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Dental Asymmetry through Time in Coastal Florida and Georgia

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Dental Asymmetry through Time in Coastal Florida and Georgia
Creator:
WILLIAMS, SHANNA E.
Copyright Date:
2008

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Subjects / Keywords:
Anthropology ( jstor )
Asymmetry ( jstor )
Dentition ( jstor )
Mandible ( jstor )
Maxilla ( jstor )
Paleoanthropology ( jstor )
Physical anthropology ( jstor )
Sex linked differences ( jstor )
Sexual dimorphism ( jstor )
Teeth ( jstor )
Tick Island ( local )

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University of Florida
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University of Florida
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Copyright Shanna E. Williams. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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4/30/2005
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656215751 ( OCLC )

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DENTAL ASYMMETRY THROUGH TIME IN COASTAL FLORIDA AND GEORGIA By SHANNA E. WILLIAMS A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF ARTS UNIVERSITY OF FLORIDA 2004

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Copyright 2004 by Shanna E. Williams

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ACKNOWLEDGMENTS I would like to acknowledge my committee members, Michael Warren (chair), Anthony Falsetti, and Thomas Hollinger, for making time in their busy schedules to offer me support and guidance. I am also grateful to Scott Mitchell of the Florida Museum of Natural History, whose permission, patience, and assistance in making skeletal material accessible to me made this endeavor possible. A special “thank you” goes to friend and co-conspirator, Alana Lynch, for her critical eye for editing, generous ear for listening, and creative mind for ideas. I sincerely thank my family. Their constant support and guidance kept me moving forward, even when I wanted desperately to stay planted in one spot. I thank them for encouraging me to strive for more, to push the boundaries, and most importantly to never settle. iii

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TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iii LIST OF TABLES............................................................................................................vii LIST OF FIGURES...........................................................................................................ix KEY TO ABBREVIATIONS AND ACRONYMS..........................................................xi ABSTRACT.....................................................................................................................xiii CHAPTER 1 INTRODUCTION........................................................................................................1 Research Criteria..........................................................................................................2 Hypothesis....................................................................................................................4 2 PREVIOUS WORK ON DENTAL ASYMMETRY...................................................6 Background...................................................................................................................6 Previous Literature........................................................................................................7 Genetic Studies......................................................................................................8 Ethnogeographic Studies.......................................................................................9 Sexual Dimorphism in Dental Asymmetry.........................................................10 3 GEOGRAPHIC AND TEMPORAL BOUNDARIES................................................12 Introduction.................................................................................................................12 Natural Environment..................................................................................................12 Cultural Periods..........................................................................................................13 Paleoindian Period...............................................................................................13 Archaic Period.....................................................................................................14 Late Prehistoric Periods.......................................................................................15 European Contact................................................................................................18 iv

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4 SITES..........................................................................................................................21 Introduction.................................................................................................................21 Bay West (8CR200)............................................................................................22 Tick Island (8VO24)............................................................................................24 Palmer (8SO2).....................................................................................................25 Bayshore Homes (8PI41)....................................................................................26 St Simons Island Sites (9GNXXXX)..................................................................27 Harrison Homestead (8NA 41)............................................................................27 Contemporary Sample.........................................................................................29 Temporal Series...................................................................................................29 5 MATERIALS AND METHODS...............................................................................31 Materials.....................................................................................................................31 Occlusal Surface Wear...............................................................................................31 Sexing.........................................................................................................................34 Dental Measurements.................................................................................................34 Statistical Methods......................................................................................................35 6 RESULTS...................................................................................................................37 Fluctuating Asymmetry..............................................................................................41 Sex Differences...................................................................................................41 Site Differences...................................................................................................41 Sex*Site Interaction.............................................................................................44 Directional Asymmetry..............................................................................................46 Sex Differences...................................................................................................46 Site Differences...................................................................................................47 Sex*Site Interaction.............................................................................................48 7 DISCUSSION AND CONCLUSIONS......................................................................49 Summary of Statistical Results...................................................................................49 Sex Differences...................................................................................................50 Site Differences...................................................................................................52 Sex*Site Interaction.............................................................................................57 Fluctuating Asymmetry vs. Directional Asymmetry...........................................58 Asymmetry in Tooth Classes...............................................................................58 Further Research.........................................................................................................61 Conclusions.................................................................................................................61 v

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APPENDIX A MAXILLA VALUES FOR FLUCTUATING ASYMMETRY.................................64 B MANDIBLE VALUES FOR FLUCTUATING ASYMMETRY..............................68 LIST OF REFERENCES...................................................................................................71 BIOGRAPHICAL SKETCH.............................................................................................78 vi

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LIST OF TABLES Table page 3-1 Relevant cultural periods in Florida and Georgia....................................................20 4-1 Site ranges and cultural periods...............................................................................30 6-1 Maxillary values for fluctuating asymmetry............................................................38 6-2 Mandibular values for fluctuating asymmetry.........................................................38 6-3 Maxillary values for directional asymmetry............................................................39 6-4 Mandibular values for directional asymmetry.........................................................39 6-5 Fluctuating asymmetry site differences....................................................................40 6-6 Directional asymmetry site differences....................................................................40 A-1 Female maxilla data.................................................................................................64 A-2 Male maxilla data.....................................................................................................64 A-3 Bay West maxilla data..............................................................................................65 A-4 Tick Island maxilla data...........................................................................................65 A-5 Palmer maxilla data..................................................................................................65 A-6 Bayshore maxilla data..............................................................................................66 A-7 St. Simons maxilla data............................................................................................66 A-8 Harrison maxilla data...............................................................................................66 A-9 Contemporary population maxilla data....................................................................67 B-1 Female mandible data...............................................................................................68 B-2 Male mandible data..................................................................................................68 B-3 Bay West mandible data...........................................................................................68 vii

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B-4 Tick Island mandible data........................................................................................69 B-5 Palmer mandible data...............................................................................................69 B-6 Bayshore mandible data...........................................................................................69 B-7 St. Simons mandible data.........................................................................................70 B-8 Harrison mandible data............................................................................................70 B-9 Contemporary population mandible data.................................................................70 viii

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LIST OF FIGURES Figure page 3-1 Line dividing northern and southern Florida............................................................16 4-1 Location of St. Simons in Georgia...........................................................................21 4-2 County location of Florida samples.........................................................................22 4-3 Map of Bay West site (8Cr200)...............................................................................23 4-4 Tick Island (8VO24) sketch.....................................................................................24 4-5 Palmer (8SO2) sketch...............................................................................................25 4-6 Bayshore Homes (8PI41) sketch..............................................................................26 4-7 Harrison Homestead (8NA41) sketch......................................................................28 5-1 Smith surface wear scoring system..........................................................................32 5-2 Scott surface wear scoring system...........................................................................33 6-1 Mandible sex differences (FA).................................................................................41 6-2 Maxilla site differences (FA) in buccolingual dimension........................................42 6-3 Maxilla site differences (FA) in the mesiodistal dimension....................................43 6-4 Mandible site differences (FA) in the buccolingual dimension...............................43 6-5 Mandible site differences (FA) in the mesiodistal dimension..................................44 6-6 Maxilla sex*site interactions (FA)...........................................................................45 6-7 Mandible sex*site interactions (FA) for PM4..........................................................45 6-8 Mandible sex*site interactions (FA) for C1.............................................................46 6-9 Mandible sex differences (DA)................................................................................46 6-10 Maxilla site differences (DA)...................................................................................47 ix

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6-11 Mandible site differences (DA)................................................................................47 6-12 Maxilla sex*site interactions (DA)..........................................................................48 7-1 Mandible sex*site interactions (DA) for instances of sex difference......................51 7-2 Fluctuating asymmetry bar graphs of site differences.............................................52 7-3 Directional asymmetry bar graphs of site differences..............................................53 7-4 Buccolingual maxillary FA across sites...................................................................54 7-5 Mesiodistal maxillary FA across sites......................................................................54 7-6 Buccolingual mandibular FA across sites................................................................55 7-7 Mesiodistal mandibular FA across sites...................................................................55 x

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KEY TO ABBREVIATIONS AND ACRONYMS FMNH Florida Museum of Natural History NAGPRA Native American Graves Protection and Repatriation Act C1 canine PM3 3rd premolar PM4 4th premolar M1 1st molar M2 2nd molar FA fluctuating asymmetry DA directional asymmetry BL buccolingual MD mesiodistal bC1 buccolingual dimension of canine bPM3 buccolingual dimension of 3rd premolar bPM4 buccolingual dimension of 4th premolar bM1 buccolingual dimension of 1st molar bM2 buccolingual dimension of 2nd molar mC1 mesiodistal dimension of canine mPM3 mesiodistal dimension of 3rd premolar mPM4 mesiodistal dimension of 4th premolar xi

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mM1 mesiodistal dimension of 1st molar mM2 mesiodistal dimension of 2nd molar xii

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Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Arts DENTAL ASYMMETRY THROUGH TIME IN COASTAL FLORIDA AND GEORGIA By Shanna E. Williams May 2004 Chair: Michael Warren Major Department: Anthropology Biological symmetry of paired structures indicates an organism’s ability to maintain developmental stability under a given set of conditions. When this stability is compromised, deviations from symmetry may ensue. By evaluating departures from expected norms, or asymmetry, scientists can examine the causal factors by which these deviations arise. Teeth are the focus of many asymmetry studies. Generally, odontometric studies investigate two forms of dental asymmetry: directional, where crown size is consistently larger on one side, and fluctuating; wherein the largest side varies among individuals of a given population. Dental asymmetry has been primarily attributed to non-specific environmental stressors, including both physical surroundings and cultural exploitation of these surroundings. Studies have revealed a pattern indicating that dental asymmetry is greatest among human populations with the highest incidences of suboptimal levels of health and nutrition. Only a few of these studies have compared skeletal populations; and xiii

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each used research populations that were not only temporally, but geographically disparate. This study, however, is unique in that it examines temporally distinct groups occupying the same landscape. By holding landscape as a constant, this work is able to assess other environmental factors, such as the impact of shifts in cultural practices, on dental asymmetry over time. Mesiodistal and buccolingual diameter measurements were collected from temporally distinct skeletal collections housed at the Florida Museum of Natural History and the Forensic Research Laboratory at the University of Florida. The sample includes six indigenous populations of coastal Florida and Georgia, ranging from the Middle Archaic Period (5000 to 3000 B.C.) to Spanish contact (1683 A.D.); as well as a contemporary population. It was hypothesized that differences would be population-based (i.e., the result of temporal differences), with the highest and lowest asymmetry levels in the Contact Period sites and the contemporary population, respectively. Furthermore, sex differences were believed to be negligible. The results of this study showed that elevated dental asymmetry levels were consistently found in only one of the two Contact Period sites and surprisingly both Woodland Period sites (500 B.C. to 1000 A.D.). Also only a few instances of sexual dimorphism were demonstrated. Overall, these trends suggest that dental asymmetry levels vary through time, and are potentially related to shifts in cultural practices (as opposed to exclusively being the product of geographic variation). Thus, dental asymmetry can serve as a valuable tool in assessing general lifestyle trends related to certain time periods; and assessing patterns of stress unique to particular populations. xiv

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CHAPTER 1 INTRODUCTION Biological symmetry represents a balance between processes that disrupt developmental instructions (developmental noise) and processes that resist this disruption (developmental homeostasis). When balance of these opposing processes is compromised, asymmetry results (Palmer 1994). Developmental homeostasis can be understood as a genotype’s ability to produce a well-formed adaptive phenotype despite developmental disturbances (Livshits and Kobyliansky 1991). Evolution has developed a mechanism that buffers against these disturbances, thereby maintaining homeostasis. This mechanism, known as stabilizing selection, has two forms. The first is normalization, which removes genotypes leading to abnormalities or extreme phenotypes. The second is canalization, which stabilizes the morphological program by buffering an organism against a variety of different environmental conditions (Waddington 1957; Livshits and Kobyliansky 1991; Palmer 1994). Biologically, this is achieved by negative feedback systems that regulate enzyme activity, hormonal regulation of non-contiguous structures, and/or central nervous regulation of non-contiguous structures (Palmer 1994). When this intricate system is compromised, asymmetry results. Asymmetry has been studied in a variety of organisms including Drosophila, mice, rats, fish, and humans. Characters such as paired organs, numbers of scales, wing venation, and skeletal variants are often investigated in these studies (Parsons 1990). 1

