The Origins of Dental Crowding in the Florida Archaic: an Anthropological Investigation of Malocclusions in Windover Pond (8BR246) Kathryn O’Donnell Miyar

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2012 The Origins of Dental Crowding in the Florida Archaic: An Anthropological Investigation of Malocclusions in Windover Pond (8BR246) Kathryn O’Donnell Miyar

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COLLEGE OF ARTS AND SCIENCES

THE ORIGINS OF DENTAL CROWDING IN THE FLORIDA ARCHAIC:

AN ANTHROPOLOGICAL INVESTIGATION OF MALOCCLUSIONS IN WINDOVER

POND (8BR246)

KATHRYN O’DONNELL MIYAR

A Dissertation submitted to the Department of Anthropology in partial fulfillment of the requirements for the degree of Doctor of Philosophy

Degree Awarded: Spring Semester, 2012 Kathryn Miyar defended this dissertation on February 29, 2012.

The members of the supervisory committee were:

Lynne Schepartz

Professor Directing Dissertation

Dennis Slice

University Representative

Glen Doran Committee Member

Rochelle Marrinan Committee Member

The Graduate School has verified and approved the above-named committee members, and certifies that the dissertation has been approved in accordance with university requirements.

I dedicate this disquisition to my ever-patient and supportive Eduardo

iii

ACKNOWLEDGEMENTS

This feat could not have been accomplished without the guidance and support of many people and I am greatly indebted to their assistance. Firstly, I would like to acknowledge my advisor and committee chairperson, Dr. Lynne Schepartz, for taking me under her wing, helping ameliorate my writing skills, and guiding me toward professionalism. I would particularly like to thank Dr. Rochelle Marrinan for being my mentor and a true champion of the anthropology graduate students during these last few tumultuous years. To Dr. Glen Doran for his insight and granting me endless access to one of the greatest skeletal collections in the world. To my outside committee member, Dr. Dennis Slice, for his patient statistical guidance and for opening new doors for me in collaborative interdisciplinary research. I would also like to thank Dr. Margo Schwadron for helping me transition into my career and for her endless support and understanding during this last very trying year.

My fellow graduate students and colleagues at the FSU anthropology department fostered an inspirational and supportive community that has helped produce some exciting research from our tiny department. I am very grateful to all of them for their insights and good company at the bar. In particular I would like to thank Alexandra Parsons for her amazing critical eye, unending listening skills, and for ultimately helping me retain my sanity. I thank Sarah Liko very much for all her encouragement. These last two years would have been difficult to manage without her ear and assurance. I am immensely grateful to Ian Pawn and Geoffrey Thomas for their guidance with my statistical analyses and helping me polish my graphs and tables. I would like to thank Timothy Parsons for his time and unsurpassable map-making skills. Though not affiliated with FSU and thousands of miles away, I could not have made it through graduate school without the support of my friend, colleague, and co-founder, Lynn Lucas.

I credit Dr. Linda Taylor for first opening my eyes to this field during her Introduction to Physical Anthropology class and for knowing I would be an anthropologist before I did. I thank her for helping make that happen. I would also like to thank my master’s advisor Dr. Clifford Brown for believing in me and making this career a reality. Without my research assistant, Jayce iv Hill, this project could not have been completed by the imposed deadline. I am very grateful for his time and efforts. I would also like to thank Joe and Maranda Kles for sharing their home with me during my days of research at the FMNH.

I would like to acknowledge the people and institutions that made this research possible: Eisele Dissertation Foundation, FSU Dissertation Research Foundation, Florida Museum of Natural History, Florida Atlantic University, University of Miami, Florida Gulf Coast University, Monica Faraldo, Dr. Arlene Fradkin, Dr. Heather Walsh-Haney, Dr. Neil Wallis, Donna Ruhl, and Dr. Dave Dickel.

The support and love from my family and friends has been remarkable. I want to thank all of you, particularly my parents, for your patience and encouragement these many years. I promise I will have a “real job” now.

Lastly, I would like to acknowledge my husband, to whom this dissertation is dedicated, who has been the iron backbone throughout my entire graduate career and who has been the most amazing friend throughout my entire life. I would never have made it to this point without him.

