Paleoepidemiology of Periodontal Disease and Dental Calculus in The

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2008 Paleoepidemiology of Peridontal Disease and Dental Calculus in the Windover Population (8BR246) Maria Therese Fashing

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COLLEGE OF ARTS AND SCIENCES

PALEOEPIDEMIOLOGY OF PERIDONTAL DISEASE AND DENTAL CALCULUS

IN THE WINDOVER POPULATION (8BR246)

MARIA THERESE FASHING

A Thesis submitted to the Department of Anthropology in partial fulfillment of the requirements for the degree of Master of Science

Degree Awarded: Fall Semester, 2008

The members of the Committee approve the Thesis of Maria Fashing defended on October 6, 2008.

______Glen Doran Professor Directing Thesis

______Frank Marlowe Committee Member

______Lynne Schepartz Committee Member

Approved:

______Glen Doran, Chair, Department of Anthropology

The Office of Graduate Studies has verified and approved the above named committee members.

ACKNOWLEDGEMENTS I would like to thank Glen Doran for his advice and assistance with this thesis. He provided me with an incredible opportunity to examine dental disease in a well-preserved skeletal sample. I would also like to thank the other members of my committee, Frank Marlowe and Lynne Schepartz, for their comments. I would like to express my gratitude to my parents for their support throughout graduate school. My mother, Gisela, provided valuable information about periodontal disease and dental calculus based on her many years as a dentist. My father Norman, a biology professor, was always willing to answer questions about statistics. Their suggestions and encouragement made the process of writing a thesis much easier. I would also like to thank Richard McCoy for his support and enthusiasm throughout this research project. He was always willing to listen to my ideas and offer suggestions. Most of all, he was patient and helpful when I spent long hours analyzing data and writing.

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TABLE OF CONTENTS LIST OF TABLES ...... vi LIST OF FIGURES ...... vii ABSTRACT ...... ix CHAPTER 1 – INTRODUCTION ...... 1 Periodontal Disease ...... 1 Etiology ...... 2 Epidemiology ...... 3 Previous Studies of Periodontal Disease in Archaeological Populations ...... 4 CHAPTER 2 – MATERIALS AND METHODS ...... 7 Windover (8BR246)...... 7 Sample...... 8 Dental Disease Assessment...... 9 Other Florida Sites ...... 15 Manasota Key Cemetery (8SO1292) ...... 18 Highland Beach (8PB11) ...... 20 Fort Center (8GL12) ...... 20 Republic Groves (8HR4) ...... 21 Bird Island (8DI52) ...... 21 CHAPTER 3 - RESULTS ...... 23 Introduction ...... 23 Locations of Alveolar Resorption ...... 23 Prevalence of Periodontal Disease in the Entire Windover Sample ...... 25 Prevalence of Periodontal Disease by Age ...... 26 Prevalence of Periodontal Disease by Sex ...... 29 Antemortem Tooth Loss ...... 32 Locations of Dental Calculus ...... 33 Prevalence of Dental Calculus in the Entire Windover Sample ...... 35 Prevalence of Dental Calculus by Age ...... 38 Prevalence of Dental Calculus by Sex ...... 41 Relationship between Dental Calculus and Periodontal Disease ...... 43

Prevalence of Dehiscences and Fenestrations ...... 44 Prevalence of Dehiscences and Fenestrations by Age ...... 44 Prevalence of Dehiscences and Fenestrations by Sex ...... 47 Locations of Dehiscences and Fenestrations ...... 50 Relationship between Dehiscences and Fenestrations and Periodontal Disease ...... 51 Relationship between Periodontal Disease and Other Dental Health Problems ...... 52 Periodontal Disease in Different Lineages at Windover...... 53 CHAPTER 4 – DISCUSSION AND CONCLUSIONS ...... 57 APPENDIX ...... 66 BIBLIOGRAPHY ...... 69 BIOGRAPHICAL SKETCH ...... 75

LIST OF TABLES Table 1: Demography of the Windover sample...... 9

Table 2: Florida archaeological sites discussed in this thesis...... 21

Table 3: Mean CEJ-AC measurements of maxillary molars in the Windover sample, ordered from highest to lowest...... 24

Table 4: Prevalence of antemortem tooth loss in the maxillary molars of the Windover sample. 33

Table 5: Minimum, maximum, and mean calculus scores for the maxillary molars in the Windover sample, ordered from highest to lowest...... 35

Table 6: Distribution of alveolar defects by tooth surface...... 51

Table 7: Periodontal disease prevalence in Florida hunter-gatherer populations...... 58

Table 8: Windover periodontal disease and dental calculus results by individual...... 66

Table 9: Summary of G Test of Independence results for dental conditions...... 68

LIST OF FIGURES

Figure 1: Buccal measurements of the distance from the cement-enamel junction to the alveolar crest...... 11

Figure 2: Dehiscence on the maxillary left first molar of individual #95.11...... 11

Figure 3: Fenestration on the maxillary right second molar of individual #36.504...... 12

Figure 4: Abscess associated with the maxillary right first molar of individual #74.42...... 14

Figure 5: Left maxillary premolars and molars exhibiting ―no periodontal disease‖ in an individual approximately 19 years of age (#69.1)...... 16

Figure 6: Right maxillary premolars and molars exhibiting ―slight to moderate‖ periodontal disease in an individual approximately 42 years of age (#501.6)...... 16

Figure 7: Left maxillary premolars and molars exhibiting ―slight to moderate‖ periodontal disease in an individual approximately 53 years of age (#121.23)...... 17

Figure 8: Right maxillary molars exhibiting ―moderate to severe‖ lingual bone loss due to periodontal disease. The individual is approximately 38 years of age (#57.077)...... 17

Figure 9: Extensive alveolar bone loss due to severe periodontal disease in an individual approximately 38 years of age (#82.3)...... 18

Figure 10: Right maxillary molars exhibiting ―probable periodontal disease‖ in an individual approximately 58 years of age (#93.3)...... 19

Figure 11: Prevalence of periodontal disease in the entire Windover sample...... 26

Figure 12: Prevalence of periodontal disease by age group...... 28

Figure 13: Prevalence of periodontal disease by sex...... 30

Figure 14: Percentage of the Windover sample with each maximum calculus score...... 37

Figure 15: Prevalence of dental calculus in the entire Windover sample...... 38

Figure 16: Prevalence of dental calculus by age group...... 40

Figure 17: Prevalence of dental calculus by sex...... 42

Figure 18: Prevalence of dehiscences by age group...... 45

Figure 19: Prevalence of fenestrations by age group...... 47

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Figure 20: Prevalence of dehiscences by sex...... 48

Figure 21: Prevalence of fenestrations by sex...... 49

Figure 22: Prevalence of periodontal disease by lineage...... 55

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ABSTRACT The skeletal remains from the Windover site (8BR246) provide a unique opportunity to understand the dental health of an Archaic period hunter-gatherer population. Windover is a mortuary pond located in Brevard County, Florida, dated to approximately 7400 years B.P. Excavations at the site recovered at least 168 individuals, which is one of the largest and best- preserved skeletal collections of this antiquity from North America. This thesis analyzes the epidemiology of periodontal disease and dental calculus at Windover. Mild forms of both of these dental health problems are common in the Windover population, with only a small percentage of individuals affected by severe periodontal disease or dental calculus. The prevalence of periodontal disease and dental calculus increases with age, but the distribution of these dental health problems is not significantly associated with sex in the Windover sample. The relationship between periodontal disease and other dental health characteristics, including alveolar bone defects, caries, and abscesses, is also assessed. The relative importance of hereditary factors in periodontal disease susceptibility is evaluated through a discussion of periodontal disease prevalence in two Windover lineages that are hypothesized to be genetically distinct. Comparison of the periodontal disease prevalence at Windover with other early Florida hunter-gatherer populations reveals variability between these populations, indicating that factors other than diet may affect susceptibility to periodontal disease. Etiological and demographic factors that may account for variability between these populations are suggested. Factors that may influence the prevalence and severity of periodontal disease in past populations include dental calculus, age, sex, alveolar bone defects, susceptibility to dental health problems, and heredity.

CHAPTER 1 – INTRODUCTION

This thesis analyzes the epidemiology of periodontal disease and dental calculus in the Windover population (8BR246). The research provides insight into the dental health of a Florida hunter-gatherer population that lived over 7000 years ago. The study will assess the distribution of periodontal disease and dental calculus in the Windover population and compare these results to other early Florida hunter-gatherer populations. The study will then identify factors that may account for variability in the prevalence of periodontal disease between these archaeological populations. This introduction first defines periodontal disease and describes the etiology and epidemiology of the disease in modern populations. Previous studies of periodontal disease in archaeological populations are then reviewed. The archaeological sites analyzed in this study and the methods for periodontal disease assessment will be discussed in the next chapter. Periodontal Disease Periodontal disease refers to two associated dental health problems, gingivitis and periodontitis. The development of these periodontal diseases causes an inflammation of the tissues that surround and support the tooth, known as the periodontium. These tissues include the cementum of the root surface, the alveolar bone of the jaw, the periodontal ligament, and the gingiva (Hillson 1996; Strohm and Alt 1998). Periodontal inflammation is primarily caused by bacteria that accumulate due to insufficient oral hygiene (Hildebolt and Molnar 1991; Strohm and Alt 1998). The early stages of periodontal disease may be limited to an inflammation of the gums, a condition known as gingivitis (Schluger et al. 1977). Early gingivitis can remain stable, but continued bacterial irritation of the gums may cause the expansion of the inflammation over time (Soames and Southam 2005). If gingivitis worsens, the inflammation may create a deeper lesion that affects the entire periodontium. Once the gingival inflammation spreads to the deeper layers and begins to destroy the periodontal ligament and the alveolar bone, the condition is called periodontitis (Molnar and Molnar 1985; Soames and Southam 2005). Expansion of the lesion leads to detachment of the periodontal ligament that would normally attach the gingiva and the alveolar bone to the cementum on the root of the tooth (Hildebolt and Molnar 1991). As a result, a periodontal pocket forms next to the tooth, exposing the cementum of the root surface and permitting the accumulation of bacteria in this area. The lesion may then stabilize for

1 several months, which is often followed by episodic enlargement of the lesion over a period of many years (Hillson 1996). During these episodes, the alveolar bone undergoes progressive resorption, leading to extensive bone loss and changes in the bone texture. Bone loss can occur in a uniform way around all teeth, known as horizontal bone loss, or can create localized defects around individual teeth (Karn et al. 1984; Saari et al. 1968). The loss of the supporting alveolar bone can eventually lead to tooth loss (Hildebolt and Molnar 1991). The severity of the disease varies greatly, however, and tooth loss does not always occur. Gingivitis and periodontitis are chronic diseases that cause progressive destruction of the periodontium at unpredictable intervals. Depending on the individual, gingivitis can remain stable for years, slowly develop into periodontitis, or progress rapidly into severe periodontitis. The progression of alveolar bone destruction in periodontitis is also inconsistent, alternating between periods of resorption and remission. Periodontal disease is therefore conceptualized as a chronic problem that involves loss of periodontal ligament attachment and alveolar bone during brief episodes at random time intervals throughout life (Soames and Southam 2005). When an individual from Windover is diagnosed with periodontal disease, then, the disease may or may not have been active at the time of death. The alveolar bone resorption observed in the individual could have occurred at any time during their life. Although the term periodontal disease includes both gingivitis and periodontitis, only the latter can be recognized in archaeological individuals. Gingivitis only causes an inflammation of the gums, without affecting the skeleton. Periodontitis, on the other hand, can be identified by examining alveolar bone loss and changes to the bone texture that occur as a result of the disease. Since individuals often live with chronic periodontitis for many years, evidence of the random destructive periods of the disease is preserved in the skeleton (Mitchell 2003). The inability to assess gingivitis in archaeological individuals probably leads to an underestimate of the actual prevalence of periodontal disease in ancient populations. Archaeological assessments of periodontal disease should therefore be regarded as estimates of the prevalence of periodontitis rather than gingivitis. Etiology Dental plaque is thought to be the main etiological agent that initiates the development of periodontal disease (Soames and Southam 2005). Plaque refers to a deposit composed primarily of bacteria that live in the mouth and obtain nutrients from saliva, gingival crevice fluid, and diet

(Hillson 1996). Modern clinical experiments have shown that plaque causes an inflammatory response in the adjacent gingiva. If the plaque is removed, the inflammation typically subsides and the gingiva returns to a healthy condition (Albandar 2002). Without plaque removal, the continuous bacterial irritation of the gums may eventually lead to established gingivitis and periodontitis. When plaque remains on the teeth for a long period of time, the plaque deposits can mineralize to form dental calculus. Calculus is a hard substance composed largely of inorganic, crystalline salts with some organic proteins, carbohydrates, and lipids (Scheie 1989). Large deposits of dental calculus often accumulate near the salivary gland ducts, both on the lingual surfaces of the mandibular anterior teeth and the buccal surfaces of the maxillary molars (Shafer et al. 1974). Dental calculus is classified into two types: supragingival and subgingival. Supragingival calculus is attached to the exposed crown of the tooth, and subgingival calculus is located beneath the gingiva. Subgingival calculus is harder, darker, and less extensive than supragingival calculus (Hillson 1996; Shafer et al. 1974). Large deposits of calculus often include both the supragingival and subgingival types. In addition to plaque, calculus may facilitate the development of periodontal disease. The rough surface of dental calculus can irritate the nearby gingival tissues during tooth movement and chewing (Shafer et al. 1974). Calculus also provides a surface for the rapid growth of additional plaque and other bacteria, provoking an inflammatory response in the gingiva (Davies et al. 1997; Schluger et al. 1977). In these ways, dental calculus deposits can facilitate the development and progression of periodontal disease. The relationship between dental calculus and periodontal disease will be discussed further in Chapter 3. Other possible etiological agents involved in the development and progression of periodontal disease will also be examined in Chapters 3 and 4. Epidemiology Periodontal disease is one of the most common dental diseases in modern populations, typically affecting over half of the adult population (Soames and Southam 2005). Most of these individuals exhibit slight to moderate levels of attachment loss, with only a small percentage of individuals suffering from severe periodontitis (Locker et al. 1998). The prevalence and severity of periodontal disease have been found to increase with age, with the elderly exhibiting considerably more periodontal disease than young adults (Albandar 2002; Goldman and Cohen