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2 Asymmetry studies examining the dentition are particularly prevalent because of the durable nature of teeth. When considering the body as a whole, teeth are by far the most resistant to chemical and physical destruction. This high rate of preservation is directly attributable to an outer layer of highly durable enamel, and a hard, mineralized tissue core of dentine (Johanson and Blake 1996). Since teeth often withstand the taphonomic effects of internment, they can serve as a valuable tool in comparisons between archaeological and modern human populations (Hillson 1996). Generally, odontometric studies consider two types of dental asymmetry: directional, where crown size is consistently larger on one side; and fluctuating, wherein the largest side varies between individuals of a given population. Research suggests that dental asymmetry can serve as an indicator of environmental stress, whereby external stressors such as nutrition, climate, and disease can negatively impact dental development in such a way to result in asymmetry. Impressed by the potential wealth of biological information imbedded within the teeth, I became interested in trying to discern how dental asymmetry can reflect environmental stressors through time. Hence I approached this issue by examining mesiodistal and buccolingual tooth diameters. These measurements were then used to quantify asymmetry within the permanent dentitions of early prehistoric, late prehistoric, contact period, and modern skeletal populations. Research Criteria This experiment consists of data generated from six Amerindian samples and one contemporary population from the Southeastern Coastal Plain, specifically regions of Florida and Georgia. These populations were chosen based on the following criteria:

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3 The need to have a large sample size for statistical analysis Access to the skeletal material Limited genetic variation The first two criteria were easily met. For over 40 years, the Florida Museum of Natural History (FMNH) has conducted archaeological excavations throughout the state. Material gathered from these excavations has been curated by the FMNH in accordance with relevant Federal regulations. Since Native American Graves Protection and Repatriation Act (NAGPRA) regulations do not apply to the remains housed at FMNH, the museum provides a unique opportunity for researchers to study precontact and contact skeletal remains. Similarly, the presence of a Forensic Research Laboratory at the University of Florida provides access to modern material. The third criterion, however, was a little more dubious. Controlling for genetic variability is virtually impossible, because of temporal variation and migration patterns. Furthermore, there is currently no clear technique to discern between modern and pre-20 th Century skeletal populations through statistical methods. However, if one were to consider the matter from a global perspective; any genetic differences, at least in the archaeological collections, would be of minor importance, given the following conditions: All noncontemporary populations were Amerindian They all resided in the same environment (i.e., Southeastern Coastal Plain) The characteristics under examination are known to vary primarily in response to environmental; as opposed to genetic stress (Kieser 1990; Hillson 1996) While the first two conditions are less applicable to the contemporary population, the third condition still holds true.

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4 Coming from the Southeastern Coastal Plain, all of these populations can be said to share similar natural environments. While the members of the respective populations originate from two different states, Florida and coastal Georgia, ecology does not recognize state lines. The area encompassing both states is predominantly subtropical and the salt marsh-barrier island coastal environment of northeastern Florida is identical to that of Georgia’s barrier islands (Milanich 1998; Hutchinson et al., 1998). While there are ecological variations within this coastal region and through time, such differences are minor from a more pandemic environmental perspective. Thus, I assume that the environment is a constant in this study. Unlike the aboriginal populations, I cannot conclusively establish that every individual from the contemporary population was a native Floridian or Georgian. However, since these people died in the last 40 years and were not indigent, I assume that they all benefited from similar levels of medical care and nutrition. Finally, given the proposed absence of a genetic component in levels of dental asymmetry (Chapter 2), the impact of the somewhat multiracial nature of the contemporary population is negligible. Thus, by examining these coastal populations of Florida and Georgia that date from the Middle Archaic Period (5000 to 3000 B.C.) to a modern population, I believe I have minimized some of the extraneous “noise” (e.g., in terms of geographic and genetic differences) found in other studies (Perzigian 1977). This allows me to discern asymmetry-related trends though time, in relation to drastic shifts in cultural practices. Hypothesis Based on previous studies on dental asymmetry (Chapter 2) and the cultural shift occurring in Florida and Georgia with European contact (Chapter 3) it was hypothesized that significantly elevated levels of dental asymmetry would be seen in populations

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5 experiencing both the transition from pre-agriculture to maize subsistence and the impact of Spanish colonization. The logic behind this hypothesis is based on the severe physiological stresses associated with these events (Chapter 3). Meanwhile, given advances in medical care and nutrition, the contemporary population would display the least degree of dental asymmetry. Furthermore, most literature suggests that sexual dimorphism, in relation to dental asymmetry is negligible (Chapter 2). Thus, a similar pattern was expected across the sites.

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CHAPTER 2 PREVIOUS WORK ON DENTAL ASYMMETRY Background Developmental homeostasis, or stability, can be viewed as an organism’s ability to produce an “ideal” form under a particular set of conditions. Decreased cellular stability increases the likelihood of departures from this “ideal” form. While a priori knowledge of “ideal” forms is rare, bilateral structures epitomize a template upon which to construct ideal, perfect symmetry. Thus, departures from bilateral symmetry can serve as a convenient tool for assessing deviations from the norm (Palmer 1994). Asymmetry within a population is the product of interactions between two independent, opposing processes; developmental stability (homeostasis) and developmental noise (i.e., developmental instability). As already mentioned, developmental stability is a suite of biological processes that buffer against disruption of the ideal ontogenetic trajectory. Developmental noise, on the other hand, is the result of biological processes that disrupt precise development; such as random differences in cellular division rates, random differences in the rate of physiological processes, and the effects of external stress on enzymatic activity. These processes serve to destabilize symmetry. As such, asymmetry can be viewed as an indirect measure of compromised developmental stability and/or high levels of developmental noise. For an individual to achieve perfect symmetry he or she must be able to buffer against developmental noise or the noise itself must be inadequate to elicit an abnormal morphological response (Palmer 1994). 6

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7 Dentition is a common structure used in analyzing departures from symmetry. Generally, asymmetry falls into two categories: Directional asymmetry (DA), which is the tendency for consistently greater development of one side over the other within a population. A certain degree of directional asymmetry (averaging 0.06 mm) is common in the human dentition. This phenomenon not only varies between teeth, but also in its direction. For instance, first premolars may be larger left than right, while canines may be larger right than left. Fluctuating asymmetry (FA), on the other hand, is small random differences between antimeric pairs. While this component is usually larger than directional asymmetry, actual differences in diameter are still not large in relation to inter-observer error and measurement precision (Hillson 1996). Previous Literature Fluctuating asymmetry (FA) is the form of human dental asymmetry most extensively noted in the literature. Studies have shown that FA will vary in magnitude among prehistoric and historic human populations (Doyle and Johnston 1977; DiBernnardo and Bailit 1978; Harris and Nweeia 1980; Townsend 1981; Mizoguchi 1986; Kieser et al.. 1986a; Kieser and Groeneveld 1988) between the dental arch, and within the tooth field (Townsend and Brown 1980; Bailit et al. 1970; Kieser et al. 1986). The presence of directional dental asymmetry (DA) in deciduous and permanent teeth has also been noted in several human populations (e.g. Corruccini et al. 1982; Sharma et al. 1986; Boklage 1987; Ben-David et al. 1992). Yet many researchers have overlooked this feature or attributed it to measurement error. However, this phenomenon was found to be significant in an adolescent white American population; manifesting as reversed directionality in opposing jaws. It has been postulated that the degree of directional asymmetry may be linked to either environment or genetic stress (Harris 1992; Sharma et al.. 1986). However, more work is needed in this area to elucidate the factors responsible for DA.

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8 Genetic Studies Dental asymmetry is approached in either genetic or ethnographic terms. Many genetic investigations examine developmental instability and the resulting asymmetry in terms of decreased genetic control over development. Studies of individuals with increased morbidity risk due to congenital abnormalities, also demonstrate elevated levels of FA in various structures, such as hand and foot breaths. Chromosomal and polygenic conditions such as Down Syndrome, familiar cleft lip (with or without palate), and schizophrenia display increased levels of FA compared to normal subjects (Livshits and Kobyliansky 1991). This suggests that a relationship exists between a genetic breakdown in developmental homeostasis and fluctuating asymmetry in humans. Increased homozygosity, as measured by the number of loci, may also be associated with decreased phenotypic variability and elevated asymmetry levels (Livshits and Kobyliansky 1991). However, inbreeding investigations have produced equivocal results. Some suggest a high genetic component (Boklage 1987), while others attribute asymmetry to environmental influences (i.e., climate, nutrition, etc.) (Potter and Nace 1976; Mizogueni 1987; Sharma and Corruccini 1987). Research by Livshits and Koloyliansky (1991) examining additive and non-additive genetic parental influences concluded that neither significantly contribute to individual FA traits. Rather, the study states that FA traits might be significantly correlated with an individual’s average degree of FA (i.e., all traits displaying asymmetry). The authors postulated that while individual FA traits acquire most of their variability from environmental influences, average FA may be potentially controlled by some genetic mechanism (Livshits and Kobyliansky 1991). Thus, while some genetic component may make one susceptible to the presence of asymmetry in

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9 various structures, external factors influence the degree to which each structural asymmetry is manifested. Ethnogeographic Studies Asymmetry is more typically examined in terms of ethnogeographic differences (i.e. environmental/cultural population dynamics). To understand asymmetry in ethnogeographic terms, it is necessary to comprehend stress as a causative agent. The word “stress” entered into the medical lexicon by the early 20 th Century. The new use of this word was based on the reaction of the sympathetic nervous system and adrenal medulla of laboratory animals subjected to adverse conditions; such as oxygen depravation, blood loss, and hypothermia. Noting that these animals produced a “stressed” physiological reaction; it was suggested that levels of stress could thereby be measurable (Kieser 1990). Asymmetry is largely discussed in terms of Selyian stress, based on work conducted by Canadian investigator; Hans Selye, in the 1970s. He documented the primary role of the anterior pituitary and adrenal cortex in the body’s response to nonspecific environmental stressors (e.g., elevated levels of heat, cold, and audiogenic stress). In addition, these stressors were found to produce asymmetrogenic effects in the teeth and limbs of lab animals (Sciulli et al. 1979). Hence, in population level investigations it has been suggested that the primary contributor of developmental noise is external; in the form of environmental stressors. These stressors, in turn, result in elevated FA levels. Because of this, many studies note the utility of FA as an indirect measure of the impact of environmental stress on populations. Bailit and his associates (1970) were among the first to test this presumed positive correlation between asymmetry and stress in non-twin human populations. Comparing the crown diameters of inhabitants of Tristan da Cunha, the Nasioi of Bougainville, the

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10 Kwaio of Malaita, and Bostonian children; the researchers anticipated that Tristanites would display the most asymmetry, Boston children the least, and the Masioi and Kwaio would fall variably in between. Their results followed this anticipated pattern; in terms of climatic, health, and nutritional differences between the groups. A comparison between precontact Native American skeletal populations and a modern cadaver population (Hamann-Todd) has also found the anticipated pattern of diminished levels FA through time, which was attributed to improved standards of living (Perzigian 1977). There appears to be a fairly consistent pattern of greater FA in the more posterior permanent teeth than the anterior teeth (Garn et al. 1981; Mayhall and Saunders 1986). Those teeth displaying greater variability in crown size also tend to display greater FA (Kieser and Groeneveld 1988). This may be potentially explained by the length of time a tooth is encased in soft tissue. The longer the developing tooth resists in the soft tissue, the more susceptible it is to environmental disturbances. While the deciduous dentition also shows some asymmetry, the anterior teeth display the greatest degree of variation (Townsend and Farmer 1998). Sexual Dimorphism in Dental Asymmetry Presented by Garn and associates (1965, 1966), the classic model of sexual dimorphism in dental asymmetry suggests the paired X chromosome confers greater dimensional control during ontogenesis. Hence females are better buffered against deviations from the norm. Nichol, Turner and Dahlberg (1984) echoed this sentiment of diminished asymmetry in males; postulating that the maternal intra-uterine environment would confer greater deleterious effects on males than females. However, most findings contradict these predictions; noting no evidence of sexual dimorphism in response to asymmetrogenic factors (Bailit et al. 1970; Perzigian 1977, Townsend and Garcia-Gody

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11 1984; Kieser et al. 1986a, 1986b; Kieser and Groeneveld 1988; Livshits and Kobyliansky 1989; Kieser 1992; Kieser and Groeneveld 1994). Thus, dental asymmetry in males and females, in particular FA; may increase due to increases in developmental noise, decreases in developmental stability, or a combination of the two. When extreme dietary, climatic, and geographic conditions exceed the buffering capability of developmental homeostasis; varying degrees of asymmetry result. Because of this, analysis of asymmetry can potentially elucidate the socio-environmental conditions under which communities exist.