TABLE OF CONTENTS

List of Tables ...... x List of Figures ...... xiv Abstract ...... xviii

1. INTRODUCTION ...... 1 Hypothesis ...... 3 Chapter Overview ...... 6

2. DENTAL CROWDING ...... 7 Growth and Developmental Factors Affecting Dental Crowding ...... 8 Deciduous and Permanent Teeth and the Formation of Occlusion...... 8 Mesial Drift ...... 9 Lingual Tipping ...... 10 Third Molar Eruption ...... 12 Continuous Eruption ...... 13 Genetic Factors...... 13 Studies of Heredity ...... 14 Dental Size and Morphology ...... 14 Arch Dimensions ...... 16 Angle Malocclusion Classes ...... 17 Dental Traits Affecting Malocclusion ...... 17 Individual Rotations and Displacements ...... 18 Supernumerary Teeth, Impaction, and Agenesis ...... 18 Environmental Etiology...... 20 Masticatory Function Hypothesis ...... 21 Disuse Theory ...... 23 Dental Wear and Attritional Occlusion...... 24

3. MATERIALS AND METHODS ...... 26 Primary and Comparative Skeletal Collections ...... 26 Windover (8BR246)...... 29 Warm Mineral Springs (8SO19) ...... 30 Little Salt Springs (8SO18) ...... 32 Republic Groves (8HR4) ...... 33 Harris Creek at Tick Island (8VO24)...... 35 Bay West (8CR200) ...... 37 Gauthier (8BR193)...... 38 Methods ...... 40 vi Age Estimation...... 40 Age Cohorts ...... 41 Sex Estimation ...... 41 Dental Crowding ...... 42 Little’s (1975) Irregularity Index ...... 44 Tooth and Arch Dimensions in Relation to Dental Crowding ...... 45 Dental Metrics ...... 45 Arch Metrics ...... 46 Arch Depth and Arch Width ...... 46 Investigation of Environmental Factors Relating to Dental Crowding ...... 47 Attrition ...... 47 Dental Wear Patterning ...... 49 Cranial Metrics...... 49 Investigation of Genetic Relatedness Using Nonmetric Traits ...... 51 Dental Nonmetric Traits ...... 51 Cranial Nonmetric Traits ...... 52 Statistical Analyses...... 54

4. WINDOVER ANALYSES ...... 56 Dental Crowding Analyses ...... 56 Dental Crowding Frequencies by Age Cohorts ...... 56 Onset of Dental Crowding ...... 59 Dental Crowding Frequencies Between Sex Cohorts ...... 60 Spatial Distribution of Crowding at Windover ...... 61 Spatial Distribution of Windover Dental Crowding Severity Ranks ...... 62 Windover Tooth Size and Arch Size Relationships to Dental Crowding ...... 67 Windover Tooth Size Analyses ...... 68 Windover MD Metrics Between Sex Cohorts...... 68 Windover Maxillary MD Metrics Between Occlusion Cohorts ...... 72 Windover Mandibular MD Metrics Between Occlusion Cohorts ...... 75 Windover Maxillary MD Metrics Between Crowding Severity Ranks ...... 77 Windover Crowding Severity Mandibular MD Metrics ...... 79 Windover Arch Depth Analyses ...... 82 Windover Arch Depth Comparisons Between Sexes ...... 82 Windover Arch Depth Comparisons Between Sex and Occlusion Cohorts ...... 83 Windover Arch Width Analyses ...... 85 Windover Arch Width Comparisons Between Sex Cohorts ...... 85 Windover Arch Width Comparisons Between Sex and Occlusion Cohorts ...... 86 Windover Cranial Metrics Analyses ...... 89 Windover Cranial Metrics Comparisons Between Sex Cohorts ...... 89 Windover Cranial Metrics Comparisons Between Sex and Occlusion Cohorts ...... 93 vii Windover Attrition Analyses ...... 94 Windover Attrition Mean Comparisons Between Age and Sex Cohorts ...... 96 Windover Attrition Mean Comparisons Between Age and Occlusion Cohorts ...... 98 Windover Non-Masticatory Dental Wear Patterns ...... 99 Interproximal Grooved Dental Wear Patterns ...... 100 Lingual Root Wear Dental Pattern ...... 102 Incisor Lingual Wear Pattern ...... 105