1973; Schluger et al. 1977). Studies have also found that males may be more susceptible to periodontal disease than females (Goldman and Cohen 1973; Hildebolt and Molnar 1991). Thus, epidemiological studies of modern populations suggest that periodontal disease increases with age and is more common in males than females, but the majority of adults exhibit some degree of periodontal disease. Previous Studies of Periodontal Disease in Archaeological Populations The prevalence of periodontal disease in archaeological populations has been a controversial topic in paleoepidemiology. Some researchers believe that periodontal disease can be accurately diagnosed in archaeological populations using either measurements of alveolar bone loss or qualitative assessments of the condition of the bone (Costa 1982; Fyfe et al. 1993; Hildebolt et al. 1988; Kerr 1991; Lavelle and Moore 1969; Lavigne and Molto 1995; Molnar and Molnar 1985; Newman and Levers 1979). Early studies of periodontal disease supported the idea that ancient populations experienced little periodontal disease, with the prevalence of periodontal disease increasing in populations during recent centuries (Lavelle and Moore 1969; Newman and Levers 1979). Lavelle and Moore (1969) found that the amount of alveolar resorption was significantly greater in mandibles from seventeenth-century London than mandibles from a sixth-century Anglo-Saxon cemetery. Newman and Levers (1979) concluded that Anglo-Saxon populations exhibited continuous eruption of the teeth to compensate for reduction in the height of the occlusal plane because of attrition, and any apparent loss of bone was related to continuous eruption rather than periodontal disease. Both Lavelle and Moore (1969:72) and Newman and Levers (1979:341) believed that periodontal disease was much lower in ancient populations than modern populations because of the soft texture of the diet during the past few centuries, which would facilitate the accumulation of the bacteria that initiate gingival irritation. The increased consumption of sugar and other refined carbohydrates might also have led to a greater prevalence of periodontal disease in recent centuries (Lavelle and Moore 1969). More recent studies of periodontal disease have recognized that the prevalence of periodontal disease has been variable between archaeological populations, with factors other than diet also influencing the development of periodontal disease (Fyfe et al. 1993; Hildebolt et al. 1988; Molnar and Molnar 1985; Whittaker et al. 1998). In a study of several prehistoric populations from Hungary, Molnar and Molnar (1985:59) found that the percentage of teeth affected by alveolar lesions varied from 29.7% to 79% in different populations. The results

4 showed a trend of increasing periodontal disease from young to old age, indicating that age was a factor in periodontitis. Hildebolt and Molnar (1988) studied prehistoric populations from the state of Missouri, concluding that variability in periodontal disease prevalence between individuals from different regions might be explained by geochemical factors. Fyfe, Chandler, and Wilson (1993) provided another explanation for the high prevalence of periodontal disease among pre-seventeenth century individuals from the Solomon Islands, suggesting that the practice of chewing betel leaf might have encouraged the development of calculus and alveolar recession. A more recent study of Romano-British and eighteenth-century London populations by Whittaker, Molleson, and Nuttall (1998) addressed the possibility that dental calculus might influence the development of periodontal disease. Although this study did not find an association between the amount of calculus and changes in alveolar bone height in these two populations, the authors acknowledge that future studies of the relationship between these two variables are necessary. Studies of periodontal disease from the past few decades have focused on identifying the possible causes of variability in disease prevalence between archaeological populations. Other researchers have argued that many of these studies overestimate the prevalence and severity of periodontal disease in archaeological populations (Clarke 1990; Clarke et al. 1986; Danenberg et al. 1991; Watson 1986; Whittaker et al. 1990). These researchers contend that many specimens presumed to have periodontal disease actually have another condition, such as dental abscesses or continuous tooth eruption. According to Clarke (1990), researchers have inadequately discriminated between bone loss associated with periodontitis and abscesses. Dental abscesses result from the death of the tooth pulp from dental caries, attrition, or trauma. The buildup of pus often leads to the development of a hole in the adjacent bone to allow drainage (Hillson 1996). These abscesses can cause loss of a large area of localized bone, which might be mistaken for periodontal disease. In a large study of skulls from thirty-four populations, Clarke and colleagues (1986) demonstrated that bone loss actually attributable to periodontal disease was present in less than 10% of teeth. The study also concluded that premodern populations exhibited much less alveolar resorption than modern populations. While Clarke (1990) has cautioned that dental abscesses rather than periodontal disease cause much of the observed bone loss, other researchers have argued that attrition and resulting continuous eruption have led to mistaken observations of periodontal disease. Watson (1986) examined

British skulls with dates ranging from the Anglo-Saxon to Tudor periods and concluded that alveolar recession probably resulted from occlusal forces and attrition rather than periodontal disease. Later studies have also found that continuous eruption of the teeth to maintain proper occlusion despite severe attrition may be responsible for the changes in alveolar bone that other researchers have referred to as periodontal disease (Danenberg et al. 1991; Whittaker et al. 1990). Therefore, all of these studies have concluded that periodontal disease prevalence in past populations is relatively low, and that continuous eruption or dental abscesses are the actual cause of much of the bone loss that has been termed periodontal disease. This study of the Windover population considers the problems addressed by all of these researchers. This assessment of periodontal disease includes analysis of dental abscesses and attrition as well as measurements of alveolar bone loss. The study also proposes possible factors that might account for differences in disease prevalence between archaeological populations. In these ways, the current study examines the controversial issues in periodontal disease research using a sample from the Windover site.

CHAPTER 2 – MATERIALS AND METHODS

Windover (8BR246) Windover is a mortuary pond located in Brevard County in central Florida near Titusville. The site was discovered when human remains and artifacts were found during construction of the Windover Farms housing development in 1982. Glen H. Doran and David N. Dickel directed excavations of the site for 17 months between 1984 and 1986 (Purdy 1991). The Windover site consists of a shallow pond with peat sediments in which burials were placed during the Early and Middle Archaic periods. Radiocarbon dates from twenty-seven samples, including human bone, peat, wooden stakes, and a gourd rind, provided radiocarbon dates which suggest that the burials date to approximately 7400 radiocarbon years B.P. (uncorrected). Interpretation of these samples has led to the suggestion that burial of human remains in the pond may have started around 8000 years B.P. and continued until approximately 7000 years B.P. (Tuross et al. 1994:290). Stable isotope and paleobotanical studies indicate that the individuals living at the site practiced a hunter-gatherer subsistence strategy, consuming riverine animals and seasonally available plants. Analysis of stable nitrogen isotopes suggests that the diet focused on such animals as duck, turtle, and catfish rather than deer and other terrestrial species. The paleobotanical evidence implies that the plant portion of the diet included the seeds of fleshy fruits, such as elderberry (Tuross et al. 1994). These results suggest complex exploitation of the surrounding environment by the people at Windover. One of the most notable features of the Windover site is the incredible preservation of both artifacts and human remains. Several organic materials were recovered, including wooden artifacts, textiles, incised bone objects, tools made from deer antlers, faunal remains, and plants (Adovasio et al. 2002; Penders 2002). At most archaeological sites with this early date, items such as wood and textiles would not have survived. The stable anaerobic environment and neutral pH in the Windover pond facilitated preservation of organic materials. The burials were placed at a shallow level in the pond with little oxygen, since all of the free oxygen was used by the decomposing plant materials on the bottom of the pond. The peat also had a relatively stable, neutral pH, possibly because of buffering by freshwater snail shells rich in calcium carbonate (Doran 2002). These ideal conditions led to the preservation of a wide array of organic materials.

Excavations at the site recovered an exceptionally well-preserved skeletal collection of at least 168 individuals, one of the oldest skeletal collections of this size in the New World. The number of individuals recovered from Windover is particularly significant because these individuals comprise almost half of the individuals in North America found in contexts predating 7000 years B.P. (Doran 2002). The skeletal collection includes a relatively equal number of males and females. The individuals represent a variety of age groups, ranging from infants to individuals older than 65 years of age. Prehistoric skeletal samples often have fewer subadults than would be expected, possibly because of preservation problems or cultural practices. The age distribution of the Windover skeletons does not show evidence of this problem, since 52% of the sample consists of subadults below 18 years of age (Purdy 1991). The large number of subadults in the collection will facilitate the epidemiological goal of this thesis, providing an adequate sample for assessment of the age of onset of periodontal disease. The exceptional preservation of alveolar bone and teeth in many of the skeletons also makes this collection ideal for research on periodontal disease, which requires measurements of intact bone. Since a well- preserved skeletal collection of this size and antiquity is rare, this study will provide information on the epidemiology of periodontal disease in one of the earliest New World populations studied thus far. Sample The skeletal sample (n = 76) used in this analysis included all individuals from Windover that had well-preserved maxillae with fully erupted permanent second molars. The sex and age estimates for these individuals were obtained from the Windover database, which used the sex and age methods in Buikstra and Ubelaker (1994). Sex was assessed using nonmetric pelvic and cranial traits as well as femoral and humeral head dimensions. Individuals were aged by examination of the pubic symphysis in combination with dental attrition scores specific to the Windover population. If these two methods of age estimation differed, the age based on attrition was used (Berbesque and Doran 2008). For the purposes of this study, individuals were grouped into ten year age cohorts. The sample included a similar number of males and females, as well as a small number of probable males and probable females (Table 1). Several of the individuals were subadults (n = 10) whose sex could not be determined. The sample included a range of ages, with the largest number of individuals between 40 and 49 years. Most age groups included a relatively equal

8 number of males and females. Although the entire sample included 76 individuals, not all individuals could be analyzed for every dental characteristic. Therefore, the sample size for each characteristic is reported in the results section.

Table 1: Demography of the Windover sample. Age Male Probable Male Female Probable Female Subadult Total 10-19 years 0 0 3 0 10 13 20-29 years 4 1 7 1 0 13 30-39 years 5 0 3 1 0 9 40-49 years 11 3 10 0 0 24 50-59 years 4 0 4 1 0 9 60+ years 3 0 5 0 0 8 Total 27 4 32 3 10 76

Dental Disease Assessment Periodontal disease was assessed in the Windover sample using a combination of measurements and qualitative examinations of the alveolar bone condition. First, digital sliding calipers were used to measure the distance from the cement-enamel junction of each tooth to the alveolar crest of the bone (CEJ-AC) in millimeters (Figure 1). The cement-enamel junction refers to the point on the tooth where the crown, which is coated with enamel, meets the root, which is coated with cement (Hillson 1996). The alveolar crest is the ridge-like cortical bone located around the subgingival surface of the teeth (Hildebolt and Molnar 1991). The measurement of the CEJ-AC distance provides an estimate of the amount of alveolar bone loss that may have occurred due to periodontal disease (Davies et al. 1969). The distance between the cement-enamel junction and the alveolar crest for a healthy individual is typically 1-2 mm (Stoner 1972; Strohm and Alt 1998). According to modern clinical studies, a distance greater than 2 mm indicates that the individual may have had periodontal disease (Fyfe et al. 1993; Stoner 1972). The CEJ-AC distance for a healthy periodontium in the Windover population was established by examining individuals between ten and twenty years of age because the molars of these individuals would have erupted recently. The CEJ-AC distances for these teeth therefore provide the best indication of the distance at the time of eruption, prior to the onset of 9 periodontal disease. The individuals in this age bracket exhibited CEJ-AC distances between 1-3 mm. Based on this result, individuals in the Windover population with CEJ-AC distances greater than three millimeters may have had periodontal disease. The distance from the cement-enamel junction to the alveolar crest was measured for all maxillary molars. The bone surrounding the molars often exhibits better preservation in archaeological remains than the alveolar bone associated with the anterior dentition, facilitating the assessment of periodontal disease in a larger number of individuals. The assessment of bone loss in the maxillary molars provides an indication of all individuals who experienced periodontal disease, since bone loss associated with periodontal disease typically occurs first in these molars (Hillson 1996; Watson 1986). For the second and third maxillary molars, CEJ-AC measurements were recorded at the center of the tooth for both the buccal and lingual sides (Figure 1). The first maxillary molar has three widely divergent roots (Hillson 1996), which were each measured separately. The measurements for the left and right first maxillary molars therefore included the mesiobuccal, distobuccal, and lingual roots of the teeth. The examination of these measurements for all maxillary molars revealed overall patterns of alveolar bone loss in the Windover population. After measuring the CEJ-AC distance, the condition of the alveolar bone was assessed by recording the presence of alveolar bone defects associated with the maxillary molars. The presence or absence of dehiscences and fenestrations in the alveolar bone associated with each maxillary molar was recorded. Dehiscences and fenestrations both refer to missing portions of alveolar bone that expose the root surface of the tooth. These two defects are differentiated based on the amount of the root surface exposed. Dehiscence describes a loss of the buccal or lingual alveolar bone, which completely denudes the root surface of the tooth (Rupprecht et al. 2001) (Figure 2). Fenestrations are defects that only expose a small portion of the root surface, often near the apex of the root (Strohm and Alt 1998) (Figure 3). Dehiscences and fenestrations can be a variation in normal bone structure rather than a consequence of periodontal disease, but the presence of these features indicates localized resorption of alveolar bone (Schluger et al. 1977:41). The recording of dehiscences and fenestrations therefore provided one method for examining the alveolar bone condition.

Figure 1: Buccal measurements of the distance from the cement-enamel junction to the alveolar crest. The arrows show the location of measurements for the first, second, and third maxillary molars. The abbreviations are as follows: B = buccal, DB = distobuccal, and MB = mesiobuccal.

Figure 2: Dehiscence on the maxillary left first molar of individual #95.11.

Figure 3: Fenestration on the maxillary right second molar of individual #36.504.