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CHAPTER 3 GEOGRAPHIC AND TEMPORAL BOUNDARIES Introduction Having established the scientific framework of dental asymmetry, I now turn to the geographic, temporal and cultural parameters of the research material. As already mentioned, time and more importantly societies rarely establish their boundaries based on the arbitrary lines that separate territories and states. That being said, the native communities of Georgia and Florida have greatly impacted each other’s history. As such, it is necessary to briefly outline the history of coastal Florida and Georgia indigenous cultures through time. This brief sketch will outline presumed cultural, nutritional, and, consequently biodevelopmental stressors endured by these peoples. Admittedly there have been climatic and geographic fluctuations in this coastal region within the last 12,000 years, most specifically elevated water levels. Nonetheless, I hold the environments constant, and view the societal exploitation of the habitat as variable. This summary is in no way fully representative of Florida and by consequence the latter part of Georgia history in its entirety. Instead, it offers an overview of the region through time; focusing on particular periods which are relevant to understanding the cultural context of the research material. Natural Environment The coastal zone encompasses the most ubiquitous part of Florida because of its peninsular shape; as well as a portion of Georgia. The coastal strip of Florida consists of the Atlantic Ocean and the Gulf of Mexico. The terrain of the beach and foreshore is 12

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13 generally inhospitable because of seasonal exposure to storms and scarce resources, especially along the Atlantic coast. More inland, the sandy beaches and dunes become muddy shores thick with vegetation, creating lagoons or marsh areas. Sheltered by the winds and fed by drainage from the interior; these areas host a diverse array of organisms, and serve as an interchange between marine and inland species (Milanich and Fairbanks 1978). Moisture rich areas; such as along streams and lakes of northern Florida and Georgia, are made up of hardwood hammocks. Characterized by a broad spectrum of plant communities, these dense forests provide shelter and food for a vast array of animals. The environment of coastal Georgia consists primarily of these types of wetlands. Chains of barrier islands are also found along the eastern coastline of Florida and Georgia containing beach and dune landscapes, live oak hammock, tidal flats, and estuary habitats (Milanich 1998, Milanich and Fairbanks 1978; Wallace 1978). Cultural Periods Paleoindian Period The earliest evidence of human occupation in Florida consists of a spear point made from the ivory tusk of a mammoth, dated to shortly after the last Ice Age. This suggests the region was inhabited at least 12,000 years ago. These inhabitants, known as Paleoindians, were skilled in hunting the large game prolific during the late Ice Age (McGoun 1993; Milanich 1998). Descended from people who crossed into North America from eastern Asia during the Pleistocene epoch (1,600,00 B.C. to 10,000 B.C.), these nomadic inhabitants were presumably drawn to the region by the Pleistocene megafauna in Florida; such as mastodons and extinct bison (Milanich 1998). Water was also prevalent in this region. This was a precious commodity at the height of the Ice Age where the sea level had fallen as much as 300 feet beneath current levels creating a cool,

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14 arid climate (McGoun 1993; Milanich 1998). Because of the nomadic nature of these people, they did not have prescribed burial grounds (Milanich and Fairbanks 1978). Archaic Period Towards the end of the Ice Age, temperatures increased and sea levels rose as glaciers began to melt. These environmental changes led to the extinction of many of the species, which had previously thrived in the area. As a consequence, patterns of food procurement were shifted. This shift in Paleoindian lifestyle ushered in the Archaic Period (7500 B.C. to 500 B.C.) (Milanich 1998). With the extinction of many big game animals, Archaic peoples developed a broader range of plant and animal subsistence; whereby they “specialized in nothing, but were versatile in attempting everything”(Farb 1968:206). With the climate becoming wetter and rising seas levels diminishing land utilization, broad exploitation became prudent. Between 3,000 and 2,000 B.C. signs of pottery manufacture arose, suggesting at least a partially sedentary existence. Within the next millennium, a small degree of horticulture also became evident. The presence of larger and more numerous archaeological sites dating to this period suggest the people adapted well to their new environment (McGoun 1993; Stepanitis 1986). Prescribed burial grounds also began to surface during this period. Some Archaic populations interred their dead within the layers of peat found at the bottoms of ponds. These burials bear little resemblance to the drier peat bogs of Europe and the British Isles. In fact, excavations of such sites have been likened to “trying to dig chocolate mousse underwater” (Milanich 1998; 16). Bodies were wrapped in fabric and anchored by wooden stakes to the pond bottom, thus submerging the remains in loosely consolidated peat (Beriault et al. 1981; Milanich 1998).

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15 Shell burial mounds have also been discovered dating to this period. These burials generally consisted of digging a shallow hole into an existing shell midden (i.e., shell refuse heap), placing the often flexed, wrapped body inside and covering it with sand. This procedure would be preformed several times over, eventually forming a mound (Milanich 1998). Late Prehistoric Periods While the southern third of Florida remained basically in the Archaic stage until the arrival of the Spanish in the sixteenth century (McGoun 1993), the rest of Florida underwent continual alterations in lifeways through interaction with other groups. This period, known as the Woodland (500 B.C. to 900 A.D.), is marked by the beginnings of settled communities, increased complexity of political and religious organization, and elaboration of ceramic and burial rituals (Milanich 1998; Milanich and Fairbanks 1978; Stepanitis 1986). These continual alterations are not currently well understood by archaeologists. As a result, there are many names for various cultural phases which make up the Woodland Period. As not all of these cultural sub-periods are relevant to this discussion, I instead focus on those cultural periods that the research materials are associated with. To assist in this endeavor, I have included a table at the end of the chapter depicting the temporal ranges, geographic distribution, and relevant characteristics of the various cultural periods mentioned (Table 3-1). Coastal dwellers living from 500 B.C. to 900 A.D. in Florida are referred to as the Manasota phase of Woodland culture. Shell middens often encircled these sites, and prolific marine resources allowed occupation of the same location for many years. After 300 A.D. shell mounds were often associated with charnel house use (Milanich 1998).

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16 Charnel houses were constructions in which the dead were macerated prior to the placement of their bundled remains within the mound (Sears 1960). This elaboration in mortuary ritual is suggestive of increased population size (Milanich 1998). Contemporary with the Manasota culture along the coast, Weeden Island cultures appeared throughout northern Florida from 250 A.D. to 750 A.D. This area (Figure 3-1) is north of a line running southwestward from Cocoa to Bradenton, or more simply the region north of the winter freeze limit (McGoun 1993). Figure 3-1. Line dividing northern and southern Florida. Weeden Island villages often included vast earthen work mounds and distinctive pottery. These peoples did not grow corn; though some gardening may have been practiced. This lack of renewable food reserves often led to a cyclical pattern of growth, expansion, and eventual decline as wild food collection was unable to sustain growing settlement populations (Milanich 1998). Trade routes allowed the sharing of goods and ideas between different groups; thus it is not unusual to come across sites containing

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17 material culture reflective of multiple groups. Sites containing material culture reminiscent of both Manasota and Weeden Island culture are often referred to as Manasota Weeden Island culture. Between 750 A.D. and 900 A.D. another shift in cultural practices occurred. As ideas flowed along trade routes, the Woodland culture era eventually morphed into what archaeologists refer to as Mississippian culture. In Florida, this change solidified by 900 A.D. and became known as the Safety Harbor phase of Mississippian culture. The distinctive difference between Woodland and Mississippian is most apparent in pottery forms and function, socio-political practices, and a marked increase in horticulture. However, as before, the people still primarily relied upon wild resources as their main source of food. The heart of Safety Harbor culture was centered around Tampa Bay in Pinellas, Manatee, Hillsborough, and northern Sarasota counties. This region was south of any agriculture that was being practiced in the area. Intensive agriculture, as practiced above the Fall Line (i.e., Macon, Georgia and up), was not possible in this area. The soil inland and around Tampa Bay was unsuitable for the slash-and-burn techniques that characterized maize agriculture from this period. Instead, wild resources were maximized through more centralized social and political systems. While centers for small chiefdoms dotted the Tampa Bay area, larger capitals never manifested (Milanich 1998). Even though the specific phases within larger cultural phases (i.e., Safety Harbor within Woodland) are given different names within Georgia, the overall socio-economic cultural context is the same as those described for Florida. Because of this, detailed discussion of Georgia’s cultural phases is not necessary.

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18 European Contact While inhabitants of Florida and Georgia experienced significant social changes in the Woodland and Mississippian periods, nothing would prepare these people for the drastic fallout experienced with the arrival of Europeans. In less than 200 years, colonialism would decimate these native groups. The first documented exploration of Florida was by the Spanish in 1513. By 1565, the Spanish had established their first permanent settlement at St. Augustine, FL. The Spanish then went on to secure a series of Roman Catholic missions along the Atlantic coast of Georgia and Florida, as well as in northern Florida (Hutchinson et al.1998). Missions were primarily composed of Spanish and native groups, such as the Guale from Georgia and Florida’s Timucua and Apalacuee. These people were farmers, and as such, had agricultural expertise that was exploited by the Jesuit monks and Franciscan friars. The Spanish used the Native Americans as a labor force in corn production and processing. Their mission system created dramatic changes in native settlement behavior, diet and labor. This system harnessed native villagers as laborers, forcing them to reorganize themselves into sedentary agricultural communities (Milanich 1995; 1998). A maize based diet, in conjunction with the elevated stress and pathogen levels encountered with the arrival of the Spanish, had severe physiological consequences on these native populations. Maize is deficient in the essential amino acids lysine and tryptophan. In addition, the presence of phytate in maize reduces the body’s available iron, leading to iron deficiency anemia. Skeletally, this manifests as increased levels of porotic hyperostosis and cribra orbitalia. These conditions are commonly seen in late prehistoric and postcontact Florida and Georgia populations (Hutchinson et al. 1998).

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19 Additionally, increased levels of osteoarthritis and biomechanical analysis of femoral and humeral diaphyses suggest the mission workload was substantially more severe than prior levels. There is also a dramatic increase in carious lesions, enamel hypolasia, and periosteal reactions in populations from this time period. All of these factors are indicative of increased physiological stress, as compared to earlier precontact populations (Hutchinson 1998). Thus, data suggest the increased consumption of maize by precontact peoples, particularly in Georgia, produced declines in health; which became magnified by the elevated workloads and Old World infections experienced during the mission period. Old World epidemics struck northern Florida missions in 1595, 1612-1617, 1649-1650, and 1655-1656, killing both missionized and nonmissionized indigenous trading groups throughout the region. Spanish officials attempted to counteract these increasing mortality rates by repopulating existing missions with more numerous native groups from Georgia and even South Carolina. Meanwhile, British militiamen and their native allies from the Carolinas further weakened Spain’s control over the region by organizing attacks on coastal missions in 1660 and 1684 (Milanich 1998). Not having the resources to defend these sites, the Guale and Timucuan missions on the Georgia coast were abandoned. By 1665 the northernmost mission was on Amelia Island, a scant 50 miles north of the original St. Augustine settlement. These scattered raiding tactics by the British continued on, and by 1710 the majority of the native population was dead or had fled to neighboring territories. A 1717 census of the remaining 10 Spanish refugee villages in Florida indicates the indigenous populations had dwindled down to 942 individuals. British occupation in 1763 resulted in the exodus of virtually all of the

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20 remaining native Floridians into Cuba. Meanwhile, the Creek Indians, also from Georgia, infiltrated inland Florida. The few remaining Timucuans and Apalachees were subsumed into the Creek population. This population was further expanded by the later arrival of runaway African slaves. Later known as the Seminole, the descendents of this population continue to reside in Florida (Milanich 1998). Table 3-1. Relevant cultural periods in Florida and Georgia Period Dates Geographic Range Settlement and Subsistence Paleoindian 12,000-7500 B.C. North America Nomadic, hunter-gathers Archaic North America Early 7500-5000 B.C. Middle 5000-3000 B.C. Late 3000-500 B.C. Broader subsistence strategy, prescribed burial grounds Woodland North America Manasota 500 B.C.-900 A.D. Coastal Florida Weedon Island 250-750 A.D. Northern Florida settled communities, elaboration of burial rituals increased complexity of political and religious organization Mississippian North America Safety Harbor 900 A.D.-contact Northern Florida Marked increase in horticulture Increase socio-political practice Contact 1500s Florida and Georgia Implementation of mission system, maize based subsistence, introduction of European pathogens Modern 20th Century Florida 20th Century

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CHAPTER 4 SITES Introduction Of the six archaeological sites utilized in this study, five (Bay West, Tick Island, Bayshore, Palmer, Harrison Homestead) are located in counties along the peninsular coast of Florida. The sixth site lies on St. Simons Island, which is part of the chain of barrier islands along the southern coast of Georgia (Figures 4-1 and 4-2). Finally, a modern sample is also included in the research material. Chronologically, these populations represent the Archaic, Manasota Weeden Island, Safety Harbor, European contact and modern periods. A brief description of the archaeological investigations conducted at these sites follows. Site numbers are provided for each of the sites investigated. Figure 4-1. Location of St. Simons in Georgia. Photo courtesy of Thomas Whitley. 21

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22 Florida Sites Harrison Homestead Tick Island Bayshore Palmer Bay West Figure 4-2. County location of Florida samples. Notice how the counties are located on the coast. Bay West (8CR200) The Bay West Site is located on the lower western portion of southern Florida in Collier County. It resides 9-10 km east of the Gulf of Mexico Coastline. The site was discovered in1980 during a dredging operation at the Bay West Nursery (Figure 4-3). Workers uncovered human bones, while removing peat from a cypress pond for use in the nursery. Unfortunately, this peat mining heavily disturbed the site before volunteers and members of the Southwest Archaeological Society could be brought in to excavate

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23 the area. Luckily the team, headed by John Beriault, was still able to collect some important information. Radiocarbon-14 dating on the wooden posts and peat associated with the burials dated from 4900 to 4000 B.C., placing it within the Middle Archaic Period (5000 to 3000 B.C.). This is consistent with the time period of projectile points also found within the pond and adjacent area. Thirty-five to forty individuals were buried in the Bay West Pond. The anaerobic conditions of such freshwater peat environments serve to inhibit decay processes and assist in preservation of remains (Beriault et al., 1981). Figure 4-3. Map of Bay West site (8Cr200). The northern portion of the map contains the mortuary pond from which the research material came from. Drawing by Shanna Williams. (Adapted from: Beriault, J. et al. 1981.“The Archeological Salvage of the Bay West Site, Collier County, Florida.” Florida Anthropologist, Vol. 34, Figure 1, p. 41.)