5. COMPARISONS OF WINDOVER TO OTHER FLORIDA ARCHAIC SAMPLES ...... 107 Dental Crowding Frequencies ...... 107 Dental Nonmetric Analyses ...... 109 Cranial Nonmetric Analyses ...... 116 Tooth Size and Arch Size Analyses ...... 119 Tooth Size Analyses ...... 120 Maxillary MD Metrics Between Occlusion Cohorts ...... 120 Mandibular MD Metrics Between Occlusion Cohorts ...... 122 Arch Depth Analyses ...... 124 Maxillary Arch Depth ...... 124 Mandibular Arch Depth ...... 125 Arch Width Analyses ...... 127 Maxillary Arch Width Analyses ...... 127 Mandibular Arch Width Analyses ...... 128 Cranial Metrics Analyses ...... 130 Dental Attrition Analyses ...... 136

6. DISCUSSION ...... 139 Dental Crowding Within Windover ...... 139 Onset of Dental Crowding in the Florida Archaic ...... 141 Dental Crowding Frequencies Between Age Cohorts ...... 142 Dental Crowding Frequencies Between Sex Cohorts ...... 144 Windover Cluster Patterns for Dental crowding and Dental Wear Patterns ...... 144 Dental Crowding Between Windover and the Comparative Samples ...... 146 Nonmetric Analyses ...... 150 Dental Metrics ...... 152 Arch Shape ...... 153 Cranial Shape ...... 154 Attrition Analyses ...... 156 Summary ...... 157

7. CONCLUSION ...... 161 viii Methodology and Future Research ...... 161 Research Results ...... 162 Summary ...... 165

APPENDICES ...... 167 Appendix I Dental Nonmetric Frequencies ...... 168 Appendix II Cranial Nonmetric Frequencies ...... 172

REFERENCES CITED ...... 176

BIOGRAPHICAL SKETCH ...... 197

LIST OF TABLES

Table 3.1 Skeletal Samples ...... 27

Table 3.2 Age Cohorts ...... 41

Table 3.3 Sex Cohorts ...... 42

Table 3.4 Dental Terminology and Definitions ...... 43

Table 3.5 LII Scores ...... 44

Table 3.6 Craniometric (CM) Abbreviations ...... 50

Table 3.7 Dental Nonmetric Traits and Presence Ranges ...... 52

Table 3.8 Cranial Nonmetric Traits and Their Presence Ranges ...... 54

Table 4.1 Windover Dental Crowding Within Age Cohorts ...... 57

Table 4.2 Windover Dental Crowding Between Sex Cohorts ...... 60

Table 4.3 Windover Dental Crowding Between East and West Portions in NE Pond Subsection ...... 63

Table 4.4 Windover Demographics for East and West Portions in the NE Pond Subsection ...... 65

Table 4.5 Windover Maxillary MD Descriptive Statistics by Sex Cohorts ...... 71

Table 4.6 Windover Mandibular MD Descriptive Statistics by Sex Cohorts ...... 71

Table 4.7 Windover Maxillary MD Metrics: Mann-Whitney U Test Results Between Sex Comparisons ...... 72

Table 4.8 Windover Mandibular MD Metrics: Mann-Whitney U Test Results Between Sex Comparisons ...... 72

Table 4.9 Windover Maxillary MD Descriptive Statistics of Permanent Anterior Teeth with Good Occlusion ...... 74

Table 4.10 Windover Maxillary MD Descriptive Statistics of Permanent Anterior Teeth with Dental Crowding ...... 74

Table 4.11 Windover Maxillary MD Metrics: Mann-Whitney U Test Results Between Occlusion Cohorts ...... 74

Table 4.12 Windover Mandibular MD Descriptive Statistics of Permanent Teeth with Good Occlusion ...... 76

Table 4.13 Windover Mandibular MD Descriptive Statistics of Permanent Teeth with Dental Crowding...... 76

Table 4.14 Windover Mandibular MD Metrics: Mann-Whitney U Test Results Between Occlusion Cohorts ...... 77

Table 4.15 Windover Maxillary MD Descriptive Statistics of Permanent Teeth with Mild/Moderate Crowding ...... 79

Table 4.16 Windover Maxillary MD Descriptive Statistics of Permanent Teeth with Severe Crowding...... 79

Table 4.17 Windover Mandibular MD Descriptive Statistics of Permanent Teeth with Mild/Moderate Crowding ...... 81