The condition of the alveolar bone was further assessed through visual inspection. In the absence of periodontal disease, the alveolar crest has a smooth knife-edged contour. Periodontal disease causes the resorption of the cortical plate at the alveolar crest, revealing the porous cancellous bone and creating a more rounded alveolar margin (Clarke et al. 1986; Clarke and Hirsch 1991). When the bone was adequately preserved, the condition of the alveolar crest was recorded as smooth and knife-edged or porous and rounded. When an individual exhibited CEJ- AC measurements greater than 3 mm as well as a porous and rounded alveolar crest, these factors were recorded as strong evidence of periodontal disease. In addition, the alveolar bone was examined for any indication of antemortem tooth loss in the maxillary molars. In advanced periodontal disease, all of the bone supporting the teeth is resorbed, leading to tooth loss. After exfoliation of the teeth, the gingival tissue heals and the

12 bone in the tooth socket is remodeled. Antemortem tooth loss can be identified in archaeological specimens by the presence of alveolar resorption and evidence of bony remodeling of the tooth socket after loss of the tooth (Larsen 1997). Loss of a tooth before death can be differentiated from postmortem loss of the tooth because loss of the tooth after death results in an empty tooth socket, which lacks evidence of remodeling. Antemortem loss of teeth may indicate advanced periodontal disease, which would eventually cause exfoliation of the teeth. However, dental caries can also lead to antemortem tooth loss by progressively demineralizing and eventually destroying the tooth (Soames and Southam 2005). Therefore, antemortem tooth loss was only used as additional support for the presence of periodontal disease in individuals with CEJ-AC distances of over 3 mm and unhealthy alveolar bone. The evaluation of periodontal disease prevalence can be complicated by two factors: continuous eruption and pulpal-alveolar lesions. Teeth may continue to erupt throughout life to maintain proper occlusion despite severe attrition (Clarke and Hirsch 1991; Danenberg et al. 1991). The continuous eruption of teeth will increase the CEJ-AC distance, since greater root exposure will occur as the tooth erupts further out of the alveolar bone. This issue was addressed by recording the degree of attrition on the maxillary molars using the wear stages published in Scott (1979). The Scott method divides the tooth into four quadrants and assigns a wear score between 1 and 10 for each quadrant, adding these together for a total tooth wear score between 4 and 40. The wear scores indicate the amount of enamel left on the tooth, with a score of 1 for a quadrant with all enamel remaining and a score of 10 for a quadrant in which all enamel has been worn away. The decision about whether a CEJ-AC distance greater than 3 mm should be attributed to periodontal disease or continuous eruption was made based on the wear stage for each tooth. If an individual had severe attrition as well as a CEJ-AC distance exceeding 3 mm, the dentition was classified as exhibiting probable periodontal disease. Definite periodontal disease was only recorded in individuals without extreme occlusal attrition. Pulpal-alveolar lesions can also be mistaken for periodontal disease. The pulp of the tooth can become necrotic due to caries, severe attrition, or trauma, resulting in an abscess (Hildebolt and Molnar 1991). Abscesses develop from the accumulation of pus, which often leads to a hole in the bone near the root apex to relieve pressure (Hillson 1996). This hole remains in the alveolar bone of the individual after death, allowing postmortem diagnosis of

Figure 4: Abscess associated with the maxillary right first molar of individual #74.42. Severe attrition and pulp exposure on the occlusal surface of this molar indicates that the loss of bone is due to an abscess.

abscesses (Figure 4). In some cases, extensive bone loss may occur because of abscesses (Hildebolt and Molnar 1991). The misinterpretation of pulpal-alveolar lesions as periodontal disease was avoided by recording the presence of pulp chamber exposure on the tooth and the presence of abscesses for each maxillary molar. If a tooth had a great degree of pulp chamber exposure or an abscess, bone loss around this tooth was attributed to pulpitis rather than periodontal disease. After recording all of the above factors, a final visual assessment of periodontal disease was made for each individual. Individuals with CEJ-AC measurements less than 3 mm and healthy alveolar bone were classified as having ―no periodontal disease‖ (Figure 5). For individuals with CEJ-AC measurements greater than 3 mm, the condition of the alveolar bone was examined, as mentioned above. Individuals exhibiting bone loss and unhealthy alveolar

14 bone were classified as having periodontal disease. Antemortem tooth loss in some of these individuals provided additional evidence for periodontal disease. The individuals with these characteristics were classified in one of three categories: slight to moderate periodontal disease, moderate to severe periodontal disease, or probable periodontal disease. ―Slight to moderate periodontal disease‖ was recorded when individuals had only minimal bone loss, with CEJ-AC measurements typically around 4 or 5 mm (Figure 6 and Figure 7). Individuals with greater CEJ-AC measurements were generally classified as having ―moderate to severe periodontal disease‖ (Figure 8 and Figure 9). Since the amount of bone loss often varies between different teeth in the same individual, however, the final assessment of periodontal disease involved a visual assessment of all maxillary molars as well as examination of the measurements. An individual was classified as having ―probable periodontal disease‖ if the dentition exhibited the characteristics of periodontal disease, but had one of three complicating factors: an insufficient portion of the alveolar bone was preserved, the molars exhibited severe attrition, or an abscess was present (Figure 10). If the alveolar bone was extensively damaged, the periodontal disease score was recorded as unobservable and the individual was excluded from the periodontal disease sample. In addition to periodontal disease, the presence and amount of dental calculus were recorded for all maxillary molars. These features were recorded separately for the buccal and lingual surfaces of each tooth. The amount of calculus was classified as none (0), small (1), moderate (2), or large (3), using the system in the Standards for Data Collection from Human Skeletal Remains (Buikstra and Ubelaker 1994), which is based on Brothwell (1981). Although some teeth exhibited dark stains that might have resulted from an antemortem calculus deposit, calculus was only recorded when a deposit still remained on the tooth. Other Florida Sites The Windover periodontal disease and dental calculus results were compared with the results of dental disease studies from five other sites. All of these sites are located in Florida and were occupied by populations practicing a hunter-gatherer subsistence strategy. The specific sites were chosen based on the availability of periodontal disease or dental calculus results from previous studies. The examination of periodontal disease in all of these populations will facilitate the identification of possible factors that might affect the development and progression

Figure 5: Left maxillary premolars and molars exhibiting “no periodontal disease” in an individual approximately 19 years of age (#69.1).

Figure 6: Right maxillary premolars and molars exhibiting “slight to moderate” periodontal disease in an individual approximately 42 years of age (#501.6).

Figure 7: Left maxillary premolars and molars exhibiting “slight to moderate” periodontal disease in an individual approximately 53 years of age (#121.23).

Figure 8: Right maxillary molars exhibiting “moderate to severe” lingual bone loss due to periodontal disease. The individual is approximately 38 years of age (#57.077).

Figure 9: Extensive alveolar bone loss due to severe periodontal disease in an individual approximately 38 years of age (#82.3).

of periodontal disease. Based on these results, this study will contribute to paleoepidemiology by suggesting possible directions for future research on periodontal disease. Manasota Key Cemetery (8SO1292) The Manasota Key Cemetery is located on a barrier island off the west coast of Florida in Sarasota County (Table 2). The site was discovered during construction work, and subsequently excavated by W. A. Cockrell. The Manasota Key people may have been a population ancestral to the Calusa people, or a distinct cultural group. Four radiocarbon dates were obtained for the site using samples of shell and bone, providing a mean date of about 1730 years B.P. These

Figure 10: Right maxillary molars exhibiting “probable periodontal disease” in an individual approximately 58 years of age (#93.3). Antemortem loss of the right maxillary second molar and alveolar resorption throughout the maxilla indicates that the individual probably had periodontal disease. Postmortem damage to the alveolar bone of the left maxilla prevents definitive assessment of periodontal disease.

dates indicate that the site was used during the Woodland period, specifically the Caloosahatchee I cultural period. According to David Dickel (1991:2), the occupants of the site were probably semi-sedentary hunter-gatherers, with a diet of marine resources, small game, and gathered plants. The consumption of marine resources was one of the main dietary differences between the Manasota Key population and the Windover population. The skeletal remains recovered from the site were in poor condition due to bioturbation and demineralization of the bone. A minimum of 120 individuals were excavated from the site, including 79 adults as well as 41 subadults below 18 years of age. The adult individuals include 24 males, 17 females, and 38 individuals of indeterminate sex. Age estimates suggested that none of the adults were over 38 years of age, though Dickel (1991:31) points out that this may be due to methodological limitations rather than an actual lack of older individuals. Since periodontal disease in modern populations tends to be correlated with age (Schluger et al. 1977), the relatively young age of the individuals in the Manasota Key sample must be recognized when using this site for comparative purposes.

Highland Beach (8PB11) The Highland Beach site consists of a burial mound that was discovered approximately 200 meters from the Atlantic Ocean in the town of Highland Beach, Florida. The site was probably a secondary burial site, which might have been associated with a settlement located approximately 4 kilometers south of the burial mound. Since the site was disturbed, many of the skeletons were fragmentary or commingled. In addition, many teeth had been lost postmortem. Based on the cultural artifacts found, the site was dated to approximately 950 years B.P. The inhabitants of the site are thought to have practiced a hunter-gatherer subsistence strategy, consuming marine resources, small animals, and wild plants (Isler et al. 1985). Although the site dates to a later time than Windover, the subsistence strategy was relatively similar. The primary difference between the diets of these two populations was the exploitation of marine resources at Highland Beach. Fort Center (8GL12) The Fort Center site is located seven kilometers from Lake Okeechobee in Glades County, in the Okeechobee Basin cultural area. The site was excavated from 1966 to 1971 by William H. Sears with the assistance of field crews from Florida Atlantic University, Colgate University, and the University of Florida. The skeletal remains from Fort Center date to approximately 1450 years B.P. (Shaivitz 1986). During this time, the site was thought to be a ceremonial complex, which included a low mound with a probable charnel house (Mound B), a living area on Mound A, and an artificial charnel pond with a platform on which bundled burials were placed. At some point, the platform burned and collapsed into the pond, causing 300 bundled burials to fall into the pond. Later, 150 of the burials were removed from the pond and interred on top of Mound B, while the other 150 were left in the pond (Purdy 1991; Sears 1982). The human skeletal remains recovered during the excavations included 121 individuals. The sample consisted of 43 males, 50 females, 11 adults of indeterminate sex, and 17 subadults of indeterminate sex (Purdy 1991). Analysis of the skeletal remains by Patricia Shaivitz (1986) indicated that these individuals might have been high status residents of a ceremonial complex. The people at the Fort Center site are thought to have practiced a hunting and gathering subsistence strategy. The foods consumed included reptiles, fish, and mammals (Shaivitz 1986). Sears (1982) argues that the Fort Center people obtained some of their food through agriculture. His argument for agriculture is based on the presence of maize pollen in a small number of

20 coprolites found at the site. However, the lack of artifacts associated with the processing of plants, macroplant corn remains, or ethnographic accounts describing the cultivation of corn suggests that the practice of agriculture at the site is unlikely (Purdy 1991). Unless further evidence of agriculture at Fort Center is discovered, the diet at the site will be assumed to consist of hunting of animals and gathering of plant foods.

Table 2: Florida archaeological sites discussed in this thesis. Site Name and Number Radiocarbon Minimum Number References Years BP of Individuals Bird Island (8DI52) 4570 36 Stojanowski 1997 Fort Center (8GL12) 1450 300 Sears 1982; Shaivitz 1986 Highland Beach (8PB11) 950 108 Isler et al. 1985 Manasota Key (8SO1292) 1730 120 Dickel 1991 Republic Groves (8HR4) 6520 – 5745 37 Saunders 1972 Windover (8BR246) 7400 168 Doran 2002

Republic Groves (8HR4) The Republic Groves site is located in Hardee County in central Florida. The majority of the radiocarbon dates indicate that the site dates to approximately 5745 to 6520 years B.P. (Purdy 1991). This early date suggests that the population at this site likely practiced a hunting and gathering subsistence strategy. The human skeletal remains recovered from the site include a minimum of 37 individuals. The sample included six children, seven young adults, and thirteen early to late middle aged adults. The age of the remaining individuals could not be determined. The sex of 19 individuals could be determined, with a result of ten males and nine females (Purdy 1991). Periodontal disease in adults from the Republic Groves site was discussed by Saunders (1972). Bird Island (8DI52) The Bird Island site is located on an island in the Gulf of Mexico in Dixie County in north Florida. Surface collections and excavations of the site have taken place periodically since the 1950s. A radiocarbon sample of human bone from the site was dated to 4570 years B.P.,

21 suggesting that the burials date to the Late Archaic period (Stojanowski 1997). Much of the skeletal material was exposed on the surface for extended periods of time after eroding out of primary context during storms. These conditions led to poor preservation of the skeletal remains, limiting the information that could be collected from the sample (Stojanowski and Doran 1998). The skeletal sample includes at least four subadults of indeterminate sex as well as 32 adults, divided into five females, seven males, and 20 individuals of indeterminate sex (Stojanowski 1997). Although poor preservation precluded analysis of periodontal disease in this sample, the prevalence of dental calculus in the sample was recorded.

CHAPTER 3 - RESULTS

Introduction This chapter discusses the results of the dental health analysis of the Windover sample. The first part of the chapter describes the periodontal disease results by reporting the tooth surfaces with the most alveolar resorption, the prevalence of periodontal disease in the entire sample and in different age and sex groups, and the amount of antemortem tooth loss. The next part discusses the prevalence of dental calculus in the Windover sample, and then analyzes the relationship between dental calculus and periodontal disease. After this, the prevalence of dehiscences and fenestrations is described, and the relationship between these alveolar bone defects and periodontal disease is assessed. The periodontal disease data are then compared with dental caries and abscess data from Windover. The final section examines the prevalence of periodontal disease in two Windover lineages that are hypothesized to be genetically distinct. Locations of Alveolar Resorption The measurements of the distance between the cement-enamel junction and the alveolar crest reveal which maxillary molars in the Windover sample were affected most by periodontal disease. The tooth surfaces with the greatest amount of bone loss were identified by calculating the mean measurement for each location, excluding tooth surfaces with dehiscences (since these may indicate bone missing due to morphological variation rather than periodontal disease). In the Windover sample, the location with the greatest amount of bone loss was the lingual surface of the first molars (Table 3). The next location exhibiting a great deal of bone loss was the lingual surface of the second molars. The lingual surface of the third molars also had bone loss indicating periodontal disease, with mean CEJ-AC measurements greater than the established healthy distance of 3 mm. Another location particularly affected by periodontal disease was the buccal surface of the maxillary first molar, including both the mesiobuccal and the distobuccal roots. The mean measurements for RM1 mesiobuccal, RM1 distobuccal, and LM1 mesiobuccal are all greater than 3 mm. The buccal surfaces of the maxillary second and third molars were less affected by bone loss, with mean measurements of less than 3 mm for both the left and right teeth.

Table 3: Mean CEJ-AC measurements of maxillary molars in the Windover sample, ordered from highest to lowest. The tooth surface labels are as follows: R = right, L = left, M = molar, and the number 1, 2, or 3 indicates the molar number. Tooth Surface n Mean (mm) Minimum Maximum Standard Standard error deviation RM1 lingual 56 4.73 0.92 11.51 0.33 2.48 LM1 lingual 58 4.60 0.85 10.06 0.29 2.22 RM2 lingual 60 4.53 0.86 11.92 0.26 2.02 RM1 mesiobuccal 41 4.42 0.94 15.25 0.48 3.08 LM2 lingual 60 4.27 0.79 8.49 0.23 1.81 RM3 lingual 40 3.93 1.32 7.88 0.26 1.63 RM1 distobuccal 43 3.59 0.82 12.05 0.40 2.59 LM3 lingual 42 3.58 0.49 6.49 0.22 1.44 LM1 mesiobuccal 39 3.15 0.97 8.19 0.26 1.62 RM2 buccal 56 2.89 0.45 14.65 0.27 2.04 LM1 distobuccal 46 2.83 0.51 7.63 0.22 1.49 RM3 buccal 38 2.51 0.76 4.56 0.13 0.78 LM2 buccal 53 2.47 0.25 4.68 0.14 1.04 LM3 buccal 38 2.32 0.52 4.34 0.12 0.77

These results suggest that, in the Windover population, periodontal disease causes the greatest amount of localized bone loss on the lingual surface of the first molar. The lingual surfaces of the maxillary second and third molars showed the next greatest amount of mean bone loss, suggesting that the lingual surfaces of the maxillary molars were the most affected by periodontal disease. The buccal surfaces of the teeth generally showed less alveolar bone loss. The buccal surfaces of the first molar, however, had a larger amount of bone loss. This result suggests that both the buccal and lingual surfaces of the first molar were the most affected by periodontal disease. The maxillary second and third molars exhibited less bone loss than the maxillary first molars. The difference in bone loss between the first molar and the second and third molars might be explained by the tooth eruption sequence. The first molar erupts around the age of six, while the second molar erupts around the age of twelve and the third molar usually erupts around the age of eighteen. Since the first molar erupts considerably earlier than the second or third molar, this tooth begins to be exposed to stressors, such as plaque and calculus, at an earlier age. The first molar therefore is at risk of bone loss associated with periodontal disease for a longer period of time than the second or third molar.