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24 Tick Island (8VO24) The Tick Island Site was excavated by Ripley R. Bullen, of the Florida State Museum (now known as the Florida Museum of Natural History), in 1961. The site consists of a prehistoric mortuary located within a large freshwater snail shell mound. Tick Island, located on the upper eastern portion of the state, is surrounded on all sides by lakes formed from the St. Johns River. The site, itself, is located on the southeast side of the island and is made up of four areas designated A to D. Area D contained the skeletal material examined in this study (Figures 4-4). This shell mound was a flat platform that was periodically enlarged during a minimum of seven construction phases. One hundred-seventy-five people were buried in the lower layers of the mound. The top of the mound was made up of discarded shell refuse from later inhabitants living at the site. Many of the elements found within the shell mound were heavily mineralized or concreted. Radiocardon dates from the burials date to 3500, 3370, and 3080 B.C.; placing them within the Middle Archaic period as well (Aten 1999; Jahn 1978). Figure 4-4. Tick Island (8VO24) sketch as it may have been before quarrying. Letters identify the main areas of the site. Area D, the large mound in the rear, contained the skeletal material used in this analysis. Drawing by Shanna Williams. (Adapted from: Aten, L.E.1999 “Middle Archaic ceremonialism at Tick Island, Florida: Ripley P. Bullen’s 1961 excavation at the Harris Creek Site.” The Florida Anthropologist, Vol. 52, No. 3, Figure 3, p. 137).

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25 Palmer (8SO2) Also excavated by Ripley Bullen and his wife Adelaide between 1959 and 1962, the Palmer Site is a prehistoric village complex located in a coastal gulf setting south of Tampa Bay, in Sarasota County. The burial mound component of the site is located in the southeastern portion of the site (Figure 4-5). This unassuming sand mound rises only four feet above the surrounding land due to recent fill, but was undoubtedly quite impressive during its use. Archaeological material found within the different zones of the mound suggests sporadic construction of the mound. Approximately 400 individuals are represented by fragmentary remains. Based on radiocarbon dates and ceramics, the mound probably dates from 250 to 750 A.D. (Bullen and Bullen 1976). Based on its location and material culture, the site falls within the Manasota Weeden Island period. Figure 4-5. Palmer (8SO2) sketch. Drawing by Shanna Williams (Adjusted from: Kozuch, L.1998. “Faunal Remains from the Palmer Site (8SO2), with a focus on shark remains.” The Florida Anthropologist. Vol. 51, No. 4, Figure 1, p. 178).

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26 Bayshore Homes (8PI41) The Bayshore Homes site lies in the Central Peninsular Gulf Coast in the northwestern corner of St. Petersburg. The site was excavated by William Sears of the FMNH in 1958 and includes three earthen work mounds and a shell midden. Of the three mounds, the one designated Mound B was the most prominent and well known (Figure 4-6). This made it a target for treasure hunters, which resulted in some disturbance of the material in the upper layers (Sears 1960). About 145 individuals are represented by the fragmentary human remains, which were examined by Charles E. Snow (Snow 1962). These individuals reflect secondary burials from a charnel house. Pottery sherds found within the site date from 750 A.D. to 1000 A.D.; placing it within the later Manasota Weeden Island period and the early Safety Harbor Period (Sears 1960). Figure 4-6. Bayshore Homes (8PI41) sketch. Drawing by Shanna Williams (Adapted from: Sears, W.H.1960 “The Bayshore Homes Site, St. Petersburg, Florida.” Contributions of the Florida State Museum. Social Sciences. No 6. University of Florida, Gainesville, Figure 1, p. 2).

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27 St Simons Island Sites (9GNXXXX) St. Simons Island is part of a chain of barrier islands along the Georgia Coast (Figure 4-1). Numerous excavations have been conducted on the island, the earliest of which was performed by Clarence Moore at the end of the 19 th Century. This site was later investigated by Ronald Wallace from the University of Florida and designated Couper Field (Pearson and Cooke 2003, Wallace 1978). Now an abandoned field, the site is on the northern part of the island. Sixteen burials were discovered in below ground grave pits. Associated pottery indicates occupation during the late precontact/ early European contact period (1450 to 1550 A.D.) (Pearson and Cooke 2003; Wallace 1978). Additional skeletal material was found at Taylor Mound (9GNXXXX); also located on the northern end of St. Simon’s Island, south of Couper Field. The initial excavation was conducted by Charles Pearson and Fred Cooke in the early 1970s and was subsequently re-excavated by Jerald Milanich in 1973. Representing a ceremonial mound with associated burials, the structure is located in a grove of trees undisturbed by plowing. The site, which contains a combination of undisturbed and re-excavated burials, dates to the historic period (1600-1650 A.D.) and is representative of nonmissionized Timucua Indians (Pearson and Cook 2003; Wallace 1978). Finally, these two Georgian sites, as of yet, have not been given an official site number and thus excavations from this area are designated as 9GNXXXX. Harrison Homestead (8NA 41) Amelia Island is located on the east coast of Nassau County near the Florida-Georgia border (Figure 4-7). The site on this island, known as Harrison Homestead, contains two 17 th Century Spanish Franciscan missions. Of interest in this study is the Santa Maria Mission Church/Cemetery. The site was excavated by archaeologists from

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28 the FMNH in conjunction with Clark Larsen in the late 1980s. The 118 burials were uncovered from within the structure of a church. The majority of the remains displayed burial postures consistent with other mission periods (i.e. heads aligned southwest and feet to the northeast). The church served a Yamasse population, which were refugees from north Georgia and South Carolina. They resided at the mission till 1683 when the Spanish crown relocated them to a mission on St. Catherine’s Island in Florida. Documentation from Franciscan priests indicates occupation of Santa Maria extended from 1675 to 1683; till raids by the English forced relocation. The high level of burials in these eight years of occupation suggests 31.25 deaths a year at Santa Maria (Saunders 1988). Figure 4-7. Harrison Homestead (8NA41) sketch. Star indicates location on Amelia Island.

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29 Contemporary Sample Finally, a modern population housed at the University of Florida’s Forensic Research Laboratory was analyzed. The sample consists of mandibles and maxillae of known sex and ancestry. Plaster casts representing a single individual are also present. Age ranges extend from late teens to late fifties. Both sexes are fairly evenly represented, and the vast majority of the individuals are classified as white. Temporal Series Table 4-1 summarizes the chronological time periods encompassed by these skeletal populations. Roughly 7000 years is covered by this material. The assumption of continuous, predominantly in situ cultural development has been made; given restricted geographic (and ecologically similar) region and temporal continuity among the members of the respective samples. In other words, the presence of environmental (i.e., cultural) stressors should be observable and measurable within these populations.

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30 Table 4-1. Site ranges and cultural periods Period Dates Sites Site Dates Paleoindian 12,000-7500 B.C. Archaic Early 7500-5000 B.C. Middle 5000-3000 B.C. Tick Island, Bay West 3500, 3370, 3080 B.C. Late 3000-500 B.C. 4900-4000 B.C. Woodland Manasota 500 B.C.-900 A.D. Weedon Island 250-750 A.D. Palmer 250-750 A.D. Mississippian Bayshore Homes 750-1000 A.D. Safety Harbor 900 A.D.-contact Contact 1500s St. Simons 1450-1550 A.D. Harrison Homestead 167-1683 A.D. Modern 20th Century Contemporary Sample 20th Century

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CHAPTER 5 MATERIALS AND METHODS Materials The archaeological samples are stored in the Florida Museum of Natural History, University of Florida, Gainesville. The contemporary population, meanwhile, is housed in the Forensic Research Laboratory, also at the University of Florida. In total, the samples represent 109 individuals; however, the total number of each tooth analyzed varies as few individuals displayed a complete dentition. Furthermore, poor preservation of bone from some of the archaeological sites precluded utilization of every individual recovered. Occlusal Surface Wear Dietary factors, such as pre-agricultural consumption of coarse foods (Smith 1984; Larsen 1995), caused elevated levels of dental attrition in the majority of the archaeological populations. Severe attrition can led to miscalculation of asymmetry levels, as the enamel which accounts for tooth length and width is worn and/or chipped away. To mitigate this phenomenon, those teeth which exceeded particular wear stages outlined by Smith (1984) and Scott (1979) were excluded. The Smith (1984) system outlines a surface wear scoring system for incisors, canines, and premolars with wear stages extending from one to eight. As the presence of enamel is necessary to provide accurate measurement of dental dimension, teeth exceeding stage 6 (Figure 5-1) were excluded. 31

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32 Figure 5-1. Smith surface wear scoring system. Drawing by Shanna Williams (Adapted from: Buikstra J.E. and Ubelaker D.H.1994. Standards for Data Collection from Human Skeletal Remains. Fayetteville, Arkansas, Arkansas Archaeological Survey Report No. 44. Figure 25, p. 52) Molar attrition rate was based on the Scott System (1979); whereby each occlusal surface is divided into quadrants. The enamel in each quadrant is scored on a scale of 1 to 10 (Figure 5-2). Using an adjusted version of this system, the boundary for molar inclusion was set at levels 1-5 and 8 (i.e., medium to thick enamel must be exhibited on each quadrant’s outer rim). Thus, teeth falling into levels 6 and 7 (i.e., lacking a complete outer rim) were excluded. Once again, this exclusion was conducted to control for inaccurate dimensional measurements due to enamel lose.

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33 Figure 5-2. Scott surface wear scoring system. Drawing by Shanna Williams. (Adapted from: Buikstra J.E. and Ubelaker D.H.1994. Standards for Data Collection from Human Skeletal Remains. Fayetteville, Arkansas, Arkansas Archaeological Survey Report, No. 44, Figure 26, p. 53). As occlusal wear increases with age (Molnar 1972), limiting the degree of attrition in the data also serves to constrain age somewhat. As it is generally agreed that asymmetry levels are related to exposure to environmental stressors during dental development, it is only relevant to establish that the research individuals have reached adulthood and still possessed intact enamel. Thus, the selection criteria for research subjects required that their dentition include fully erupted permanent second molars and

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34 have the outer rim of enamel present. As the presence of intact incisors was rare, and the majority of those available displayed partial or complete lose of enamel, incisor data was not used in this analysis. The third molar was also excluded. As the most unpredictable of all the tooth types, the third molar tends to deviate in form, if not missing entirely (Hillson 1996). Therefore, only the canine, premolars, and first and second molar antimeres were measured when present. The following notations will be used to represent each tooth type: C1 = canine; PM3 = third premolar; PM4 = fourth premolar; M1 = first molar; and M2 = second molar. For those unfamiliar with the PM3 and PM4 notation, PM3 represents teeth 4, 12, 21, and 28, while PM4 represents teeth 5, 13, 20, 29 in the Universal/National System used by the American Dental Association. Sexing As sex differences may potentially contribute to variation in asymmetry levels, sex was determined using the scoring system for sexually dimorphic cranial features as outlined by Buikstra and Ubelaker (1994). Sex determination based on os coxae morphology (Buikstra and Ubelaker 1994) was also used in conjunction with cranial features when available, and site documentation indicated the absence of commingling. To expedite the process, reliable published and unpublished sexing information on the sites was used when available (Saunders 1988; Sears 1960; Wallace 1978;). Any individuals who could not be sexed with certainty were excluded from the analysis. Dental Measurements Buccolingual and mesiodistal diameters of dental crowns were recorded following measurement definitions outlined by Mayhall (1992). Mesiodistal diameter (MD) was defined as the maximum width of the tooth crown in the mesiodistal plane. This method allows for greater accuracy in cases of interproximal wear and malocclusion (Mayhall

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35 1992). Buccolingual diameter (BL), meanwhile, was defined as the widest diameter of the tooth, measured perpendicular to the mesiodistal plane. Measurements were recorded to an accuracy of 0.01 mm using digital calipers (Mitutoyo Digital Point Jaw Calipers) interfaced to a microcomputer. This technique was repeated at a later date on 30 randomly selected subjects to assess measurement error. In no instance did the mean difference between double determinations differ significantly from zero at p<0.05. Therefore, the replicability trial indicates no significant observer bias in the tooth measurement data. Statistical Methods Dental asymmetry was quantified in two ways. First, fluctuating asymmetry (FA) was calculated by transforming the data to eliminate individual size differences among teeth (Harris and Nweeia 1980). Under this rescaled asymmetry measure, (d*), the absolute value of the side difference is divided by the mean size of the left and right teeth. d* = L R ( L + R) 2 Second, directional asymmetry (DA) was calculated by subtracting the crown size difference between left and right antimeres, (d = L – R), in the mesiodistal and buccolingual dimension. A negative value for DA indicates the right antimere to be larger than the left. Since the sites are serving as a proxy for time, the datasets were compared against on another. The presence of significant time and sex differences within and between the samples was assessed by multiple regression analysis for each tooth in the buccolingual and mesiodistal dimensions.

