (star) has a drastically high dental caries prevalence in comparison to PA and EC communities, fitting best with LC communities.

Future Research

While these research findings suggest that age, sex, and tooth class are all important contributors to oral health in the Fallen Tree community, the scope of oral pathology data collection was quite limited for this study, especially for periodontal 37 disease. As voiced by DeWitte (2012), the common measurement of periodontal disease in bioarchaeological samples as the distance between the CEJ and alveolar bone is problematic. This method is unable to account for standard human variation in the distance between the CEJ and alveolar bone and does not account for non-pathological continued eruption of the teeth, leading to perceived higher rates of periodontal disease.

Therefore, future research incorporating new methodologies that address these issues, such as micro-CT scans, may provide a more accurate representation of overall oral health in the Fallen Tree community.

Furthermore, while this study identified connections between age, sex, and tooth class and dental caries, periodontal disease, antemortem tooth loss, and abscesses, there are likely several other factors that contribute to oral health, including the composition of the oral microbiome and immune competency. Future studies documenting these more complex contributors to oral health in bioarchaeological contexts will provide greater insight into the complexity of oral health.

Chapter 5. Conclusion

Prior to the discovery of the Fallen Tree site, a distinct gap in knowledge of Guale health persisted between the end of the precontact preagricultural period and the establishment of the Santa Catalina de Guale mission on St. Catherines Island. The results of this study suggest there are clear relationships between age, sex, and tooth class and oral pathology presence in Guale individuals from the Fallen Tree community. Guale adults had higher rates and severity of oral pathology than juveniles, and adult females had additional risks of developing oral pathology. Furthermore, incisors and molars exhibited the highest rates of oral pathology, likely due to a soft, high carbohydrate diet.

Although relationships between age, sex, and tooth class have been well- documented in many populations, Guale individuals from the Fallen Tree community experienced the additional stress of living during the beginning of Spanish colonialism in the American southeast (Stojanowski, 2005). This likely augmented already heightened behavioral and biological predispositions for oral pathologies, resulting in higher rates and severity of oral pathologies in the Fallen Tree community compared to earlier Guale communities on St. Catherines Island (Larsen et al., 2002). This trend of increasing rates and severity of oral pathologies continued in later Guale populations forcibly aggregated into Spanish missions (Larsen, 1990), underscoring the amplified stresses of colonialism under Spanish rule.

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Appendix A. Oral Pathology Scoring Systems

Table 2. Description of Caries Severity Scoring System (Adapted from Larsen (1985))

Score Description 0 Unobservable 1 Small pit 2 Medium (clearly discernable pit; sizeable, but lesion still restricted to enamel) 3 Large (pit extending into pulp chamber) 4 Total destruction (ranging from destruction or near destruction of tooth crown to only tooth root tip present)

Table 3. Description of Periodontal Disease Scoring System for CEJ to Alveolar Bone Measurement (Adapted from DeWitte and Bekvalac (2009) and Fujita and Adachi (2017))

Score Description 0 Unobservable 1 <2 mm (Normal) 2 2-3 mm (Mild PD) 3 3-5 mm (Moderate PD) 4 >5 mm (Severe PD)

Table 4. Description of Antemortem Tooth Loss Scoring System (Adapted from Buikstra and Ubelaker (1994))

Score Description 0 Unobservable 1 Tooth present, in occlusion 2 Tooth present, not in occlusion (postmortem tooth loss) 3 Tooth absent, no alveolar resorption 4 Tooth absent, some alveolar resorption 5 Tooth absent, alveolar bone fully resorbed