The results presented in Table 3 indicate that many individuals in the Windover sample experienced bone loss greater than 3 mm on several tooth surfaces. This amount of bone loss generally indicates some degree of periodontal disease. The next section will describe the actual percentage of the sample affected by periodontal disease. Prevalence of Periodontal Disease in the Entire Windover Sample Periodontal disease affected a large number of the individuals buried at the Windover site (Figure 11). Of the fifty-six individuals with maxillae that could be assessed, only twenty-five individuals lacked evidence of alveolar resorption associated with periodontal disease. The remaining thirty-one individuals (55.4% of the individuals examined) exhibited evidence of some degree of periodontal disease, based on the presence of alveolar bone loss, unhealthy alveolar bone and, in some cases, antemortem tooth loss. Of the individuals with periodontal disease, the largest number (n = 16) exhibited slight to moderate periodontal disease. Only five individuals (8.9%) had alveolar bone loss indicative of moderate to severe periodontal disease. Ten individuals (17.9%) were classified as having probable periodontal disease, without further classification of the severity of the disease. These individuals most likely suffered from periodontal disease, but inadequate preservation, severe attrition, or abscesses prevented definitive assessment (see Chapter 2 for details). Since the probability that these individuals had periodontal disease is high, this group is included in the estimated percentage of the sample affected by the disease. These results suggest that over half of the Windover population may have suffered from periodontal disease. However, only a small number of individuals probably experienced severe periodontal disease, which would have eventually resulted in antemortem tooth loss. The majority of individuals would have experienced slight alveolar bone resorption that possibly caused discomfort but did not threaten tooth loss. Since periodontal disease does not always progress to a severe stage, this result alone cannot predict whether these individuals would ever have experienced a more severe form of periodontal disease with increasing age. The association between periodontal disease and age in the Windover sample will be discussed in the next section.

Figure 11: Prevalence of periodontal disease in the entire Windover sample. The numbers indicate the percentage of the total sample (n = 56) with each condition.

Prevalence of Periodontal Disease by Age Periodontal disease is strongly associated with increasing age in the Windover sample. Figure 12 shows the prevalence of periodontal disease in each ten year age group. None of the thirteen individuals between the ages of 10-19 years exhibited any evidence of periodontal disease. All individuals in this age bracket had healthy CEJ-AC distances of less than 3 millimeters. Between the ages of 20-29 years, three of eleven individuals exhibited periodontal disease. This result suggests that the earliest age of onset of periodontal disease in the Windover population was likely sometime between 20-29 years of age. The individuals in this age group with periodontal disease only experienced a small degree of alveolar resorption. Two of these individuals had slight to moderate periodontal disease, and one individual had probable periodontal disease. Most of the other individuals between the ages of 20 and 29 years exhibited slight lingual bone loss, with CEJ-AC measurements between 3-4 millimeters. This degree of bone loss was insufficient to confidently label as periodontal disease, but could have been an indicator of very early or developing periodontal disease. These individuals could have 26 experienced gingivitis as a precursor to more severe periodontal disease but, since the gum tissue was not preserved, this condition cannot be assessed. The observation that individuals between the ages of 20 and 29 years had slight lingual bone loss or slight to moderate periodontal disease does, however, suggest that this may have been the earliest age of onset of periodontal disease in the Windover population. The results for individuals between 30-39 years of age provide additional support for the idea that bone loss began between 20-29 years. By 30-39 years, all individuals had periodontal disease ranging from slight to severe. Four individuals in this age group exhibited slight to moderate periodontal disease and two individuals showed moderate to severe periodontal disease. These results suggest that individuals at Windover may have started to experience bone loss in their late twenties, with bone loss becoming more severe by the time they reached their thirties. Since bone loss associated with periodontal disease occurs intermittently, however, the loss of bone should not be regarded as a constant problem affecting individuals in their twenties and thirties. The loss of bone may have occurred in sporadic intervals throughout these decades of life. The onset of periodontal disease in the Windover population in the twenties and thirties is similar to the age of onset of periodontal disease in modern populations (Schluger et al. 1977). In the 40-49 year age group, most individuals had some degree of periodontal disease. Three of fifteen individuals, however, exhibited no evidence of periodontal disease. This result shows that a small percentage of the population was not affected by periodontal disease, even at older ages. In the 40-49 year age group at Windover, slight to moderate periodontal disease was relatively common, while severe cases of periodontal disease were rare. Eight of the individuals in the 40-49 year age group had slight to moderate periodontal disease, and only one individual had moderate to severe periodontal disease. Three other individuals were classified as having probable periodontal disease. By the time individuals in the Windover population reached the age of 49 years, most of them had experienced some alveolar resorption. All individuals between the ages of 50-59 years exhibited symptoms of periodontal disease. Four of the individuals were classified as having probable periodontal disease, while two individuals had slight to moderate periodontal disease. Only one individual was classified as manifesting moderate to severe periodontal disease. These results suggest that individuals in the Windover population between 50 and 59 years of age typically suffered from periodontal disease.

Figure 12: Prevalence of periodontal disease by age group.

The majority of individuals over 60 years also experienced bone loss from periodontal disease. Two of the individuals in this age group had probable periodontal disease, and one individual had moderate to severe periodontal disease. However, one of the individuals still had healthy alveolar bone, without any evidence of periodontal disease. Therefore, periodontal disease does not appear to have affected the entire population, regardless of age. Since the sample size for this age group is small, generalizations about the effects of periodontal disease on individuals in the Windover population over the age of 60 years cannot be made. Unfortunately, many of the individuals buried at Windover died prior to this age, and few of the individuals who lived beyond the age of 60 years had intact maxillae. The results from the few maxillae that could be measured indicate that at least some individuals over the age of 60 years had periodontal disease. The proposed association between age and periodontal disease is also supported by the results of a statistical test of the independence of these variables. Using a G Test of Independence (Sokal and Rohlf 1981), the association between these two variables was found to

28 be highly significant (G=58.194, df=15, P=0.000). Therefore, periodontal disease is strongly associated with age in the Windover sample. Other epidemiological studies have also found an association between periodontal disease and increasing age (Albandar 2002). For modern populations, Schluger et al. (1977:78) report a positive correlation between both the extent and prevalence of gingivitis and periodontal disease and increasing age. Periodontal disease affected only 10 percent of the individuals between the ages of 18 and 25 years. By the age of 65-74 years, 60 percent of individuals were affected by periodontal disease. These results show a dramatic increase in periodontal disease with increasing age. Goldman and Cohen (1973:58) also find increasing periodontal disease severity with increasing age from 18 to 79 years in modern populations. The results from Windover indicate that the trend of periodontal disease increasing with age was also present in a population from thousands of years earlier. Prevalence of Periodontal Disease by Sex When the prevalence of periodontal disease was examined in relation to sex, no association was found between these two variables. This sample (n=46) included only individuals whose sex could be assessed as male, probable male, female, or probable female. Subadults were excluded from this sample, since the sex of young individuals is difficult to accurately determine. Figure 13 shows the prevalence of periodontal disease by sex in the Windover sample. Both males and females experienced all levels of severity of periodontal disease, and both groups included some individuals without any noticeable symptoms of periodontal disease. The percentage of individuals without periodontal disease was higher in females than in males. Out of eighteen males, only four (22.2%) lacked any evidence of periodontal disease. In contrast, eleven (47.8%) out of twenty-three females had healthy alveolar bone without any indication of periodontal disease. These results suggest that fewer females than males may have suffered from periodontal disease in the Windover population. Males also may have been affected by more severe periodontal disease than females at Windover. In the entire Windover sample, only five individuals suffered from moderate to severe periodontal disease. Four out of these five individuals were male, with only one female experiencing moderate to severe periodontal disease. Six males and all three of the individuals classified as probable males had slight to moderate periodontal disease, indicating that many of the males at Windover experienced at least some alveolar resorption associated with periodontal

Figure 13: Prevalence of periodontal disease by sex.

disease. Four males were classified as having probable periodontal disease, which suggests that these individuals also experienced some symptoms of the disease. Overall, 77.8% of males and 100% of probable males at Windover exhibited some degree of periodontal disease. In contrast, 52.2% of females and 100% of probable females at Windover had some degree of periodontal disease. Five of the total twenty-three females had probable periodontal disease, and six females exhibited slight to moderate periodontal disease. As mentioned above, only one female had alveolar bone loss extensive enough to be classified as severe periodontal disease. The two probable females in the sample had probable periodontal disease and slight to moderate periodontal disease, respectively. The results suggest that a smaller number of females than males may have suffered from periodontal disease at Windover, and that periodontal disease among females was generally less severe than among males. The difference in periodontal disease prevalence between males and females at Windover might be explained by cultural practices. Males and females could have consumed different foods or varying amounts of the same foods. Dietary variation between males and females might

30 indicate that access to particular foods was restricted based on sex. Discrepancies in the type and texture of foods consumed might have led to periodontal disease differences between males and females at Windover (Lavelle and Moore 1969; Newman and Levers 1979). Thus, the observed variation in periodontal disease between the sexes at Windover might reflect differential access to foods. Although the results seem to indicate an association between sex and periodontal disease at Windover, analysis of the results using a G Test of Independence found the proposed association to be statistically insignificant (G=14.187, df=9, P=0.116). Therefore, the potential association between sex and periodontal disease discussed above was not statistically significant. The lack of statistical significance could be due to the relatively small size of the Windover sample, which only included forty-six individuals. The results of other studies on periodontal disease suggest that the relationship between sex and periodontal disease is complex. Studies of the United States population conducted between 1960 and 1962 found that men were more susceptible to periodontal disease than women (Goldman and Cohen 1973). In modern populations, juvenile periodontal disease is also known to affect males more often than females. This form of periodontitis begins during the teenage years and is found four times more often in males than in females, suggesting that males are more susceptible to at least certain types of periodontal disease (Hildebolt and Molnar 1991). The results of studies of modern populations therefore indicate that sex may be associated with periodontal disease. Studies conducted on archaeological populations, however, have not been as conclusive about the relationship between sex and periodontal disease. At Manasota Key Cemetery, Dickel (1991:91-92) found that 67% of males and 82% of females were affected by periodontal disease. These differences were, however, found to be statistically insignificant. Therefore, there may not have been an association between sex and periodontal disease at Manasota Key Cemetery. In contrast, Isler et al. (1985:142) found that severe alveolar resorption was more common in males than females at Highland Beach. Slight alveolar resorption of up to 3 mm was exhibited by 71% of females and only 46% of males. More severe alveolar resorption of 3 mm to 6 mm was found in 44% of males and 19% of females. These results were statistically significant, suggesting that males were more susceptible to severe alveolar bone loss than females at Highland Beach.

Based on the results of these studies, the relationship between sex and susceptibility to periodontal disease may not be as predictable as the relationship between increasing age and periodontal disease. The possible trend of more severe periodontal disease in males in the Windover population was similar to the patterns found at Highland Beach and in modern populations, but different from the pattern found at Manasota Key Cemetery. Therefore, the relationship between sex and periodontal disease is highly complex and may not be uniform across populations. Antemortem Tooth Loss The examination of antemortem tooth loss provides additional information about the extent of periodontal disease in the Windover sample. Antemortem tooth loss can result from either severe periodontal disease or dental caries. The cause of antemortem tooth loss cannot be determined from skeletal remains, but extensive loss of adjacent alveolar bone may suggest that periodontal disease led to tooth loss. This study found minimal antemortem tooth loss in the maxillary molars in the Windover sample (Table 4). Only seven out of 76 individuals (9.2%) displayed evidence of alveolar bone remodeling from antemortem tooth loss of the maxillary molars. The teeth lost included all molars except the maxillary left second molar. Loss of the maxillary third molar appears to have been the most common, with four of the seven individuals losing this tooth. Only one individual lost more than one maxillary molar, displaying loss of both the maxillary first molars. Overall, loss of the maxillary molars was uncommon in the Windover population. The small percentage of individuals affected by antemortem tooth loss suggests that periodontal disease in the Windover sample was relatively mild. Even though thirty-one individuals exhibited symptoms of periodontal disease on the maxillary molars, only seven individuals lost any of these molars antemortem. These results indicate that alveolar bone resorption from periodontal disease was rarely severe enough to cause tooth loss. As mentioned above, only five individuals in the Windover sample exhibited moderate to severe periodontal disease. The limited number of individuals with antemortem tooth loss thus supports the idea that most individuals in the Windover population experienced either no periodontal disease or slight to moderate periodontal disease.

Table 4: Prevalence of antemortem tooth loss in the maxillary molars of the Windover sample. The left column indicates which teeth were lost, and the next column shows how many individuals lost a particular tooth. The tooth labels are as follows: L = left, R = right, M = molar, and the number 1, 2, or 3 indicates the molar number. Tooth Lost Antemortem Frequency (# of Individuals) Percent of Sample LM1 1 1.3 LM3 2 2.6 RM2 1 1.3 RM3 2 2.6 LM1 and RM1 1 1.3 None 69 90.8

Locations of Dental Calculus

Since dental plaque is often an etiological agent in the development of periodontal disease in modern populations (Soames and Southam 2005), examination of the locations of dental calculus (mineralized plaque) may provide insight into the etiology of periodontal disease in the Windover population. The locations with the largest amount of dental calculus were determined by calculating the mean score for each tooth surface. As described in the methods chapter, the amount of dental calculus was scored using the following system: 0 = none; 1 = small; 2 = moderate; 3 = large. Therefore, higher mean calculus amounts indicate a larger amount of calculus on a particular tooth surface. The results for the Windover sample are listed in Table 5. The tooth surfaces affected most by dental calculus are the buccal portions of all maxillary molars. The mean calculus scores for the buccal surfaces of the molars range from 0.51 – 0.89. The mean calculus scores for the lingual surfaces of the molars, in contrast, range from 0.17 – 0.36. These results indicate that calculus was more common on all buccal surfaces than any of the lingual surfaces. The accumulation of dental calculus on the buccal surface of the maxillary molars can be explained by the proximity of this tooth surface to the parotid duct. The locations with the largest accumulations of calculus are the surfaces of the teeth that are nearest to the openings of the major salivary gland ducts. As a result, calculus tends to accumulate in two locations: the lingual surfaces of the mandibular anterior dentition opposite the submaxillary and sublingual gland openings, and the buccal surfaces of the maxillary molars opposite the opening of the parotid duct. The distribution of calculus in these locations can be

33 confined to a specific tooth or generalized across several tooth surfaces (Shafer et al. 1974). In the sample of maxillary molars analyzed in this study, calculus appears to have been generalized across the buccal surface of all maxillary molars. The maxillary molar with the largest amount of calculus was the first molar, on both the right and left sides. The second molars were affected by calculus to nearly the same degree as the first molars. The first and second molars were the only teeth with a maximum calculus score of 3, indicating a large amount of calculus. The third molars exhibited a smaller amount of calculus than either the first or second molars. The mean calculus score for the third molars was lower, and the maximum score for these teeth was 2, indicating a moderate amount of calculus. These results are similar to the teeth most affected by alveolar bone loss (see Table 3 above), with the first molars being the most common location. This suggests that the first and second molars were affected the most by both periodontal disease and dental calculus. The third molars were a less common location for both of these dental problems, although periodontal disease and dental calculus sometimes occurred on the third molars. The results for the locations of dental calculus and periodontal disease suggest that these two dental problems may have been associated in the Windover population. The occurrence of both dental calculus and alveolar bone loss on the first and second molars indicates a possible association between these two conditions. Soames and Southam (2005) argue that dental plaque, which can mineralize to become calculus, is the main etiological agent responsible for causing chronic periodontal disease. This argument is supported by the results of epidemiological studies, which have found a positive association between dental plaque and both the prevalence and severity of periodontal disease. Dental plaque is not, however, the only risk factor for periodontal disease, as demonstrated by the above discussions of periodontal disease prevalence in relation to age and sex. The following sections will further discuss the prevalence of dental calculus in the entire Windover sample, as well as different demographic groups. This study will then assess whether there is an association between periodontal disease and dental calculus in the Windover sample.