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36 The two factor design model for this data is yijk = + i + j + ()ij + ijk where yijk = the kth tooth measurement for the ith sex group and the jth site = the overall mean i = main effect of sex j = main effect of site ()ij = first order interaction term between of site and sex between pairs of treatments ijk = the residual error terms This multiple regression assumes normal distribution with mean zero and variance . Furthermore, this model serves to partition the variation in the observations into parts based on each main effect (sex and site) and the first order interaction between the two effects. This partition leads to a series of F-tests which assess the significance of the various components. The model was applied to the calculated values of d* and d, respectively, using SAS version 8.2. Due to the unbalanced design, Type III Sums of Squares results represented the contribution of each factor and the interaction effect, with significance levels set a p<0.05. To assist in interpretation, the mean results for DA and FA were visualized using the graphing function in Excel.

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CHAPTER 6 RESULTS The results are organized in terms of fluctuating asymmetry and directional asymmetry. Within each category the effects of sex, site and sex*site interaction are presented. Tables 6-1 through 6-4 provide the ANOVA results for both types of asymmetry. Highlighted cells denote p-values which reached significance at the overall multiple comparison level of less than 0.05, with yellow indicating p<0.05 and blue indicating p<0.01. Tables 6-5 and 6-6 provide the mean difference between sites demonstrating significant differences for FA and DA. The graphs included within this chapter represent only those teeth which were found to be significant by ANOVA. In cases where the x-axis represents sites, the sites are arranged in temporal order, with the Bay West population at the far left and the contemporary population at the far right. While infrequent, there are some instances of mean values which visually appear to be significant. This can be accounted for by such mathematical phenomena as small sample sizes (particularly in the Bay West population), and/or large variance values within some of the samples. Specific FA values for each site and by sex can be found in Appendix A and B. In order to prevent these cases from distracting the reader, I have included arrows or circles in the figures to indicate which specific sites or teeth demonstrate statistical significance. Finally, as more cases of statistical significance were seen in the analysis of fluctuating asymmetry, the results for this form of asymmetry are discussed first, followed by a similar discussion for directional asymmetry. 37

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38 Table 6-1. Maxillary values for fluctuating asymmetry Buccolingual Mesiodistal Tooth Effects DF F P DF F P C1 Sex 1 0.01 0.928 1 0.33 0.5685 Site 6 1.39 0.2322 6 1.61 0.1606 Sex*Site 4 0.25 0.906 4 0.18 0.9464 M3 Sex 1 0.18 0.6749 1 3.63 0.0609 Site 6 1.22 0.3048 6 2.8 0.0168 Sex*Site 5 1.1 0.3685 5 0.43 0.825 PM4 Sex 1 0.14 0.7072 1 3.35 0.0712 Site 6 2.33 0.0411 6 1.95 0.0831 Sex*Site 5 0.73 0.6068 5 0.97 0.444 M1 Sex 1 0.91 0.3427 1 0 0.9939 Site 6 1.03 0.4161 6 0.68 0.6648 Sex*Site 5 2.54 0.0357 5 1.75 0.1354 M2 Sex 1 2.07 0.155 1 3.04 0.086 Site 6 5.63 <.0001 6 2.54 0.0288 Sex*Site 5 2.53 0.0374 5 2.18 0.0668 Note: BL = buccolingual. MD = mesiodistal. DF = degree of freedom. Sex*Site= interaction effect. F = F value. P = P value. Table 6-2. Mandibular values for fluctuating asymmetry Buccolingual Mesiodistal Tooth Effect DF F P DF F P C1 Sex 1 1.97 0.1664 1 2.07 0.1559 Site 4 1.15 0.344 4 4.94 0.0017 Sex*Site 4 0.66 0.6199 4 2.88 0.0307 PM3 Sex 1 0.57 0.4522 1 0.01 0.9125 Site 6 1.65 0.1462 6 2.45 0.033 Sex*Site 5 0.84 0.5268 5 0.57 0.7224 PM4 Sex 1 4.57 0.0361 1 9.9 0.0024 Site 6 4.44 0.0007 6 4.36 0.0009 Sex*Site 5 7.4 <.0001 5 4.75 0.0009 M1 Sex 1 0.03 0.859 1 1.1 0.2981 Site 6 0.92 0.4848 6 1.35 0.25 Sex*Site 5 1.31 0.27 5 1.96 0.0971 M2 Sex 1 0.51 0.479 1 0.49 0.4883 Site 6 0.45 0.8405 6 1.31 0.2672 Sex*Site 5 0.49 0.7843 5 0.43 0.8241

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39 Table 6-3. Maxillary values for directional asymmetry Buccolingual Mesiodistal Tooth Effects DF F P DF F P C1 Sex 1 0.09 0.7694 1 3.83 0.0549 Site 6 1.13 0.3551 6 0.76 0.6037 Sex*Site 4 0.5 0.7345 4 0.61 0.6553 PM3 Sex 1 1.42 0.2368 1 0 0.9778 Site 6 1.26 0.2896 6 2.53 0.0279 Sex*Site 5 0.15 0.98 5 0.63 0.6803 PM4 Sex 1 0.06 0.8061 1 0.22 0.6399 Site 6 0.65 0.6929 6 1.86 0.0978 Sex*Site 5 0.32 0.8982 5 1.32 0.2629 M1 Sex 1 0.05 0.8203 1 0.09 0.7665 Site 6 0.69 0.657 6 1.37 0.2403 Sex*Site 5 0.28 0.9243 5 2.82 0.0223 M2 Sex 1 0.79 0.3779 1 0.03 0.8593 Site 6 0.64 0.6963 6 0.91 0.4927 Sex*Site 5 0.39 0.8567 5 0.19 0.9669 Note: BL = buccolingual. MD = mesiodistal. DF = degree of freedom. Sex*Site= interaction effect. F = F value. P = P value. Table 6-4. Mandibular values for directional asymmetry Buccolingual Mesiodistal Tooth Effect DF F P DF F P C1 Sex 1 0.16 0.6864 1 3.31 0.0742 Site 4 0.41 0.8011 4 2.01 0.1054 Sex*Site 4 0.37 0.828 4 0.66 0.6248 PM3 Sex 1 0.01 0.9157 1 0.22 0.6398 Site 6 2.28 0.0461 6 2.35 0.04 Sex*Site 5 1.1 0.3696 5 1.2 0.3178 PM4 Sex 1 0.92 0.3406 1 4.06 0.0479 Site 6 0.42 0.861 6 1.02 0.4185 Sex*Site 5 0.27 0.9257 5 0.34 0.8881 M1 Sex 1 0.01 0.9125 1 0.06 0.8127 Site 6 1.97 0.0831 6 0.36 0.9014 Sex*Site 5 1.5 0.2022 5 1.25 0.2966 M2 Sex 1 0.38 0.5414 1 4.2 0.0446 Site 6 0.41 0.8699 6 1.4 0.2295 Sex*Site 5 1.51 0.2 5 0.57 0.7262

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40 Table 6-5. Fluctuating asymmetry site differences tooth Site Comparison Mean Difference between sites Simultaneous 95% Confidence Intervals BUCCOLINGUAL Mandible PM3 St. Simons vs Forensic 0.023433 0.00017 0.047*** PM4 Palmer vs Bayshore 0.044674 0.0046-0.085*** Palmer vs Harrison 0.044947 0.0018 0.088*** Palmer vs St Simons 0.046412 0.0070 0.086*** Palmer vs Forensic 0.055064 0.016 0.094*** MESIODISTAL CI Bayshore vs Forensic 0.041938 0.011 0.072*** PM3 Palmer vs Forensic 0.0401 0.0061 0.074*** PM4 Bayshore vs Forensic 0.0638 0.016 0.11*** Palmer vs Forensic 0.04014 0.00440.076*** Maxilla BUCCOLINGUAL PM4 St Simons vs Forensic 0.019736 0.0010 0.038*** St Simons vs Tick Is. 0.022515 0.0017-0.043*** M1 St Simons vs Forensic 0.014152 0.0012 0.027*** M2 Bayshore vs Forensic 0.018237 0.0048 0.032*** St Simons vs Forensic 0.018236 0.0064 0.030*** MESIODISTAL PM3 St Simons vs Forensic 0.04127 0.0078 0.075*** M2 Bayshore vs Forensic 0.033876 0.0015 0.066*** Note: *** = comparisons significant at the 0.05 level. Table 6-6. Directional asymmetry site differences tooth Site Comparison Mean Difference between sites Simultaneous 95% Confidence Intervals BUCCOLINGUAL Mandible PM3 St Simons vs Forensic 0.21385 0.0059 0.422*** MESIODISTAL PM3 Bayshore vs Tick 0.51833 0.057 0.98*** Maxilla MESIODISTAL PM3 St Simons vs Forensic 0.32528 0.0078 0.64*** St Simons vs Palmer 0.34592 0.032 0.66***

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41 Fluctuating Asymmetry Sex Differences In terms of fluctuating asymmetry no significant sexual dimorphism was found between males and females in the maxilla. However, the mandible did show sex differences in both dimensions of PM4. In both instances, males display greater levels of FA than females, with the differences slightly less in the buccolingual dimension than in the mesiodistal dimension (Figure 6-1). 00.010.020.030.040.05CIPM3PM4M1M2ToothRescaled Mean Females Males 00.010.020.030.040.05CIPM3PM4M1M2ToothRescaled Mean Females Males A B Figure 6-1. Mandible sex differences (FA). A) Mesiodistal dimension. B) Buccolingual dimension. Note: Circles indicate significant sex differences for a particular tooth. Site Differences In the maxilla, three of the five teeth examined in the buccolingual dimension display significant site effects, most notably the posterior cheek teeth (PM4, M1 and M2). All three cases involve the teeth from St. Simons being larger than those same teeth in the contemporary population. This indicates that St Simons has significantly more asymmetry in these teeth than the contemporary population. St. Simons also has more FA than Tick Island in PM4. Finally, in addition to St. Simons, Bayshore also has more FA than the contemporary population for M2 (Figure 6-2).

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42 0.010.020.030.04escaled mean 0Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiter Buccolingual 00.010.020.030.04Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiterescaled mean Buccolingual A B C 00.010.020.030.04Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiterescaled mean Buccolingual Figure 6-2. Maxilla site differences (FA) in buccolingual dimension. A) PM4. B) M1. C) M2. The mesiodistal dimension of the maxilla demonstrates significant site differences in PM3 and M2. These site differences include St Simons, Bayshore, and the contemporary population. In the case of PM3, St Simons has more FA than the contemporary population. Meanwhile, Bayshore has more FA than the contemporary population for M2. As with the pattern established in the BL dimension, both St Simons and Bayshore possess higher fluctuating asymmetry values than the contemporary population for these teeth (Figure 6-3a and 6-3b).

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43 00.020.040.06Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiterescaled mean Mesiodistal 0.020.040.06 mean A B 0Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiterescaled Mesiodistal Figure 6-3. Maxilla site differences (FA) in the mesiodistal dimension. A) PM3. B) M2. Similar to the maxilla, three of the five teeth analyzed in the mandible exhibit site differences. However for this arcade the teeth involved are more anteriorly positioned (i.e., C1, PM3, and PM4). The mandible contains a single case of significant site difference in the BL dimension involving PM4 (Figures 6-4). In this instance, Bayshore has significantly more fluctuating asymmetry than Palmer, St. Simons, Harrison, and the contemporary population. In the mesiodistal dimension; Bayshore has more FA than the contemporary population for C1 (Figure 6-5a), Palmer has more FA than the contemporary population for PM3 (Figure 6-5b), and both Palmer and Bayshore have more FA than the contemporary population for PM4 (Figure 6-5c). 00.020.040.060.08Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiterescaled mean Buccolingual Figure 6-4. Mandible site differences (FA) in the buccolingual dimension. Tooth PM4 represented.

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44 0 0.02 0.04 0.06 0.08Bay W. Tick Is. Palmer Bayshore St. Sim. Harrison Modernsiterescaled mean Mesiodistal 0 0.02 0.04 0.06 0.08Bay W. Tick Is. Palmer Bayshore St. Sim. Harrison Modernsiterescaled mean Mesiodistal 0 0.02 0.04 0.06 0.08Bay W. Tick Is. Palmer Bayshore St. Sim. Harrison Modernsiterescaled mean Mesiodistal Figures 6-5. Mandible site differences (FA) in the mesiodistal dimension. A) C1. B) PM3. C) PM4. Note: Absence of data point indicates no tooth data was available for that site. Sex*Site Interaction Teeth in both the mandible and the maxilla exhibit significant sex*site interactions for fluctuating asymmetry. These interactions involve one or both dimensions of every tooth with the exception of PM3. The maxilla demonstrates sex*site interactions in the buccolingual dimension of M1 and M2. A marked sex difference can be seen at St. Simons for M1. This is denoted by a drastic increase in FA for males and a slight decrease in females at this site, in relation to the other sites (Figure 6-6a). The second molar, on the other hand, exhibits a marked increase in females at Bayshore and St. Simons followed by a steady decline at the subsequent sites. At the same time, males exhibit a slight leveling out followed by a decrease in FA through the sites (Figure 6-6b). A B C

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45 00.0050.010.0150.020.0250.030.035Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiterescaled mean F M B 0.0050.010.0150.020.0250.030.035escaled mean 0Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiter A F M Figure 6-6. Maxilla sex*site interactions (FA) for the molars. A). M1 mesiodistal. B) M2 mesiodistal. The mandible displays significant sex*site interactions in both dimensions of PM4 and the MD dimension of C1. Both dimensions of PM4 exhibit marked differences between males and females at Bayshore (Figure 6-7). Of the two sexes, males display a sudden spike in asymmetry levels for this population. A similar spike for this population is seen in C1. However in this instance, females; as opposed to males, display a sudden elevation in fluctuating asymmetry (Figure 6-8). The other sites for these teeth display a fairly steady level of FA for both sexes (Figures 6-7 and 6-8). 00.050.10.150.2Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiterescaled mean F M 00.050.10.150.2Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiterescaled mean F M B A Figure 6-7.Mandible sex*site interactions (FA) for PM4. A) Mesiodistal dimension. B) Buccolingual dimension.

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46 00.050.10.150.2Bay W.Tick Is.PalmerBayshoreSt. Sim.HarrisonModernsiterescaled mean F M Figure 6-8. Mandible sex*site interactions (FA) for C1 in mesiodistal dimension. Directional Asymmetry Sex Differences As with FA, the maxilla also displays no significant sex differences in relation to DA. However, the MD dimension of PM4 and M2 demonstrates sex differences with males being more left-side dominant and females more right-side dominant for this dimension (Figure 6-9). As a reminder, positive values indicate left side dominance, while negative values indicate right side dominance in the graphs that follow. -0.15-0.1-0.0500.050.1CIPM3PM4M1M2tooth(L-R) Mean Female Male Figure 6-9. Mandible sex differences (DA).

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47 Site Differences Only the mesiodistal dimension of PM3 in the maxilla displays a significant site difference. In this instance, St. Simons is highly left sided in relation to the contemporary population, Palmer, and Bayshore (though not significantly for Bayshore) (Figure 6-10)). In the mandible, both dimensions of PM3 exhibit the site differences. Buccolingually, St. Simons is more right-sided than the contemporary population, while mesiodistally Bayshore and Tick Island are significantly equidistant from the contemporary population in opposing directions. -0.4-0.200.20.4ABCDEFGsite(L-R) Mean Mesiodistal Figure 6-10. Maxilla site differences (DA) for PM3. (A=Bay West B= Tick Island C= Palmer D= Bayshore Homes E= St. Simons F= Harrison G= Modern). -0.4-0.200.20.4ABCDEFGsite(L-R) Mean Buccolingual -0.4-0.200.20.4ABCDEFGsite(L-R) Mean Mesiodistal A B Figure 6-11. Mandible site differences (DA) for PM3. A) Buccolingual dimension. B) Mesiodistal

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48 Sex*Site Interaction The maxilla displays the only instance of significant sex*site interaction for DA. For this arcade, the mesiodistal dimension of M1 shows significance. A graph of this tooth measurement indicates a general trend of reversal in directionality between the sites for males and females, such that when males are left dominant females tend to be right dominant, and vice versa across the sites (Figure 6-12). -0.6-0.4-0.200.20.4ABCDEFGsite(L-R) Mean F M Figure 6-12. Maxilla sex*site interactions (DA). Tooth M1 in mesiodistal dimension. (A=Bay West B=Tick Island C=Palmer D=Bayshore Homes E=St. Simons F=Harrison G=Modern).

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CHAPTER 7 DISCUSSION AND CONCLUSIONS The overall goal of this research was to evaluate fluctuating and directional asymmetry through time in geographically similar populations. While previous examination of temporally distinct populations has been conducted (Suarez 1974; Perzigian 1977), none of these studies utilized geographically similar populations, thus making this research novel. Dental variables in this assessment of asymmetry include mesiodistal (MD) and buccolingual (BL) dimensions for C1 through M2 in both arcades. The ensuing dataset was partitioned into biological variables (sex) and dimensional variables (site as a proxy for time), as well as the combination of these variables. This study addresses the following hypotheses on levels of dental asymmetry in relation to environmental stress through time: Males and females manifest stress in the dentition similarly Asymmetry levels are differentially experienced based on temporal grouping. More specifically, sites representative of European Contact display the greatest levels of asymmetry while modern populations display the least Since sex difference is predicted to be insignificant, the interaction between sex and grouping is therefore negligible Summary of Statistical Results The results presented in the previous chapter can be summarized as follows. A decided lack of sexual dimorphism is evident for FA and DA across the sites, with each having only two of the twenty measurements for this factor display significance. 49

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50 Significant site differences are found in eight of the twenty FA measurements, and three of the twenty DA measurements. St. Simons (Contact Period) has the largest level of asymmetry in half of the significant site difference comparisons. The remaining comparisons find the largest asymmetry levels in Bayshore and/or Palmer (Woodland Period). Meanwhile, the contemporary population consistently displays the lowest levels of asymmetry There are significant sex*site interactions for both types of asymmetry (FA = 5; DA = 4). While not directly addressed in the major hypotheses above, the data also demonstrate more instances of significant differences in FA across sex, site, and sex*site interactions (n=15), than DA (n=6) Additionally, the more distal teeth (premolars and M2) within the tooth fields display the most asymmetry Because of variation in statistical methods between studies and the way in which this dataset is organized, it would be inappropriate to make direct comparisons between the numerical values calculated in this research and those presented in other studies. However, this does not preclude discussion of the patterns seen within this data, in relationship to those found by other researchers. Sex Differences Most population studies for groups as vast as Amerindians, Modern whites, and Australian aboriginals (Perzigian 1977, Townsend 1981; Kieser et al 1986a; 1986b) have found that females and males do not differ significantly in their response to asymmetrogenic factors. This was also found to be the case in this study, with no sex differences occurring in the maxilla. For DA, there are only two instances of sex differences in the mandible that reached significance: PM4 (MD) and M2 (MD). However, it was suspected that this asymmetry between the sexes could in fact be attributed to extreme differences within only a few sites; as opposed to all of the sites. This was confirmed by graphing each sex’s significant mean DA values by site (i.e.,

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51 examining the sex*site interaction). As one can see, most of the sites have asymmetry levels for the sexes which are close to zero (Figure 7-1). However, there are noticeable deviations from this pattern for certain sexes at specific sites, such as females at Harrison for PM4 or both sexes at Bayshore for M2. Thus, as opposed to the presence of a directional trend in the sexes through the sites, particular sites show drastic sex outliers, in relation to the rest of the data. As with DA, there were only two instances of sexual dimorphism for FA. Once again, the graphing of these means across the seven populations reveals elevated sexual dimorphism typically in only one or two sites, as opposed to across sites (Figures 6-6, 6-7, and 6-8). Thus, the dataset as a whole does not show the presence of sexual dimorphism for either directional or fluctuating asymmetry. However, examining the interaction effects in teeth also showing sexual dimorphism suggests some sort of sexual dimorphism may be occurring within particular sites, such as at Bayshore and St. Simons for FA and potentially DA. -0.4-0.200.20.4ABCDEFGsitemean Females Males -0.4-0.200.20.4ABCDEFGsitemean Females Males A B Figure 7-1. Mandible sex*site interactions (DA) for instances of sex difference. A) PM4. B) M2. (A=Bay W, B=Tick Island, C=Palmer, D= Bayshore, E=St. Simons, F=Harrison, G=Modern)

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52 Site Differences Almost half of the measurements for FA show significant site differences (8 out of 20), while only 3 out of 20 are significant for DA. Within these differences there is a marked trend toward the significant site comparisons involving either St. Simons and the contemporary population, or a Woodland Period site (i.e., Bayshore or Palmer) and the contemporary population (Figures 7-2 and 7-3). In these cases, the asymmetry values for St. Simons or the Woodland Period sites are consistently the largest, while the contemporary population always has the smallest value. Occasionally other sites show significant site differences (Figure 7-2a, 7-2c and 7-3b). However in these instances, St. Simons, Bayshore, and Palmer still tend to display the highest level of asymmetry. 00.020.040.060.08PM3PM4toothrescaled mean St Simons Modern Bayshore Palmer Harrison 00.020.040.060.08CIPM3PM4toothrescaled mean Modern Bayshore Palmer 00.020.040.060.08PM4M1M2toothrescaled mean Tick Bayshore St Simons Modern 00.020.040.060.08PM3M2toothrescaled mean Modern Bayshore St. Simons A B C D Figure 7-2. Fluctuating asymmetry bar graphs of site differences. A) Mandible in the buccolingual dimension. B) Mandible in the mesiodistal dimension. C) Maxilla in the buccolingual dimension. D) Maxilla in the mesiodistal dimension.

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53 -0.3-0.2-0.100.10.20.30.4PM3mean St Simons Modern Palmer -0.3-0.2-0.100.10.20.3PM3mean Bayshore Tick -0.3-0.25-0.2-0.15-0.1-0.050PM3mean St Simons Modern A B C Figure 7-3. Directional asymmetry bar graphs of site differences. A). Maxilla in mesiodistal dimension. B) Mandible in mesiodistal dimension. C) Mandible in buccolingual dimension. Furthermore, when examining the site data in totality for FA, the contemporary population displays the lowest level of asymmetry (i.e., values close to zero); in relation to temporally older sites (Figure 7-4 and 7-5). This diminished asymmetry level is not surprising considering greater access to medical care and food for modern individuals, which in turn leads to diminished stress levels. There are, however, exceptions to this pattern, in terms of the mandibular dimensions, (Figure 7-6 and 7-7). In these instances, the Bay West Site (Archaic Period) has a similarly low level of asymmetry for many of the teeth. However, considering the marked improvement in quality of life for modern populations, I suspect these results are not in fact indicative of lower stress levels within the Archaic population. Instead, they most probably can be viewed as statistical artifacts related to Bay West’s small sample size (n 4 for all measurements). In addition, FA displays a pattern whereby St. Simons and/or the Woodland sites have the highest levels of asymmetry. This pattern is in keeping with the overall trend for the teeth, whereby these sites show the greatest levels of FA (Figures 7-4 through 7-7).

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54 00.010.020.030.040.050.060.070.08CIPM3PM4M1M2toothrescaled mean Bay West Tick Island Palmer Bay Shore St Simons Harrison Modern Figure 7-4. Buccolingual maxillary FA across sites. Note: Lines thickened for St. Simons, Palmer, Bayshore, and the contemporary population. 00.010.020.030.040.050.060.070.08CIPM3PM4M1M2toothrescaled mean Bay West Tick Island Palmer Bay Shore St Simons Harrison Modern Figure 7-5. Mesiodistal maxillary FA across sites.

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55 00.010.020.030.040.050.060.070.08CIPM3PM4M1M2toothrescaled mean Bay West Tick Island Palmer Bay Shore St Simons Harrison Modern Figure 7-6. Buccolingual mandibular FA across sites. Note: Lines thickened for St. Simons, Palmer, Bayshore, and the Contemporary sample. 00.010.020.030.040.050.060.070.08CIPM3PM4M1M2toothrescaled mean Bay West Tick Island Palmer Bay Shore St Simons Harrison Modern Figure 7-7. Mesiodistal mandibular FA across sites.