Table 5: Minimum, maximum, and mean calculus scores for the maxillary molars in the Windover sample, ordered from highest to lowest. The tooth surface labels are as follows: R = right, L = left, M = molar, and the number 1, 2, or 3 indicates the molar number. Tooth Surface n Min. Max. Mean Standard Error Standard Deviation RM1 buccal 73 0 3 0.89 0.074 0.636 LM1 buccal 75 0 3 0.83 0.090 0.778 LM2 buccal 71 0 3 0.82 0.093 0.780 RM2 buccal 72 0 3 0.78 0.087 0.736 RM3 buccal 51 0 2 0.55 0.090 0.642 LM3 buccal 47 0 2 0.51 0.080 0.547 LM3 lingual 47 0 2 0.36 0.077 0.529 RM1 lingual 73 0 2 0.27 0.059 0.507 RM3 lingual 51 0 2 0.25 0.073 0.523 RM2 lingual 72 0 2 0.24 0.054 0.459 LM2 lingual 71 0 2 0.21 0.053 0.445 LM1 lingual 75 0 2 0.17 0.048 0.415

Prevalence of Dental Calculus in the Entire Windover Sample The majority of the Windover sample had dental calculus on at least one tooth surface. Of the 76 individuals that could be scored for dental calculus, 71 individuals (93.4%) exhibited calculus in at least one location. Although most individuals in the sample had dental calculus, large amounts of calculus were rare. The maximum calculus score was determined for all individuals by finding the highest score observed in at least one of the twelve locations that were examined for dental calculus. The number and percentage of individuals exhibiting each maximum score was then determined. The results provide an indication of the relative amounts of calculus experienced by the Windover population. The percentage of the sample with each maximum calculus score is shown in Figure 14. Only five out of seventy-six individuals (6.6%) lacked any dental calculus. The majority of individuals (41 individuals or 53.9%) in the Windover sample had a maximum amount of calculus that was scored as small, indicating that calculus was only present in minimal quantities. Another twenty-five individuals (32.9%) had a moderate amount of calculus as the maximum score, suggesting that this amount of calculus was slightly less common in the Windover sample. Only five individuals (6.6%) exhibited a large amount of calculus, which implies that most individuals in the Windover sample were not affected by severe calculus. Alternatively, the lack of large calculus deposits could reflect

35 postmortem loss of some calculus. Since postmortem loss cannot be assessed, however, these calculus amounts are assumed to be a relatively accurate indicator of premortem amounts. Therefore, the Windover population seems to have been primarily affected by small to moderate amounts of calculus. Since most individuals in the Windover sample had at least one tooth surface with dental calculus, the results were further analyzed by determining the percentage of maxillary molar tooth surfaces affected by dental calculus for each individual. The tooth surfaces analyzed included the buccal and lingual portions of each maxillary molar, for a maximum total of twelve observed tooth surfaces. Since some individuals were missing certain maxillary molars, the number of tooth surfaces observed for each individual was variable. The percentage of tooth surfaces affected by dental calculus was determined by using the following procedure for each individual (n=76):

Number of tooth surfaces with calculus x 100 = % of tooth surfaces affected by calculus Number of tooth surfaces observed

The percentages were then grouped into five different categories, based on the relative percentage of tooth surfaces affected by calculus. Individuals with 0% of tooth surfaces affected by dental calculus were classified as ―None.‖ Individuals with 1.00-24.99% of tooth surfaces exhibiting dental calculus were classified as ―Less than ¼.‖ Individuals with 25.00-49.99% of tooth surfaces affected by calculus were classified as ―¼ to less than ½.‖ Individuals with 50.00- 74.99% of tooth surfaces exhibiting calculus were classified as ―½ to less than ¾.‖ The final category included individuals with 75.00-100.00% of tooth surfaces affected by dental calculus, and these were classified as ―¾ to all.‖ These categories will be used throughout the rest of the discussion of dental calculus.

Figure 14: Percentage of the Windover sample with each maximum calculus score. The different colors reflect the amount of calculus.

The prevalence of dental calculus in the entire Windover sample is shown in Figure 15. As mentioned above, five of the seventy-six individuals (6.6%) did not have any dental calculus. An additional fifteen individuals (19.7%) had calculus on less than ¼ of their tooth surfaces. The largest number of individuals (23 individuals or 30.3%) exhibited dental calculus on ¼ to less than ½ of their tooth surfaces. The next largest category was composed of individuals with ½ to less than ¾ of their tooth surfaces affected by calculus, which included a total of 22 individuals (28.9%). Only eleven individuals (14.5%) had ¾ to all of their tooth surfaces affected by dental calculus. These results indicate that over half of the Windover sample exhibited dental calculus on ¼ to less than ¾ of their tooth surfaces. A smaller percentage of individuals had no calculus or calculus on less than ¼ of their tooth surfaces. The total percentage of the sample with calculus on over ¾ of their tooth surfaces is relatively small. Thus, most of the Windover population seems to have been affected by calculus, but calculus

37 was rarely distributed across all tooth surfaces. Often, calculus was found on one-quarter to less than three-quarters of the maxillary molar tooth surfaces in each individual.

Figure 15: Prevalence of dental calculus in the entire Windover sample. The numbers indicate the percentage of the total sample (n=76) in each calculus category. The categories listed in the legend reflect the percentage of tooth surfaces affected by calculus.

Prevalence of Dental Calculus by Age In the Windover sample, the percentage of tooth surfaces affected by calculus is associated with the age of the individual. Figure 16 shows the number of individuals who were affected by each calculus category, divided into age groups. These results show a trend of increasing calculus with increasing age. In the 10-19 year age group, one out of thirteen individuals (7.7%) did not have any dental calculus. Eleven individuals (84.6%) had less than ¼ or between ¼ and less than ½ of their tooth surfaces of affected by calculus. Only one individual (7.7%) had calculus on ½ to less than ¾ of tooth surfaces. None of the individuals in the 10-19

38 year age group had calculus on greater than ¾ of tooth surfaces. These results indicate that most individuals between 10-19 years of age had calculus on less than ½ of their tooth surfaces. By 20-29 years, individuals in the Windover sample began to exhibit more dental calculus. While three out of thirteen individuals (23.1%) did not exhibit any calculus, most of the remaining individuals had dental calculus on more than ¼ of their tooth surfaces. Only one individual (7.7%) had calculus on less than ¼ of tooth surfaces. Another five individuals (38.5%) had calculus on ¼ to less than ½ of their tooth surfaces. Three individuals (23.1%) exhibited calculus on ½ to less than ¾ of their tooth surfaces. In this age group, one individual (7.7%) had dental calculus on ¾ of tooth surfaces, indicating that the distribution of calculus across almost all maxillary molar tooth surfaces began to occur in the 20-29 year age range. In the 30-39 year age group, a larger percentage of individuals exhibited calculus on over half of their tooth surfaces. Only one out of nine individuals lacked any dental calculus. Two individuals had less than half of their tooth surfaces affected by calculus. The majority of individuals (66.6%) exhibited calculus on at least half of their tooth surfaces. Three individuals had calculus on ½ to less than ¾ of their tooth surfaces, and another three individuals exhibited calculus on ¾ or more of these surfaces. These results suggest that, by 30-39 years of age, two- thirds of individuals had calculus on over half of their tooth surfaces. By 40-49 years of age, all individuals in the Windover sample exhibited calculus on at least some of their teeth. Seven out of twenty-four individuals (29.2%) had calculus on less than ¼ of their tooth surfaces. Another eight individuals (33.3%) exhibited calculus on ¼ to less than ½ of tooth surfaces. The remaining nine individuals (37.5%) had calculus on greater than half of their tooth surfaces, with two of these individuals exhibiting calculus on ¾ to all of the surfaces observed. These results indicate that all individuals had dental calculus by 40-49 years of age, but the percentage of surfaces affected by calculus was variable. In the 50-59 year age group, all individuals exhibited some amount of dental calculus. Only two out of nine individuals (22.2%) had less than half of their tooth surfaces affected by calculus. The remaining seven individuals (77.8%) had calculus on more than half of their tooth surfaces, with three of these individuals exhibiting calculus on ¾ or more of their tooth surfaces. By 50-59 years of age, then, most individuals had calculus on more than half of their tooth surfaces.

Figure 16: Prevalence of dental calculus by age group. The different bar colors indicate the percentage of tooth surfaces affected by dental calculus.

The results for the 60+ year age group were similar, with most individuals displaying calculus on more than half of their teeth. All individuals had some amount of dental calculus, and only one out of eight individuals (12.5%) exhibited calculus on less than ¼ of their tooth surfaces. One other individual had calculus on ¼ to less than ½ of tooth surfaces. The other six individuals (75.0%) had calculus on greater than half of their tooth surfaces, with two of these individuals exhibiting calculus on ¾ to all of their tooth surfaces. These results indicate that all individuals greater than 60 years of age in the Windover sample had calculus on some of their teeth, and most of these individuals had calculus on more than half of the tooth surfaces observed. The percentage of tooth surfaces affected by calculus was found to be significantly associated with age using a G Test of Independence (G=32.516, df=20, P=0.038). The results indicate that the percentage of tooth surfaces exhibiting dental calculus increases with age. This association between calculus and increasing age was also found at the Manasota Key Cemetery. Dickel (1991:84) reports that only one individual under 17.5 years of age had any dental calculus. In contrast, approximately 51% of adults exhibited calculus, typically on more than 40 one tooth. The percentage of individuals exhibiting calculus was therefore lower than the 93.4% of individuals observed in the Windover sample. The relationship between dental calculus and age was, however, found at both of these sites. This result suggests that individuals in both early Florida populations became more susceptible to dental calculus as their age increased. The relationship between age and dental calculus in Florida hunter-gatherer populations may be more complex than these results suggest, however. In an examination of dental calculus in individuals from the Bird Island site, Stojanowski (1999:67) did not find an association between the percentage of teeth affected by dental calculus and age. These results are however, based on a small sample size and the inclusion of more individuals in the sample might have produced different results. The differing results from Bird Island, Windover, and Manasota Key Cemetery suggest that variation in the presence and amount of dental calculus between individuals may result from a combination of factors. Another possible demographic factor influencing the prevalence of dental calculus is the sex of the individual, which will be examined in the next section. Prevalence of Dental Calculus by Sex In the Windover sample, the sex of the individual does not appear to be related to the percentage of teeth affected by dental calculus. Figure 17 shows the prevalence of varying amounts of calculus by sex. The results for males and females do not show any clear patterns related to sex. Only one out of twenty-seven males (3.7%) lacked any dental calculus. Five males (18.5%) had calculus on less than ¼ of their tooth surfaces, and six males (22.2%) exhibited calculus on ¼ to less than ½ of their tooth surfaces. The largest number of males (eleven individuals or 40.7%) were affected by calculus on ½ to less than ¾ of tooth surfaces. An additional four individuals (14.8%) had calculus on ¾ or more of their tooth surfaces. The percentage of tooth surfaces affected by dental calculus for probable males ranged from less than ¼ to less than ¾ of observed surfaces. These results indicate that the percentage of tooth surfaces affected by calculus for both males and probable males was relatively variable. The results for females were similarly variable, without any clear patterns. Two out of thirty-two females (6.3%) did not have any dental calculus. Another six females (18.8%) had calculus on less than ¼ of their tooth surfaces. Ten females (31.3%) exhibited calculus on ¼ to less than ½ of tooth surfaces, and eight females (25.0%) had calculus on ½ to less than ¾ of tooth surfaces. The remaining six females (18.8%) displayed calculus on ¾ or more of their

Figure 17: Prevalence of dental calculus by sex. The different bar colors indicate the percentage of tooth surfaces affected by dental calculus.

tooth surfaces. Of the three probable females, one individual lacked any calculus and one individual had ¼ to less than ½ of tooth surfaces affected by calculus. The other probable female had ¾ or more of tooth surfaces exhibiting calculus. Based on these results, no apparent patterns emerge for females in relation to dental calculus. The percentage of females exhibiting each amount of calculus was relatively similar, suggesting that sex may not influence the distribution of dental calculus. The results of a G Test of Independence supported the above evidence that sex and dental calculus are not significantly associated (G=10.148, df=12, P=0.603). The lack of an association between these two variables has also been found at the Manasota Key Cemetery and the Bird Island site. At the Manasota Key Cemetery, Dickel (1991:84) did not find a significant difference between calculus occurrence in males and females. Approximately 58% of the males in the sample and 64% of females exhibited calculus on at least one tooth. These results suggest that calculus was relatively evenly distributed between males and females at the Manasota Key Cemetery. Similarly, Stojanowski (1999:67) found that there was not a significant relationship

42 between the presence of calculus and the sex of the individual at the Bird Island Site. The epidemiological results from Windover, Manasota Key Cemetery, and Bird Island indicate that sex is not a significant predictor of the presence or number of teeth affected by dental calculus. Relationship between Dental Calculus and Periodontal Disease The relationship between dental calculus and periodontal disease has been extensively discussed in the modern clinical literature (Albandar et al. 1998; Davies et al. 1997; Goldman and Cohen 1973; Martínez-Canut et al. 1999; Schluger et al. 1977; Shafer et al. 1974; Soames and Southam 2005). Dental plaque is now thought to be the principal etiological agent in the development of periodontal disease (Soames and Southam 2005), and calculus is defined as mineralized plaque. Studies of modern populations have found a significant association between the presence or absence of dental calculus and the type and severity of periodontal disease (Martínez-Canut et al. 1999). Calculus influences the development of periodontal disease through two mechanisms: bacterial infection and mechanical irritation. Although calculus is mineralized, various microorganisms live on the surface, allowing new plaque to form (Gaare et al. 1990; Schluger et al. 1977). The bacteria that have been isolated from human dental plaque have been shown to cause periodontal disease when placed in the mouths of other animals (Soames and Southam 2005). Thus, the relationship between the bacteria in plaque and the development of periodontal disease has been clearly demonstrated. Calculus also facilitates the development of periodontal disease by irritating the gingival tissues. When chewing places pressure on the gingiva, rough calculus deposits can cause irritation and inflammation of the gingiva. The inflammatory reaction initiates the development of a gingival pocket (Shafer 1974), which can eventually progress to more severe periodontal disease. In modern populations, then, the association between dental calculus as a bacterial and mechanical irritant and the development of periodontal disease has been definitively established. The results from the Windover sample indicate that dental calculus may also have been a major etiological factor in the development of periodontal disease in this early Florida population. In the Windover sample, dental calculus was found to be significantly associated with periodontal disease using a G Test of Independence (G=30.448, df=12, P=0.002). Individuals with more dental calculus tended to have more extensive periodontal disease. These results suggest that dental plaque has remained one of the main etiological agents responsible for the development of periodontal disease for thousands of years.

Prevalence of Dehiscences and Fenestrations The presence of dehiscences or fenestrations indicates resorption of alveolar bone, which may be considered either a normal variation in bone structure (Hildebolt and Molnar 1991; Schluger et al. 1977) or a periodontal defect (Strohm and Alt 1991). Dehiscences completely expose the root surface on the buccal or lingual side, and fenestrations expose only a small portion of the root (Rupprecht et al. 2001). This section examines the percentage of the Windover sample with dehiscences and fenestrations, and the next section looks at the prevalence of these types of alveolar bone defects in different age groups and sexes. The following section identifies the tooth surfaces most susceptible to dehiscences and fenestrations. The final section discusses the relationship between dehiscences, fenestrations, and periodontal disease in the Windover sample. Of the 76 individuals in the Windover sample, 28 individuals (36.8%) had at least one dehiscence. The other 48 individuals (63.2%) lacked any evidence of dehiscences. Fenestrations were less common, with only 14 individuals (18.4%) exhibiting at least one fenestration. The remaining 62 individuals (81.6%) did not have any fenestrations. The lower percentage of fenestrations might be attributed to one of two causes. First, Elliott and Bowers (1963) argue that fenestrations may be a transitional stage between normal bone architecture and dehiscences. In this case, fenestrations would utimately become dehiscences and the number of fenestrations could therefore be lower than the number of dehiscences. Second, fenestrations may have been more difficult to distinguish from postmortem damage than dehiscences. Small portions of alveolar bone are sometimes missing due to taphonomic processes, and these can mimic fenestrations that were present premortem. Therefore, the number of fenestrations could have been underestimated. Although these possible complications are acknowledged, this analysis of the sample indicates that dehiscences were more common than fenestrations. Prevalence of Dehiscences and Fenestrations by Age The prevalence of alveolar bone defects in the Windover sample was further analyzed in relation to age. This study found a statistically significant association between the presence or absence of dehiscences and age group using a G Test of Independence (G=15.016, df=5, P=0.010). The number of dehiscences found in each age group is shown in Figure 18. In the youngest age group, 10-19 years, only one individual out of fourteen had a dehiscence. This result suggests that dehiscences were uncommon in young individuals in the Windover sample.

Dehiscences are more prevalent in the 20-29 year age group, with eight out of thirteen individuals (61.5%) exhibiting at least one dehiscence. The 30-39 year age group has the largest percentage of individuals exhibiting dehiscences, with six out of nine individuals (66.7%) with at least one dehiscence. Dehiscences become less common by 40-49 years of age, with only six out of twenty-four individuals (25.0%) exhibiting dehiscences. Between 50-59 years, dehiscences become more prevalent again, with four out of nine individuals (44.4%) showing this type of bone defect. Three out of the five individuals (60.0%) over the age of 60 years had dehiscences. The results for individuals over the age of 60 years may not be reflective of the overall trends in the population, however, because of the small sample size. When the data for the different age groups are compared, the results indicate that most subadults at Windover lacked dehiscences, and the prevalence of dehiscences increased between 20-39 years of age. The percentage of individuals with dehiscences peaked in the 30-39 year age group. In individuals over the age of 40 years, the percentage of people experiencing dehiscences decreased. Thus, the prevalence of individuals with dehiscences decreases with age in the Windover sample.

Figure 18: Prevalence of dehiscences by age group. Blue indicates the absence of dehiscences and red indicates the presence of at least one dehiscence.

The Windover results are similar to patterns of dehiscence prevalence found in studies conducted on modern populations. Other studies have found a decrease with age in the prevalence of teeth that exhibit a dehiscence or fenestration (Davies et al. 1974; Rupprecht et al. 2001). These studies have attributed this result to increased tooth loss in older individuals, which leads to fewer teeth that could have dehiscences. In the Windover population, antemortem loss of the maxillary molars was uncommon. Therefore, the decrease in dehiscence prevalence with age might instead relate to the loss of alveolar bone associated with periodontal disease. Extensive alveolar resorption would eventually lead to loss of the bone surrounding the root surface, which would eliminate the possibility of dehiscences in individuals with more advanced periodontal disease. Since periodontal disease increased with age in the Windover population, dehiscences might be more difficult to observe in older individuals with periodontal disease. As a result, the prevalence of dehiscences observed in the Windover sample decreased in individuals older than 39 years of age. In contrast to dehiscences, fenestrations were not found to have a statistically significant association with age using a G Test of Independence (G=3.742, df=5, P=0.587). Figure 19 reflects the lack of apparent patterning in the prevalence of fenestrations by age group. Only one out of fourteen individuals between 10-19 years of age exhibited a fenestration, which is similar to the results discussed above for dehiscences. Alveolar bone defects appear to be uncommon in very young individuals in the Windover population. In the 20-29 year age group, three out of thirteen individuals (23.1%) exhibited at least one fenestration. This result indicates that fenestrations are far less common than dehiscences in individuals between 20-29 years of age. The prevalence of fenestrations did not increase by 30-39 years of age, with only two out of nine individuals (22.2%) exhibiting fenestrations. Four out of 24 individuals (16.7%) in the 40-49 year age group had fenestrations, indicating that fenestrations were even less common in this age group. Individuals between 50-59 years of age were relatively unaffected by fenestrations, with only one out of nine individuals (11.1%) exhibiting a fenestration. Individuals over the age of 60 years had the highest prevalence of fenestrations, with three out of eight individuals (37.5%) affected by at least one fenestration. These results show a sudden increase in the percentage of individuals affected by fenestrations in the oldest age group. This larger percentage may, however, reflect the small sample size available for this age group. Overall, the statistical

46 insignificance and lack of patterning suggest that the presence of fenestrations was unassociated with the age of the individual.

Figure 19: Prevalence of fenestrations by age group. Blue indicates the absence of fenestrations and red indicates the presence of at least one fenestration.

Prevalence of Dehiscences and Fenestrations by Sex The presence of alveolar bone defects does not appear to be associated with sex in the Windover sample. The prevalence of dehiscences for each sex is shown in Figure 20. Fourteen out of twenty-seven males (51.9%) exhibited at least one alveolar dehiscence, indicating that dehiscences were relatively common in males. In addition, one out of four (25.0%) probable males had a dehiscence. A smaller percentage of females had a dehiscence in at least one location. Only ten out of thirty-two females (31.3%) exhibited a dehiscence, suggesting that dehiscences may have been less common among females. Two out of three (66.7%) probable females also had a dehiscence. Although these results suggest that there may be a difference in the percentage of males and females affected by dehiscences, the results were not found to be 47 statistically significant using a G Test of Independence (G=3.841, df=3, P=0.279). Since the Windover sample does not show a significant association between the presence of dehiscences and sex, age may be a stronger predictor for dehiscences in the Windover population.

Figure 20: Prevalence of dehiscences by sex. Blue indicates the absence of dehiscences and red indicates the presence of at least one dehiscence.

Similarly, fenestrations were not found to be significantly associated with sex using a G Test of Independence (G=2.187, df=3, P=0.535). Figure 21 shows the results for fenestrations when grouped by sex. The graph appears to lack any patterning of fenestrations in relation to the sex of the individual. Only six out of twenty-seven males (22.2%) exhibited at least one fenestration. None of the four probable males had any fenestrations. Thus, fenestrations were probably uncommon in males in the Windover population. The results for females were similar to the percentage of males exhibiting fenestrations. Six out of thirty-two females (18.8%) had at least one fenestration. One out of three (33.3%) probable females had a fenestration. These

48 results indicate that fenestrations were also relatively uncommon in females in the Windover population.

Figure 21: Prevalence of fenestrations by sex. Blue indicates the absence of fenestrations and red indicates the presence of at least one fenestration.

The lack of association between sex and alveolar bone defects is similar to the results reported by Rupprecht et al. (2001) for modern populations. The authors of this study found that both dehiscence and fenestration defects were more common in females than males. However, only the results for fenestrations were found to be statistically significant. Therefore, sex was not a predictor of the number of dehiscences observed in their sample. The number of fenestrations may, however, have been associated with sex. Examination of the results of the Rupprecht et al. (2001) study and the Windover sample suggests that the association between sex and alveolar bone defects may be complex. Factors other than sex, such as age, may play a more important role in predicting the presence of dehiscences and fenestrations. 49

Locations of Dehiscences and Fenestrations Examination of the tooth surfaces most affected by dehiscences and fenestrations can be compared with data regarding the locations of pathological alveolar resorption to obtain a better understanding of alveolar bone loss in the Windover sample. The percentage of teeth with dehiscences and fenestrations, presented separately for each tooth surface, is displayed in Table 6. These results indicate that the largest number of dehiscences (43) occurred on the mesiobuccal and distobuccal surfaces of the right and left first molars. Only four lingual dehiscences were observed, all of which were located on either the left or right first molar. Five dehiscences were located on the buccal surface of the second molars, and the third molars lacked any evidence of dehiscences. Based on these results, the first molar was affected the most by dehiscences, with 47 of the 52 observed dehiscences located on the mesiobuccal, distobuccal, or lingual surface of the first molar. The first molar also exhibited the largest number of fenestrations. Of the 16 fenestrations observed, 13 were located on the first molar. All 13 fenestrations affecting the first molar were located on the mesiobuccal or distobuccal surfaces. Only one lingual fenestration was observed in the Windover sample, and this fenestration was found on the lingual surface of the maxillary second molar. Neither of the maxillary third molars exhibited fenestrations. The pattern of fenestrations was similar to the pattern of dehiscences, with the mesiobuccal and distobuccal surfaces of the first molar affected the most, and the lingual surface rarely affected by bone defects. The results from Windover are similar to a study by Davies et al. (1974), which found that fenestrations were most common on the maxillary first molars, and that these teeth were also one of the most common locations of dehiscences. When the alveolar bone defect results are compared with the locations of pathological alveolar resorption discussed earlier in this chapter, a pattern of alveolar bone loss on the first molar emerges. The mean CEJ-AC measurement for teeth without dehiscences was greatest on the lingual surface of the first molars, with the mesiobuccal and distobuccal surfaces of the first molars also exhibiting alveolar bone loss. Dehiscences and fenestrations both occurred most often on the mesiobuccal and distobuccal surfaces of the first molars. The second and third molars were less common locations of pathological alveolar resorption. The second molars were also a less common location for dehiscences and fenestrations, and the third molars lacked evidence of either type of periodontal defect. These results indicate that the same teeth (the first

50 molars) are most affected by alveolar resorption, suggesting that there could be an association between periodontal disease and dehiscences and fenestrations in the Windover sample. This association is tested statistically in the next section.

Table 6: Distribution of alveolar defects by tooth surface. The tooth surface labels are as follows: R = right, L = left, M = molar, and the number 1, 2, or 3 indicates the molar number. Tooth Surface N Teeth with % Teeth with N Teeth with % Teeth with Dehiscence/ Dehiscence Fenestration/ Fenestration N Teeth Observed N Teeth Observed LM1 mesiobuccal 15/55 27.3 2/58 3.4 LM1 distobuccal 11/57 19.3 7/58 12.1 LM1 lingual 1/66 1.5 0/66 0 LM2 buccal 2/58 3.4 1/58 1.7 LM2 lingual 0/63 0 1/63 1.6 LM3 buccal 0/39 0 0/39 0 LM3 lingual 0/42 0 0/43 0 RM1 mesiobuccal 8/52 15.4 1/53 1.9 RM1 distobuccal 9/54 16.7 3/54 5.6 RM1 lingual 3/65 4.6 0/65 0 RM2 buccal 3/59 5.1 1/61 6.1 RM2 lingual 0/65 0 0/65 0 RM3 buccal 0/42 0 0/42 0 RM3 lingual 0/41 0 0/41 0

Relationship between Dehiscences and Fenestrations and Periodontal Disease The possible association between dehiscences and periodontal disease was tested statistically using a G Test of Independence. Analysis of the results indicates that there is a significant association between dehiscences and periodontal disease (G=13.685, df=3, P=0.003). In several other studies, more alveolar bone defects (dehiscences and fenestrations) were found in skulls with thin alveolar bone (Abdelmalek and Bissada 1973; Edel 1981; Rupprecht et al. 2001). Individuals with thin alveolar bone could be more susceptible to alveolar resorption and, ultimately, periodontal disease. Since the etiologic factors involved in the development of dehiscences are not well understood, the statistical results provide important evidence to support the idea that periodontal disease could be one of the factors contributing to the presence of