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56 The fact that St. Simons separates out from the other sites on numerous occasions is consistent with the third hypothesis; wherein Contact Period sites display the most asymmetry, due to the severe stress native groups were subjected to via increased maize agriculture and Spanish missionization. It is interesting to note that the other Contact Period site, Harrison Homestead, showed only one instance of significant site difference which manifested as Harrison having less FA than Bayshore (Figure 7-2a). I believe the overall insignificance of the Harrison site, in relation to the other temporally different populations, can be accounted for by several factors. First, Harrison was only occupied for eight years. Thus, the majority of individuals measured at this site reached adulthood elsewhere, possibly outside of the Spanish system. We currently know little about the lifeways of this population prior to Harrison Homestead. Therefore, it is possible that these individuals developed in a fairly stress-free environment. Additionally, given the short occupation time of this site, asymmetry may simply have not had an opportunity to manifest dentally. Furthermore, the death rates at this site were quite severe. Occupied by only a few hundred individuals, Harrison Homestead had a mortality rate of over 30 deaths a year (Saunders 1988). This suggests that stress levels (potentially in terms of diet, labor, disease, etc.) were elevated such that they exceeded the body’s coping ability and were thus often fatal. While stress must be excessive enough to have developmental repercussions (i.e., asymmetry) it also must not be so severe as to become lethal, in order for asymmetry to manifest. Thus, there were almost certainly elevated stress levels at Harrison, evident in the high mortality rates. However, the site’s short occupation precluded the manifestation of asymmetry in these individuals.

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57 While the significantly elevated levels of asymmetry for St. Simons and the diminished levels of the contemporary population were not surprising, the significantly pronounced asymmetry levels from the Woodland sites (i.e., Bayshore and Palmer) were unexpected, and quite intriguing. Unfortunately, the Woodland Period is not as well-documented as other time periods; and descriptions of it are generally limited to characterizations of continuation and refinement of previous cultural motifs (Caldwell 1958; Wauchope 1966; Dragoo 1975; Garrow 1975; White 1988). Because of the informational void for this time period, it is uncertain what phenomena may be accounting for the significantly high asymmetry means found at these sites in relation to the other periods, especially in lieu of the well-documented stress present for Contact Period sites. It would be worthwhile to further pursue what factors may be accounting for these elevated asymmetry levels in the Woodland sites. Sex*Site Interaction All of the sex*site interactions show marked mean asymmetry outliers at St. Simons and Bayshore. Males represent these outliers in three of the five significant FA interactions, with females accounting for the remaining two (Figures 6-7, 6-8, and 6-9). Meanwhile, both males and females exhibit spikes in DA for these sites (Figure 6-12). These results; in conjunction with the significant site differences, indicate that the asymmetry seen at the Bayshore and St. Simons is not only significantly greater than many of the other sites, but also manifests as sexual dimorphism within the sites. As previously mentioned, these significant sex differences are probably not caused by sexual dimorphism across the sites. Instead, they are the result of extreme sex differences within Bayshore and St. Simons skewing the pooled sex data.

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58 Fluctuating Asymmetry vs. Directional Asymmetry Fluctuating asymmetry represents 15 of the 21 cases of statistical significance. Of the remaining six which are DA, it is suspected that the two sex effects are related to differences within particular sites, much like those seen in FA. Hence, this study indicates FA accounts for a greater proportion of the asymmetry within sites and through time. Asymmetry in Tooth Classes Having discussed the possible implications of site and sex variability in the seven populations as related to differences in asymmetry, I now turn to the particular teeth displaying significant asymmetry. Of the teeth displaying significant levels of asymmetry, the premolars consistently show differences within both dimensions (PM3 = 8; PM4 = 5). The next most prevalent tooth type is M2 (n = 4), followed by M1 (n = 2) and a single instance of C1 involvement. The question then becomes, why do premolars appear to show more significant variation in tooth asymmetry than the molars and canines? Variability profiles constructed by Kieser and Groeneveld (1988) for Australian men and Lapp women reveal the greatest variability in crown size for I2, M3 (which were excluded from this study) followed by PM4 in the maxilla. The canines, I1, and M1 represent the least variable teeth in this arcade. While there is less of a consistent pattern, the premolars have often been found to be the most variable in the mandible (Hillson 1996). Scholars have attempted to explain this phenomenon via three different approaches. The first is known as field theory. Postulated by Butler (1939), it suggests this variation may be the result of development gradients existing within the different tooth fields; which he designated as incisor, canine, and molars. Butler presupposed that all tooth

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59 germs possess the same set of instructions. This results in the development of a generalized teeth form. It is, then, the presence of some factor external to the tooth germ that provides them with the instructions to create a particular tooth type. This factor, which he referred to as a field substance or morphogen, issues different instructions to tooth germs at different positions in the jaw, resulting in specific tooth types: incisor, canine, molar and the teeth lying posterior to them (premolars would be in the post-canine field). Each morphogen diffuses in a gradient around this central or polar tooth germ, such that neighboring tooth germs and their resulting tooth types are more strictly controlled by the morphogens; than more distant germs. This is seen in the trend of upper I2 tending to be more variable than I1, and molars increasing in variability from M1 to M3 (Hillson 1996). Hence, in the canine field, C1 would be the least variable and PM4 would be the most variable when subjected to equal levels of external stress. Kollar and Baird (1971) demonstrated that the surrounding mesenchyme is responsible for issuing instructions for tooth type. Thus, properties within the mesenchyme may be serving as these so-called morphogens. Meanwhile, Osborn (1978) explained this pattern of tooth variability using the clone theory. He suggested that the dental papilla, cells which eventually form the dentin and pulp in teeth, develop by division from three different populations (clones) of mesenchyme cells. These clones, in turn, are associated with specific classes of teeth. The anterior clone produces the deciduous and permanent incisors. The canine clone produces the deciduous and permanent canine, and finally the posterior clone produces the deciduous molars and permanent premolars and molars. Growth of a tooth germ is controlled from within (i.e., no morphogens); whereby each clone starts from a stem

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60 precursor. This precursor is cloned and the resulting clone is then copied. This pattern of cloning continues in both the anterior and posterior directions; till all the teeth within the particular clone type are formed. Because of this, successive teeth in a series undergo more cellular divisions, leading to increased accumulation of variability due to diminished mitotic control (Osborn 1978). For the molar clone, then, the most anterior premolar (PM3) and the most posterior molar (M3) have the most asymmetry in response to environmental stress. Finally, Kieser and Groenveld (1988) suggest the relatively longer period of time the more posterior teeth (premolars, M2, and M3) spend in the soft tissue stage of crown formation may leave them more susceptible to environmental disturbances. This has been noted particularly in the deciduous dentition (Mizoguchi 1980; 1983; Townsend and Farmer 1993). The common theme in all these theories is that the potential for stability to be compromised is elevated in the more distal portions of the tooth fields; thus making these teeth (i.e., premolars, M2, M3) particularly susceptible to external stressors. The results of this inquiry follow this pattern, whereby the majority of significant site differences occur in the upper and lower premolars. The involvement of the more stable teeth (lower C1 and upper M1) along with the premolars and M2 in St. Simons and Woodland sites further indicates the drastic environmental stress which must have been occurring at these locales. It should be noted that despite the apparent patterns in the results, the small sample size could be responsible for creating false positives or potentially masking instances of significance in other teeth, thereby compromising the validity of some of these

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61 interpretations. Similarly, the influence of some unknown external variable or the interaction between factors such as preservation, curation, and attrition rates may also be skewing the results. However, regardless of these possible limitations, the results do display several patterns which have been found in other populations. Further Research An interesting line of further inquiry would be to collect data on other oral dimensions, such as arcade length and width from these populations. This data could then be used to analyze the relationship between tooth asymmetry and arcade dimensions. As research suggests the size of the jaw serves as a contributing factor in patterns of relative molar calcification (Kieser and Groenveld 1988), it would also be interesting to uncover what role, if any, such dimensions play in levels of dental asymmetry. In addition, it might be worthwhile to evaluate other indicators of environmental stress, such as enamel hypoplasias and cribra orbitalia within the samples to reveal a broader picture of how these populations manifest stress. In that same vein, this data could also be used in conjunction with other faunal (Kozuch 1998), flora (Newsom 1998) or isotope analyses (Cabinera 1999) that have been conducted on these collections to create a representative view of health through time. Finally, it should be emphasized that different avenues of analysis could be performed on this data set. While beyond the scope of this study, this is something I hope to pursue further in the future. Conclusions The primary goal of my project was to identify trends of elevated stress in populations from different time periods. This was accomplished by examining dental asymmetry. Dental asymmetry is believed to be the result of developmental instability

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62 due to environmental, as opposed to genetic, stress (Garn et al. 1965; Kollar and Baird 1971; Harris and Nweeia 1980; Mizoguchi 1980; Mizoguchi 1982; Mayhall and Saunders 1986; Livshits and Kobyliansky 1989; Kieser 1990; Mayhall 1992; Hillson 1996). Thus, an evaluation of this phenomenon can measure the degree of environmental stress experienced by a population. The populations I studied originated from coastal Florida and Georgia. By constraining physical surroundings, I was able to exclude this component of environmental stress, thereby allowing me to attribute population variation in dental asymmetry to other environmental factors, such as the way different cultures exploit their surroundings. An inter-site research design identified significantly elevated levels of dental asymmetry at an early Contact Period site (St. Simons: 1450 to1550 A.D.) and the two Woodland Period sites (Palmer and Bayshore Homes: 250 to 1000 A.D.). Meanwhile, the contemporary population displayed the lowest level of asymmetry, in relation to the other populations. Overall, sexual dimorphism was also found to be minor across and within the populations. These results partially support my research hypothesis, which proposed the most dental asymmetry in the Contact Period populations and the least in the modern population. However, the elevated levels among the Woodland populations were unexpected. Yet with further research, these results may serve as a valuable glimpse into the living conditions of people during this time period. Of the populations examined, only Bayshore Homes and St. Simons displayed any instances of sexual dimorphism in dental asymmetry. These cases also produced the few significant interactions between sex and site in the sample. While this may be the result of statistical error (i.e., due to small sample size) these sex differences may speak to some

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63 unique environmental phenomenon occurring at these locales. Finally, significant differences in asymmetry were found primarily in the more distal teeth in the tooth fields (premolars and posterior molars). This is consistent with research (Butler 1939; Osborn 1978; Kieser and Groenveld 1988) indicating these teeth are more likely to display diminished ontogenetic control and increased susceptibility to environmental stress. In conclusion, the results of my research indicate that dental asymmetry, and by extension environmental stress, in fact, varies through time. Hence, dental asymmetry can reveal patterns of stress related to the living conditions associated with a particular time period, as well as trends through time. Similarly, dental asymmetry studies can allow one to step away from the realm of temporal generalizations and display asymmetry patterns unique to the life history of a particular population. For instance, this sort of analysis can reveal undocumented periods of severe stress which are isolated to a single population, resulting from factors such as illness or crop destruction. Thus, my research suggests analysis of dental asymmetry may provide a viable measure of nutritional, socio-economic, and bio-developmental differences in human populations across time.

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APPENDIX A MAXILLA VALUES FOR FLUCTUATING ASYMMETRY Table A-1. Female maxilla data Measurement Mean # of tooth pairs Std Dev bC1 0.019 33 0.019 bPM3 0.017 39 0.017 bPM4 0.021 40 0.019 bM1 0.010 36 0.010 bM2 0.013 31 0.017 mC1 0.035 33 0.046 mPM3 0.039 40 0.042 mPM4 0.049 41 0.070 mM1 0.020 36 0.019 mM2 0.026 31 0.030 Table A-2. Male maxilla data Measurement Mean # of tooth pairs Std Dev bC1 0.016 39 0.015 bPM3 0.016 42 0.025 bPM4 0.020 47 0.018 bM1 0.011 48 0.015 bM2 0.010 46 0.010 mC1 0.031 39 0.031 mPM3 0.025 45 0.024 mPM4 0.034 48 0.030 mM1 0.024 47 0.033 mM2 0.021 46 0.030 64

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65 Table A-3. Bay West maxilla data Measurement Mean # of tooth pairs Std Dev bC1 0.025 2 0.011 bPM3 0.014 2 0.015 dPM4 0.017 3 0.014 bM1 0.007 3 0.001 bM2 0.015 3 0.013 mC1 0.029 2 0.001 mPM3 0.028 2 0.013 mPM4 0.032 2 0.018 mM1 0.020 3 0.005 mM2 0.031 3 0.040 Table A-4. Tick Island maxilla data Measurement Mean # of tooth pairs Std Dev bC1 0.019 6 0.015 bPM3 0.017 9 0.016 dPM4 0.010 11 0.011 bM1 0.009 10 0.010 bM2 0.008 7 0.010 mC1 0.031 6 0.022 mPM3 0.024 10 0.021 mPM4 0.045 11 0.042 mM1 0.026 10 0.023 mM2 0.020 7 0.016 Table A-5. Palmer maxilla data Measurement Mean # of tooth pairs Std Dev bC1 0.022 19 0.020 bPM3 0.025 19 0.034 dPM4 0.021 19 0.015 bM1 0.011 19 0.011 bM2 0.011 18 0.010 mC1 0.032 19 0.036 mPM3 0.039 19 0.041 mPM4 0.032 20 0.029 mM1 0.022 19 0.017 mM2 0.024 18 0.023

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66 Table A-6. Bayshore maxilla data Measurement Mean # of tooth pairs Std Dev bC1 0.025 8 0.019 bPM3 0.014 11 0.008 dPM4 0.021 11 0.016 bM1 0.008 13 0.008 bM2 0.020 10 0.018 mC1 0.056 9 0.060 mPM3 0.041 11 0.047 mPM4 0.068 12 0.081 mM1 0.024 13 0.021 mM2 0.039 10 0.038 Table A-7. St. Simons maxilla data Measurement Mean # of tooth pairs Std Dev bC1 0.017 16 0.019 bPM3 0.021 14 0.024 dPM4 0.032 17 0.027 bM1 0.020 14 0.023 bM2 0.020 15 0.017 mC1 0.044 15 0.046 mPM3 0.052 16 0.037 mPM4 0.053 18 0.038 mM1 0.034 4 0.054 mM2 0.029 15 0.040 Table A-8. Harrison maxilla data Measurement Mean # of tooth pairs Std Dev bC1 0.021 4 0.016 bPM3 0.012 9 0.010 dPM4 0.020 10 0.019 bM1 0.015 7 0.013 bM2 0.011 4 0.013 mC1 0.028 4 0.030 mPM3 0.023 9 0.013 mPM4 0.051 10 0.096 mM1 0.017 6 0.019 mM2 0.042 4 0.049

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67 Table A-9. Contemporary population maxilla data Measurement Mean # of tooth pairs Std Dev bC1 0.008 17 0.008 bPM3 0.008 17 0.008 dPM4 0.013 16 0.009 bM1 0.005 18 0.006 bM2 0.002 20 0.003 mC1 0.013 17 0.014 mPM3 0.010 18 0.012 mPM4 0.009 16 0.011 mM1 0.011 18 0.015 mM2 0.005 20 0.007

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APPENDIX B MANDIBLE VALUES FOR FLUCTUATING ASYMMETRY Table B-1. Female mandible data Measurement Mean # of tooth pairs Std Dev bC1 0.012 27 0.011 bPM3 0.024 38 0.019 bPM4 0.024 34 0.023 bM1 0.013 33 0.013 bM2 0.015 33 0.015 mC1 0.029 27 0.042 mPM3 0.028 38 0.022 mPM4 0.025 35 0.028 mM1 0.024 33 0.026 mM2 0.017 31 0.017 Table B-2. Male mandible data Measurement Mean # of tooth pairs Std Dev bC1 0.019 40 0.031 bPM3 0.021 42 0.028 bPM4 0.028 48 0.044 bM1 0.011 44 0.012 bM2 0.024 43 mC1 0.023 39 0.023 mPM3 0.030 44 0.043 mPM4 0.040 48 0.052 mM1 0.020 45 0.017 mM2 0.022 45 0.025 0.059 Table B-3. Bay West mandible data Measurement Mean # of tooth pairs Std Dev C1 . 4 . bPM3 0.019 1 . bPM4 0.006 1 . bM1 0.006 4 0.0068703 bM2 0.008 4 0.0021304 mC1 . 0 . mPM3 0.058 1 . mPM4 0.002 1 . mM1 0.011 4 0.0089918 mM2 0.030 4 0.0219086 68

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69 Table B-4. Tick Island mandible data Measurement Mean # of tooth pairs Std Dev bC1 0.026 11 0.055 bPM3 0.023 15 0.025 dPM4 0.025 17 0.032 bM1 0.012 21 0.014 bM2 0.035 20 0.085 mC1 0.035 11 0.022 mPM3 0.049 15 0.059 mPM4 0.044 17 0.060 mM1 0.025 21 0.019 mM2 0.022 20 0.018 Table B-5. Palmer mandible data Measurement Mean # of tooth pairs Std Dev bC1 0.020 18 0.012 bPM3 0.035 17 0.036 dPM4 0.023 19 0.020 bM1 0.014 17 0.013 bM2 0.018 15 0.015 mC1 0.032 17 0.038 mPM3 0.024 18 0.017 mPM4 0.038 19 0.037 mM1 0.026 17 0.020 mM2 0.025 15 0.027 Table B-6. Bayshore mandible data Measurement Mean # of tooth pairs Std Dev bC1 0.031 5 0.020 bPM3 0.025 6 0.021 bPM4 0.031 7 0.024 bM1 0.014 8 0.012 bM2 0.014 9 0.017 mC1 0.026 5 0.017 mPM3 0.047 6 0.030 mPM4 0.042 7 0.028 mM1 0.029 8 0.020 mM2 0.027 9 0.038

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70 Table B-7. St. Simons mandible data Measurement Mean # of tooth pairs Std Dev bC1 0.010 9 0.006 bPM3 0.020 9 0.015 bPM4 0.070 7 0.093 bM1 0.019 7 0.011 bM2 0.022 6 0.016 mC1 0.050 9 0.054 mPM3 0.037 9 0.043 mPM4 0.068 7 0.076 mM1 0.032 7 0.037 mM2 0.020 6 0.014 Table B-8. Harrison mandible data Measurement Mean # of tooth pairs Std Dev bC1 0 12 0 bPM3 0.021 10 0.018 bPM4 0.025 11 0.022 bM1 0.011 5 0.005 bM2 0.014 7 0.010 mC1 . 12 0.000 mPM3 0.032 11 0.021 mPM4 0.041 11 0.028 mM1 0.018 6 0.026 mM2 0.013 8 0.008 Table B-9. Contemporary population mandible data Measurement Mean # of tooth pairs Std Dev bC1 0.008 24 0.010 bPM3 0.012 22 0.013 dPM4 0.014 20 0.022 bM1 0.007 15 0.012 bM2 0.012 15 0.017 mC1 0.008 24 0.006 mPM3 0.009 22 0.012 mPM4 0.004 21 0.005 mM1 0.010 15 0.008 mM2 0.006 14 0.008

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73 Harris, E.F. 1992 “Laterality in human odontometrics: analysis of a contemporary American White series” ” in Culture, Ecology and Dental Anthropology, J Lukacs (ed.) Kamla Raj, Delhi, p. 157-170. Harris, E.F. and M. Nweeia 1980 “Dental Asymmetry as a Measure of Environmental Stress in the Ticuna Indians of Colombia.” American Journal of Physical Anthropology. Vol. 53, p. 133-142. Hillson, S. 1996 Dental Anthropology. Cambridge University Press, Cambridge. Hutchinson, D.L., Larsen C.S., Schoeninger M.J., and L. Norr 1998 “Regional Variation in the Pattern of Maize Adoption and Use in Florida and Georgia.” American Antiquity, Vol. 63, No. 3, p. 397-416. Jahn, O.F. and R.P. Bullen 1978 “The Tick Island Site, St. Johns River, Florida.” Florida Anthropological Society Publications. Johanson D. and E. Blake 1996 From Lucy to Language. New York: New York. Kieser, J.A. 1990 Human adult odontometrics: The study of variation in adult toothsize. Cambridge University Press, Cambridge. Kieser, J.A. and H. T. Groeneveld 1988 “Fluctuating Odontometric Asymmetry in an Urban South African Black Population.” Journal of Dental Research, Vol. 67, p. 1200-1205. 1994 “Effects of Prenatal Exposure to Tobacco Smoke and Developmental Stability in Children.” Developmental Biology, Vol. 14, p. 43-47. Kieser, J.A., Groeneveld, H.T. and C.B. Preston 1986a. “Fluctuating Odontometric Asymmetry in the Lengua Indians of Paraguay. Annals of Human Biology, Vol. 13, p.489-498. 1986b. “Fluctuating Dental Asymmetry as a Measure of Odontogenic Canalization in Man.” American Journal of Physical Anthropology, Vol. 71, p. 437-444. Kollar, E.J. and G.R.Baird 1971 “Tissue Interactions in Developing Mouse Tooth Germs” in Dental Morphology and Evolution, A.A. Dahlberg (ed.) University of Chicago Press, Chicago, pp. 15-29

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74 Kozuch, L. 1998 “Faunal Remains from the Palmer Site (8SO2), with a focus on shark remains.” The Florida Anthropologist. Vol 51, No. 4, p. 208-222. Larsen, C.S. 1995 “Biological Changes in Human Populations with Agriculture.” Annual Review of Anthropology, Vol. 24, p. 185-213. Livshits, G. and E. Kobyliansky 1989 “Study of Genetic Variance in Fluctuating Asymmetry of Anthropometrical Traits.” Annals of Human Biology, Vol. 16, p. 121-129. 1991 “Fluctuating Asymmetry as a Possible Measure of Developmental Homeostasis in Humans: a Review. Human Biology, Vol. 63, No. 4, p. 441-466. Mayhall, J. 1992 “Techniques for the study of dental morphology” in Skeletal Biology of Past Peoples: Research Methods, S.R. Saunders and M.A. Katzenberg, (eds.), Wiley-Liss, New York. Mayhall, J.T., and S.R. Saunders 1986 “Dimensional and discrete dental trait asymmetry relationships.” American Journal of Physical Anthropology, Vol. 69, p. 403-411. McGoun, W.E. 1993 Prehistoric Peoples of South Florida. The University of Alabama Press,Tuscaloosa. Milanich, J.T. 1995 Florida Indians and the Invasion from Europe. University Press of Florida,Gainesville. 1998 Florida’s Indians from Ancient Times to the Present. University Press of Florida,Gainesville. Milanich, J.T., and C.H. Fairbanks 1978 Florida Archaeology. Academic Press, New York. Mizoguchi, Y. 1980 “Factor Analysis of Environmental Variation in the Permanent Dentition.” Bulletin of the National Science Museum Tokyo, Vol. 6, p. 29-46. 1983 “Influences of the Earlier Developing Teeth upon the Later Developing Teeth. Bulletin of the National Science Museum Tokyo, Vol.9, p.33-45.

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75 1986 “Correlated Asymmetries Detected: The Tooth Crown Diameters of Human Permanent Teeth.” Bulletin of the National Science Museum Tokyo, Vol. D 12, p.15-45. Molnar, S. 1972 “Tooth Wear and Culture: A Survey of Tooth Functions among Some Prehistoric Populations.” Current Anthropology, Vol. 13, No. 5, p.511-526. Newsom, L.A. 1998 “Archaeobotanical Research at Shell Ridge Midden, Palmer Site (8SO2), Sarasota County, Florida.” The Florida Anthropologist,. Vol. 51, No. 4, p. 208-222. Nichol, D.R., Turner, C.G., and T.R. Gest 1984 “Variation in the Convexity of the Human Maxillary Incisor Surface.” American Journal of Physical Anthropology, Vol. 63, p. 361-370. Osborn, J.W. 1978 “Morphologenetic Gradients: Fields Versus Clones” in Development, Function and Evolution of Teeth, P.M Butler and K.A. Joysey (eds), Academic Press, London, p. 171-199. Palmer, A.R. 1994 “Fluctuating Asymmetry Analyses: A primer” in Developmental Instability: Its Origins and Evolutionary Implications. T.A. Markow (ed), Kluwer, Dordrecht, Netherlands. Parsons, P.A. 1990 “Fluctuating Asymmetry: an Epigenetic Measure of Stress.” Biology Review, Vol. 65, p. 131-145. Pearson, C, and F.C. Cook 2003 “Clarence Bloomfield Moore’s Unpublished Excavations on St. Simons Island, Georgia: 1898.” Early Georgia. Vol. 31, No.1, p.25-39. Perzigian, A.J. 1977 Fluctuating Dental Asymmetry: Variation among Skeletal Populations. American Journal of Physical Anthropology, Vol. 47, p. 81-88. Potter, R.H.Y. and W.E. Nance 1976 “A Twin Study of Dental Dimensions: Descordance, Asymmetry and Mirror Imagery. American Journal of Physical Anthropology. Vol. 44, p. 391-6.

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BIOGRAPHICAL SKETCH In 2001 Shanna E. Williams received her Bachelor of Arts degree in anthropology and biology from The University of Virginia, Charlottesville, Virginia. That year, she was accepted into the anthropology graduate program at the University of Florida, where she plans to stay in pursuit of a doctoral degree. 78