51 dehiscences (Rupprecht et al. 2001). Alternatively, dehiscences could be a risk factor for developing periodontal disease. These test results only suggest that dehiscences and periodontal disease are associated, without indicating the nature of this relationship. The results of a G Test of Independence for fenestrations and periodontal disease indicate that these two conditions are not significantly associated (G=5.457, df=3; P=0.141). Fenestrations are typically very small alveolar defects, which expose only a minimal portion of the root surface. Therefore, fenestrations represent a much smaller amount of alveolar resorption than dehiscences. The development of tiny fenestrations in the alveolar bone may be unrelated to the development of much larger areas of alveolar resorption due to periodontal disease. Alternatively, the lack of association between these two conditions may result from the small number of fenestrations (n=14) found in the sample. If the sample size had been larger, the results might have been different. Based on the results from this sample, however, fenestrations and periodontal disease were probably unassociated in the Windover population. Relationship between Periodontal Disease and Other Dental Health Problems Since periodontal disease is often associated with poor oral hygiene, the relationship between periodontitis and other dental health problems was also investigated. Individuals with periodontal disease might also have suffered from other dental problems, such as caries and abscesses. The data for caries and abscesses in the Windover sample, which included the total number of caries and abscesses per individual, were obtained from Wentz (2006). These data were converted into information about the presence or absence of caries and abscesses for each individual. The resulting data were then compared to periodontal disease data obtained from the current study. Dental caries is a bacterial disease that affects the calcified tissues of the teeth, destroying enamel, dentine and cement (Hillson 1996; Soames and Southam 2005). The destruction of these tissues leads to the development of a cavity in the crown or the root surface. Similar to periodontal disease, the primary etiological agent in the development of dental caries is the bacteria in dental plaque. These bacteria ferment dietary carbohydrates, initiating the formation of acid that causes progressive decalcification of the tooth and disintegration of the organic substance of the tooth (Soames and Southam 2005). Since dental caries and periodontal disease both result from the bacteria in dental plaque, the presence of both of these diseases in the same individual might be expected. In the Windover sample, however, these two diseases were not

52 found to be significantly associated. The results of a G Test of Independence indicated that the presence or absence of dental caries was independent of periodontal disease (G=2.198, df=3, P=0.532). The lack of association between dental caries and periodontal disease in the Windover sample indicates that the etiology of these two diseases is probably multifactorial. While the bacteria in plaque play a primary role in the initiation of these two diseases, factors such as diet and heredity may also impact the development and progression of dental caries and periodontal disease. Severe dental caries, attrition, or trauma can cause the pulp of the tooth to become necrotic, leading to an abscess (Hildebolt and Molnar 1991; Soames and Southam 2005). As discussed in the methods chapter, abscesses are formed by the accumulation of pus during pulp death. A hole often develops in the alveolar bone near the root apex to relieve the pressure. Therefore, both abscesses and periodontal disease can cause extensive alveolar bone loss. In the Windover sample, a significant association was found between the presence or absence of abscesses and periodontal disease using a G Test of Independence (G=16.906, df=3, P=0.001). These results indicate that individuals with periodontal disease often have abscesses. This association might be explained by the tendency for individuals with poor oral hygiene to develop these two conditions. Both of these health problems would lead to alveolar bone loss and eventual loss of teeth. The presence of both periodontal disease and abscesses in the same individuals in the Windover sample indicates that individuals in this population often suffered from multiple dental health problems. Periodontal Disease in Different Lineages at Windover The pattern of periodontal disease in different biological lineages at Windover was also examined. The comparison of periodontal disease between lineages will indicate whether the etiology of periodontal disease at Windover was influenced by hereditary factors. Studies of modern populations have found that genetics may play a role in periodontitis (Corey et al. 1993; Hart and Kornman 1997; Kinane et al. 2005). While bacteria are the primary etiological agent responsible for the onset and progression of periodontal disease, genetics may influence susceptibility to the disease (Corey et al. 1993). The severity of periodontitis may also be affected by hereditary factors. The levels of oral microbial flora explain less than 20% of the variability in the expression of periodontal disease (Hart and Kornman 1997), suggesting that other factors such as heredity might be important for explaining differences in the progression of

53 periodontitis between individuals. Studies have estimated that as much as 50% of the variability in attachment loss may be due to hereditary factors (Schenkein 2002). Therefore, comparison of periodontal disease in different lineages at Windover can provide insight into the relative importance of genetics as a risk factor for periodontitis in the population. Stojanowski and Schillaci (2006) have argued that two genetically distinct lineages might be present at Windover. These researchers examined several craniometric, dental morphology, cranial non-metric, odontometric, and malocclusion variables, as well as dental anomalies, in the Windover population. The analysis of these variables using cluster analysis of Gower similarity coefficients led to the suggestion that two genetically different subgroups were buried in Pond C at Windover, with individuals from each each subgroup primarily buried on a particular side of the pond (west or east). The results of this analysis group individuals into two possible lineages, which will be used to evaluate potential genetic factors involved in periodontal disease patterns. The two lineages exhibited some variability in the presence and degree of periodontal disease (Figure 22). The Pond C west lineage included a total of seventeen individuals that could be assessed for periodontal disease, and the east lineage included eleven individuals. In the Pond C west lineage, six out of the seventeen individuals (35.3%) did not have periodontal disease. The remaining eleven individuals (64.7%) exhibited some degree of periodontal disease. Five of these individuals (29.4% of the entire lineage) had probable periodontal disease, and another five (29.4%) had slight to moderate periodontal disease. Only one individual (5.9%) in the west lineage exhibited moderate to severe periodontal disease. Since genetic factors are thought to influence the progression and severity of the disease, this result suggests that individuals in the west lineage may not have been particularly susceptible to severe periodontal disease. These individuals were, however, susceptible to slight to moderate alveolar resorption from periodontal disease. The Pond C east lineage showed a different pattern, with a greater percentage of individuals experiencing moderate to severe periodontal disease. Only three of the eleven individuals (27.3%) in this lineage did not show any symptoms of periodontal disease. The other eight individuals (72.7%) exhibited varying degrees of periodontal disease. One individual (9.1% of the entire lineage) had probable periodontal disease. Four individuals (36.4%) had slight to moderate periodontal disease. Three out of the eleven individuals (27.3%) suffered from moderate to severe periodontal disease. This result indicates that the number of individuals

Figure 22: Prevalence of periodontal disease by lineage. The different bar colors indicate the level of severity of periodontal disease.

with extensive alveolar bone loss was the same as the number of individuals without periodontal disease in the east lineage. The percentage of individuals experiencing moderate to severe periodontal disease in the east lineage (27.3%) was also much higher than the west lineage (5.9%). Therefore, individuals in the east lineage appear to have been highly susceptible to periodontal disease, including more severe forms than individuals in the west lineage. These results suggest that genetics may have been one of the factors that influenced the severity of periodontal disease in the Windover population. Although initial examination of the results from the Windover sample suggests that genetic factors may have impacted periodontal disease, the possible association between lineage and periodontal disease was not found to be statistically significant using a G Test of Independence (G=3.793, df=3, P=0.285). The statistical insignificance may result from the small sample size (n=28) for both lineages. Although Stojanowski and Schillaci (2006) identified forty-nine individuals who belonged to these lineages, only twenty-eight of them had

55 intact maxillae that could be assessed for periodontal disease. A larger sample size might have shown a statistically significant association between lineage and periodontal disease. Even though the results of this study were not statistically significant, the observed variability in the severity of periodontal disease in different lineages at Windover suggests that future bioarchaeological studies should examine the contribution of genetic factors to the development and progression of periodontitis.

CHAPTER 4 – DISCUSSION AND CONCLUSIONS

The results of this study indicate that periodontal disease was a common dental health problem in the Windover population. Approximately 55% of the fifty-six individuals in the Windover sample exhibited some degree of periodontal disease, but only a small number of these individuals experienced severe periodontal disease. These results provide insight into the prevalence of dental disease in an early hunter-gatherer population from Florida. When this percentage is compared with other hunter-gatherer populations from Florida, the prevalence of periodontal disease is found to vary between populations. Therefore, factors other than diet may influence the development and progression of periodontal disease. This concluding chapter first compares the overall percentages of periodontal disease from Windover with other hunter- gatherer and modern populations. The chapter then suggests factors other than diet that may account for the variability in periodontal disease percentages between populations, including dental calculus, age, sex, alveolar bone defects, susceptibility to dental health problems, and heredity. Possible explanations for variability in periodontal disease results based on research methodology and interobserver error are also examined. The thesis concludes with a discussion of the complex etiology and epidemiology of periodontal disease and suggestions for future research. The percentage of the Windover population that experienced periodontal disease may have been considerably lower than the hunter-gatherer population at Highland Beach (Table 7). Isler et al. (1985) examined a sample of 108 adult individuals (defined as individuals with erupted permanent second molars) from Highland Beach. This society was thought to practice a subsistence strategy involving the hunting of small animals, use of marine resources, and the gathering of plants. This strategy was probably similar to the subsistence activities carried out by the Windover population. Despite similar subsistence practices, the percentage of periodontal disease in the two populations may have differed greatly. Isler, Schoen, and Iscan report that 92% of the alveoli from Highland Beach showed some degree of resorption (1985:142). Since this result is based on the total number of teeth rather than the total number of individuals, the actual number of individuals with periodontal disease is probably slightly lower. However, this result still implies that the prevalence of periodontal disease was greater in the Highland Beach population than at Windover. As mentioned above, approximately 55% of the individuals from

Windover exhibited some amount of alveolar resorption. The discrepancy in these results suggests that diet may be only one of many etiological factors involved in the development of periodontal disease. The population from the Fort Center site also exhibited a higher percentage of alveolar resorption than the Windover population. Based on faunal and floral remains recovered from the Fort Center site, the population probably practiced a hunting and gathering subsistence strategy, consuming reptiles, fish, mammals, and plants (Shaivitz 1986). Thus, the subsistence strategy was relatively similar to the strategy practiced at Windover. In contrast to Windover, however, examination of the skeletal remains from Fort Center led to the conclusion that 94.7% of the 630 alveoli studied showed some amount of alveolar resorption (Shaivitz 1986:105). Similar to Highland Beach, this study examined the total number of teeth rather than the total number of individuals exhibiting alveolar resorption, and the percentage of individuals with periodontal disease may therefore have been lower. This result is still much higher than the Windover population, however, and more closely resembles the Highland Beach population. The results from these three sites suggest that the prevalence of periodontal disease varies greatly between hunter-gatherer populations. The Republic Groves population may have exhibited the highest periodontal disease prevalence of any Florida hunter-gatherer group. After examination of the human skeletal remains from the site, Saunders (1972) concluded that every adult in the sample showed some evidence of periodontal disease. While Saunders did not carry out a systematic study of alveolar recession, her conclusion indicates that as much as 100% of the adult population at Republic Groves might have suffered from some degree of periodontal disease.

Table 7: Periodontal disease prevalence in Florida hunter-gatherer populations. Site n Percentage of Adult Sample Reference with Periodontal Disease Windover 56 individuals 55.4% Highland Beach 108 individuals 92% Isler et al. (1985) Fort Center 630 teeth 94.7% Shaivitz (1986) Republic Groves Unspecified Up to 100% Saunders (1972) Manasota Key Cemetery 41 individuals 58% Dickel (1991)

The epidemiology of periodontal disease in hunter-gatherer populations from Florida is further complicated by the comparison of Windover, Highland Beach, Fort Center, and Republic Groves with the Manasota Key Cemetery. This site is located on the west coast of Florida and dates to 1730 years B.P. (Dickel 1991:22). Although later than Windover, the people at Manasota Key are thought to have consumed a similar diet focused on marine resources, with additional hunting of small game and gathering of plant foods. Dickel (1991) recorded the presence of periodontal resorption for 41 adult individuals, concluding that 58% of the total sample exhibited signs of periodontal disease. This percentage is similar to the result of the current study, which found that 55% of the Windover sample showed symptoms of periodontal disease. These results suggest that diet may be one factor in the development of periodontal disease. Comparison with the results from Highland Beach, Fort Center, and Republic Groves, however, suggests that diet is not the only factor affecting periodontal disease. The variability in periodontal disease prevalence between these five sites has implications for the epidemiological study of past populations. Archaeologists often discuss changes in the prevalence of health problems such as dental caries rates as a result of the shift from hunting and gathering to agricultural subsistence (Larsen 1983; Larsen et al. 1991; Temple and Larsen 2007; Turner 1979). While the discussion of all hunter-gatherer populations as a group may be useful for caries rates, periodontal disease prevalence varies greatly between different hunter-gatherer populations. Therefore, the range of periodontal disease prevalence in hunter-gatherer groups would likely overlap with the range for agriculturalists. The observed differences between hunter-gatherer groups in the prevalence of periodontal disease may occur for one of several reasons. First, this variability may indicate that these hunter-gatherer groups had substantially different diets. Second, individuals within each population may have consumed different foods, based on such factors as sex. Wentz (2006) suggests that males and females at Windover could have had unequal access to food, leading to health differences between the sexes. The possibility of inter- and intra-group differences in diet might account for some of the inconsistency in periodontal disease results between hunter- gatherer populations. However, differences in diet probably only account for a small amount of the variability between hunter-gatherer populations. The following comparison of periodontal

59 disease between the Windover population and modern populations provides evidence that diet may not be a strong predictor of periodontal disease. The results from Windover may be similar to epidemiological studies conducted by the World Health Organization on modern populations (Rudko 1973). These studies determined that 75% of adults older than 18 years of age have periodontal disease. Based on this result, the percentage of the population suffering from periodontal disease at Windover may have been as much as 20% lower than modern rates. Although the percentage of individuals experiencing periodontal disease at Windover has been estimated at 55%, this result may not be directly comparable to the World Health Organization study for two reasons. First, the actual percentage of individuals experiencing some stage of periodontal disease at Windover may have been higher because the disease often starts as gingivitis, which is unobservable in archaeological maxillae. Second, the Windover sample includes individuals younger than 18 years of age. When individuals under the age of 18 years are excluded from the sample, 70.5% of the remaining forty-four individuals from Windover exhibit symptoms of periodontal disease. The similarity of this percentage to the results found by the World Health Organization suggests that changes in diet may not have greatly impacted the percentage of the overall population affected by periodontal disease. This result provides support for the possibility that periodontal disease has a multifactorial etiology. Study of the Windover sample revealed many factors that may influence the development and progression of periodontal disease in all populations, regardless of diet or time period. Any study that examines the etiology and epidemiology of periodontal disease should consider the variables described below. Dental calculus may have been one of the most important etiological agents involved in the development of periodontal disease at Windover. This study found a highly significant association between the percentage of tooth surfaces affected by dental calculus and the severity of periodontal disease in the Windover sample. These results suggest that individuals with extensive dental calculus were more likely to develop periodontal disease. The bacteria in dental plaque, which mineralizes to form dental calculus, have been identified as one of the primary etiological agents that facilitate the onset of periodontal disease in modern populations (Soames and Southam 2005). The accumulation of dental plaque causes inflammation of the nearby gingival tissues, which can lead to chronic periodontitis and eventual loss of alveolar bone (Albandar 2002). In modern populations, the removal of dental plaque effectively stops the

60 inflammation (Gaare et al. 1990). Therefore, the assumption that extensive dental calculus was one of the primary causes of periodontal disease at Windover is valid. Individuals with dental calculus had a large surface that would readily support the multiplication of the bacteria that initiated periodontal disease. Individuals with only small amounts of dental calculus would be less likely to develop large colonies of these bacteria on their teeth. These results indicate that dental calculus is a major risk factor for periodontal disease, both at Windover and in modern populations. Thus, differences in the amount of dental calculus may account for part of the observed variability in the prevalence of periodontal disease between hunter-gatherer populations. The study of periodontal disease in past populations should therefore include an examination of the amount of dental calculus in each population. Another factor that must be assessed in bioarchaeological studies of periodontal disease is age. In the Windover sample, the association between age and the presence and severity of periodontal disease is highly significant. This increase in the prevalence of periodontal disease with age has been demonstrated in other archaeological and modern populations (Albandar 2002; Goldman and Cohen 1973; Molnar and Molnar 1985; Schluger et al. 1977). Modern epidemiological studies have found that severe periodontal disease occurs more often in older age groups (Locker et al. 1998), a conclusion which is supported by the data from Windover. Although the association between increased age and periodontal disease has been established, the precise nature of this relationship is poorly understood. Older individuals might demonstrate more bone loss than younger individuals because periodontal disease is a progressive disease, with the loss of alveolar bone occuring during periodic intervals (Locker et al. 1998). Individuals may initially develop periodontal disease at a younger age, but the bone loss may not become severe until the individual reaches old age. Therefore, the presence of periodontal disease in older individuals does not mean that the age of onset occurs at an older age. Additionally, increasing age should not be regarded as a cause of periodontal disease. One of the four individuals over 60 years of age at Windover did not exhibit any symptoms of periodontal disease. These results, in combination with modern studies (Burt 1994), suggest that periodontal disease is not a necessary consequence of aging. Increasing age is, however, considered a potential risk factor for the development of periodontal disease (Locker et al. 1998). Part of the variability in the prevalence of periodontal disease in hunter-gatherer populations might be explained by differences in the age distributions of each sample. Since the observed prevalence

61 of periodontal disease might be affected by the age distribution of the sample, bioarchaeologists studying periodontal disease should determine the relative proportions of each age group in the sample. An additional factor that might influence variability in periodontal disease prevalence between populations is sex. This study found that periodontal disease may have been more prevalent among males than females at Windover, and periodontal disease was often more severe in males. This possible association between periodontal disease and sex was not statistically significant, however. These results suggest that the presence and severity of periodontal disease may not have been related to sex in the Windover sample. Despite these results, the sex of individuals with periodontal disease should still be investigated in other populations because archaeological and modern clinical studies have found an association between these two variables. In the Highland Beach sample, males exhibited severe alveolar resorption more often than females (Isler et al. 1985). Studies of modern populations have also found that periodontal disease occurs more often in males than females (Goldman and Cohen 1973; Hildebolt and Molnar 1991). These results suggest that periodontal disease may be more common in males than females. Some of the variability between hunter-gatherer populations analyzed in this study might be attributed to differences in the relative representation of males and females in the samples. Therefore, the study of periodontal disease in past populations should include a description of the sex distribution of the sample. Alveolar bone defects are another factor that may be related to periodontal disease in the Windover population. The category of alveolar bone defects includes both dehiscences, which expose the entire root surface on one side, and fenestrations, which expose a small section of the root (Rupprecht et al. 2001). In the Windover sample, only dehiscences were significantly associated with periodontal disease. Despite the lack of a significant association between fenestrations and periodontal disease, fenestrations should still be examined in studies of periodontal disease because fenestrations may be a transitional stage in the development of dehiscences (Elliot and Bowers 1963). In contrast to dental calculus, dehiscences and fenestrations are not causes of periodontal disease. However, individuals with thinner bone often exhibit more alveolar bone defects (Abdelmalek and Bissada 1973; Edel 1981; Rupprecht et al. 2001). Thin bone might make individuals more susceptible to alveolar resorption, which is one of the characteristics of periodontal disease. Alternatively, periodontal disease might increase

62 the likelihood of developing dehiscences because the disease leads to alveolar bone loss (Rupprecht et al. 2001). The exact nature of the association between periodontal disease and dehiscences is not well understood. Dehiscences could be a risk factor for periodontal disease, or a consequence of the disease. Although the relationship between these two variables is uncertain, these two variables were clearly associated with each other in the Windover sample. Therefore, examination of alveolar bone defects in paleoepidemiological studies of periodontal disease may provide useful information about the dental characteristics of individuals with periodontal disease. Bioarchaeological studies of periodontal disease might also be enhanced by investigating other dental health problems that occur in individuals with periodontal disease. In the Windover sample, individuals with periodontal disease were likely to exhibit abscesses. This association indicates that individuals who were susceptible to alveolar resorption from periodontal disease were also susceptible to loss of bone due to abscesses. The study did not, however, find a significant association between periodontal disease and dental caries in the Windover sample. These results suggest that, although dental caries and periodontal disease are both initiated by the bacteria in dental plaque (Soames and Southam 2005), these two oral health problems do not necessarily develop in the same individual. The etiologies of dental caries and periodontal disease involve a complex combination of factors, and plaque alone does not explain the prevalence of both diseases. These results indicate that periodontal disease is not associated with all other dental pathologies, but may be associated with susceptibility to abscesses. The examination of both periodontal disease and other dental problems such as abscesses offers a more complete understanding of the overall dental health in an archaeological population. The final variable that should be addressed in studies of periodontal disease in past populations is heredity. Stojanowski and Schillaci (2006) hypothesized that two genetically distinct lineages might be present at Windover, based on the analysis of several skeletal and dental variables. When periodontal disease prevalence and severity was compared in these two possible lineages, individuals from the lineage that was primarily buried on the east side of the pond seemed to have more severe periodontal disease. The potential association between the severity of periodontal disease and lineage was, however, statistically insignificant. The sample size for the lineage analysis was only twenty-eight individuals, and a larger sample size might have produced significant results. The differences in the percentage of individuals affected by

63 severe periodontal disease in the two lineages at Windover suggests that heredity studies of periodontal disease in other archaeological populations might find an association between these variables. To date, other bioarchaeological studies of periodontal disease have not examined heredity as a potential explanation for the distribution of the disease in the population. This might be due to the difficulty of assessing genetic relationships in archaeological populations. Modern clinical studies, however, provide extensive evidence for the important role of genetic factors in the development and progression of periodontal disease (Hart and Kornman 1997; Kinane et al. 2005). Genetics may influence both initial susceptibility to periodontal disease and the severity of the disease (Corey et al. 1993). Recent studies suggest that as much as half of the variability in the loss of attachment due to periodontal disease may be related to heredity (Schenkein 2002). Therefore, a large percentage of the variability in the prevalence of periodontal disease between Florida hunter-gatherer populations might be explained by genetic factors. Future paleoepidemiological studies should address the possible contribution of heredity to the prevalence and severity of periodontal disease in archaeological populations. While any combination of these etiological and demographic factors might explain some of the variability in the prevalence of periodontal disease between hunter-gatherer populations, differences in research methodology might also account for part of the observed variability. As discussed in Chapter 1, researchers often disagree about the appropriate methods for identifying periodontal disease in archaeological populations (Hildebolt and Molnar 1991). The lack of standardized methods for the assessment of periodontal disease has led to the use of different methods in many studies, often preventing comparison of the results of multiple studies. Part of the variability between the hunter-gatherer groups discussed in this study might therefore result from methodological discrepancies. The Highland Beach and Fort Center studies relied entirely on quantitative measurements of alveolar resorption for the analysis of periodontal disease (Isler et al. 1985; Shaivitz 1986). The studies of periodontal disease at Republic Groves and Manasota Key Cemetery relied on subjective assessment of the alveolar bone, but these studies did not provide a detailed description of the methods used to make this assessment (Dickel 1991; Saunders 1972). Therefore, these results may not be directly comparable with the results of studies of periodontal disease in other populations. The current study of the Windover sample used a combination of quantitative measurements of alveolar bone loss and subjective assessment of the alveolar bone condition. The observed variability in periodontal disease

64 prevalence between these hunter-gatherers populations might be partially explained by discrepancies in the methods used by different researchers. Another possible explanation for the variability in the results of these studies is interobserver error. The assessment of periodontal disease in skeletal remains is ultimately subjective, and different observers might reach dissimilar conclusions about the periodontal health of the same individual (Clarke 1990). Although interobserver error cannot be eliminated, a standardized methodology for the assessment of periodontal disease might reduce the degree of error. The current study provides a methodology for the comprehensive assessment of periodontal disease, including analysis of etiological and demographic factors that may influence the prevalence of periodontal disease in a particular population. Future studies might be able to use this methodology, which could be improved by the addition of a method to reduce the subjectivity of the final assessment of periodontal disease. This study of periodontal disease in the Windover population reveals the complex etiology, epidemiology, and methodology involved in the assessment of this dental problem. The prevalence of periodontal disease varies greatly between early Florida hunter-gatherer populations, suggesting that bioarchaeologists should not assume uniformity in the disease profiles of these groups simply because of a similar subsistence strategy and geographical location. Part of the variability in these results might be explained by differences in methodology between studies. Etiological and demographic factors typically associated with periodontal disease might also account for part of the variability between hunter-gatherer populations. Bioarchaeological studies of periodontal disease should address these factors, which include dental calculus, age, sex, alveolar bone defects, susceptibility to dental health problems, and heredity. Examination of the many etiological and demographic factors associated with periodontal disease will provide insight into the possible reasons for variability in dental health between archaeological populations.

APPENDIX

Table 8: Windover periodontal disease and dental calculus results by individual. Individual Approximate Sex Periodontal Disease % Tooth Surfaces Age with Calculus 36.293 32 Probable Female Slight to moderate None 36.432 35 Female Slight to moderate ¾ to all 36.504 45 Female Slight to moderate ¾ to all 52.3 25 Probable Male Slight to moderate ¼ to less than ½ 53.68 41 Female Slight to moderate ¼ to less than ½ 57.077 38 Male Moderate to severe ½ to less than ¾ 57.300 25 Male None None 68.029 41 Male ¼ to less than ½ 69.084 58 Male Probable ½ to less than ¾ 69.1 19 Female None ¼ to less than ½ 71.4 49 Female Less than ¼ 72.41 48 Female ¼ to less than ½ 74.42 44 Female Probable Less than ¼ 75.3 10 Subadult None ¼ to less than ½ 76.11 22 Female Slight to moderate None 78.35 69 Female ½ to less than ¾ 79.18 15 Subadult None ¼ to less than ½ 81.11 23 Female None 82.3 38 Female Moderate to severe ½ to less than ¾ 83.66 22 Probable Female Probable Less than ¼ 84.2 12 Subadult None ¼ to less than ½ 86.26 15 Subadult None Less than ¼ 87.10 45 Male Moderate to severe ½ to less than ¾ 91.33 46 Probable Male ½ to less than ¾ 92.9 50 Male Moderate to severe ½ to less than ¾ 93.3 58 Female Probable ½ to less than ¾ 94.54 49 Male Slight to moderate Less than ¼ 95.1 23 Male None ½ to less than ¾ 97.113 30 Male Slight to moderate ¾ to all 99.322 20 Female None ¼ to less than ½ 99.50 18 Female None Less than ¼ 99.84 22 Female None ¼ to less than ½ 101.1 23 Female None ¼ to less than ½ 103.32 37 Female ¾ to all 104.21 56 Probable Female ¾ to all 109.12 65 Female ½ to less than ¾ 109.86 29 Male None ½ to less than ¾ 110.24 46 Probable Male Slight to moderate ½ to less than ¾ 112.38 14 Subadult None ¼ to less than ½

Table 8—continued. Individual Approximate Sex Periodontal Disease % Tooth Surfaces Age with Calculus 114.22 39 Male Less than ¼ 115.2 40 Female Less than ¼ 116.1 11 Subadult None ¼ to less than ½ 117.1 46 Female Probable ¼ to less than ½ 118.2 37 Male Slight to moderate ¼ to less than ½ 119.51 62 Male Probable Less than ¼ 120.1 35 Male ½ to less than ¾ 121.23 53 Female Slight to moderate ¾ to all 122.1 17 Subadult None ½ to less than ¾ 123.5 42 Male Less than ¼ 124.15 66 Female ¾ to all 125 62 Female Probable ¾ to all 127.8 56 Female Less than ¼ 128.1 61 Male ¼ to less than ½ 129.25 17 Subadult None None 130.12 28 Male ¾ to all 131.1 65 Male Moderate to severe ½ to less than ¾ 136.9 46 Female None ½ to less than ¾ 138.24 21 Female None ¼ to less than ½ 142.42 56 Male Probable ½ to less than ¾ 143.40 18 Female None Less than ¼ 145.27 46 Male ½ to less than ¾ 147.1 45 Male ¼ to less than ½ 150.4 13 Subadult None Less than ¼ 151.14 63 Female None ½ to less than ¾ 152.1 51 Male Slight to moderate ¾ to all 153.1 49 Male Less than ¼ 154.11 47 Male Probable ¼ to less than ½ 155.4 40 Male None ¼ to less than ½ 156.4 10 Subadult None ¼ to less than ½ 157.1 46 Female Slight to moderate ¼ to less than ½ 158.31 45 Male Slight to moderate ¾ to all 162.1 49 Probable Male Slight to moderate Less than ¼ 163.1 41 Female None ½ to less than ¾ 246.10 51 Female Probable ¼ to less than ½ 501.1 24 Female None ½ to less than ¾ 501.6 42 Male Slight to moderate ½ to less than ¾

Table 9: Summary of G Test of Independence results for dental conditions. Results significant at P<0.05 are indicated by an asterisk. Variables G Value df P Periodontal Disease and Age 58.194 15 0.000* Periodontal Disease and Sex 14.187 9 0.116 Dental Calculus and Age 32.516 20 0.038* Dental Calculus and Sex 10.148 12 0.603 Periodontal Disease and Dental Calculus 30.448 12 0.002* Dehiscence and Age 15.016 5 0.010* Fenestration and Age 3.742 5 0.587 Dehiscence and Sex 3.841 3 0.279 Fenestration and Sex 2.187 3 0.535 Dehiscence and Periodontal Disease 13.685 3 0.003* Fenestration and Periodontal Disease 5.457 3 0.141 Dental Caries and Periodontal Disease 2.198 3 0.532 Abscesses and Periodontal Disease 16.906 3 0.001* Lineage and Periodontal Disease 3.793 3 0.285

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BIOGRAPHICAL SKETCH Maria Therese Fashing was born on July 13, 1983 in Williamsburg, Virginia. She is the daughter of Norman and Gisela Fashing. Maria received her B.A. from the College of William and Mary in 2005, with an Anthropology major and Music minor. She developed an interest in dental anthropology as an undergraduate, conducting research on dental caries in an Anglo- Saxon population from Sedgeford, England. She also became interested in zooarchaeology, writing an Honors thesis that interpreted variability in diet on eighteenth-century Virginia plantations based on faunal remains. After graduating, she worked for the Colonial Williamsburg Foundation and Jamestown-Yorktown Foundation for a year. In the fall of 2006, she started graduate school at Florida State University. Her research has focused on the epidemiology of dental pathologies in the Windover population.