Table 4.18 Windover Mandibular MD Descriptive Statistics of Permanent Teeth with Severe Crowding...... 82

Table 4.19 Windover Arch Depth Means by Sex Cohorts ...... 83

Table 4.20 Windover Female Arch Depth in Groups with Good Occlusion and Dental Crowding...... 84

Table 4.21 Windover Males Arch Depth in Groups with Good Occlusion and Dental Crowding...... 85

Table 4.22 Windover Arch Width Descriptive Statistics by Sex Cohort ...... 86

Table 4.23 Windover Female Arch Width in Groups with Good Occlusion and Dental Crowding...... 87

Table 4.24 Windover Male Arch Width in Groups with Good Occlusion and Dental Crowding...... 88

Table 4.25 Windover Cranial Metrics: Between Sex Cohorts ...... 90

Table 4.26 Windover Cranial Metrics: Mann-Whitney U Results Between Sex Comparisons ...... 91

Table 4.27 Windover Cranial Metrics: Mann-Whitney U Test Results Between Occlusion Cohorts in Females and Males ...... 94

Table 4.28 Windover Maxillary Attrition Scores: Descriptive Statistics for Age Cohorts ...... 95

Table 4.29 Windover Mandibular Attrition Scores: Descriptive Statistics for Age Cohorts ...... 95

Table 4.30 Windover Maxillary Attrition Scores: Descriptive Statistics for Age cohorts by Sex ...... 97

Table 4.31 Windover Mandibular Attrition Scores: Descriptive Statistics for Age Cohorts by Sex ...... 97

Table 4.32 Windover Attrition Scores: Mann-Whitney U Test Results for Age Cohorts by Sex ...... 97

Table 4.33 Windover Maxillary Attrition Scores: Descriptive Statistics for Age and Occlusion Cohorts ...... 99

Table 4.34 Windover Mandibular Attrition Scores: Descriptive Statics for Age and Occlusion Cohorts ...... 99

Table 5.1 LII Dental Crowding Comparison of Samples ...... 109

Table 5.2 Frequencies of Maxillary Second Molar Enamel Extensions ...... 110

Table 5.3 Nonmetric Frequencies ...... 111

Table 5.4 Dental Nonmetric Frequencies ...... 114

Table 5.5 Cranial Nonmetric Frequencies ...... 118

Table 5.6 Cranial Nonmetric Frequencies Continued ...... 119

Table 5.7 Aggregate Sample Maxillary MD Descriptive Statistics of Permanent Teeth with Good Occlusion ...... 122

Table 5.8 Aggregate Sample Maxillary MD Descriptive Statistics of Permanent Teeth with Dental Crowding ...... 122

Table 5.9 Aggregate Sample Mandibular MD Descriptive Statistics of Permanent Teeth with Good Occlusion ...... 124

xii Table 5.10 Aggregate Sample Mandibular MD Descriptive Statistics of Permanent Teeth with Dental Crowding ...... 124

Table 5.11 Maxillary Arch Depth Descriptive Statistics Between Windover and Comparative Samples ...... 125

Table 5.12 Mandibular Arch Depth Descriptive Statistics Between Windover and Comparative Samples ...... 127

Table 5.13 Maxillary Arch Width Descriptive Statistics for Windover and Comparative Samples ...... 128

Table 5.14 Mandibular Arch Width Descriptive Statistics for Windover and Comparative Samples ...... 130

Table 5.15 Cranial Metrics: Descriptive Statistics Between Comparative Samples ...... 131

Table 5.16 Cranial Metric Significant Differences Between Samples ...... 131

Table 5.17 Cranial Metrics: Mann-Whitney U Test Results Between Windover and Republic Groves ...... 132

Table 5.18 Cranial Metrics: Mann-Whitney U Test Results Between Windover and Gauthier...... 134

Table 5.19 Cranial Metrics: Mann-Whitney U Test Results Between Windover and Bay West ...... 136

Table 5.20 Young Adult Maxillary Attrition Descriptive Statistics Between Windover and Comparative Samples ...... 137

Table 5.21 Middle-Aged Adult Maxillary Attrition Descriptive Statistics Between Windover and Comparative Samples ...... 137

Table 5.22 Older Adult Maxillary Attrition Descriptive Statistics Between Windover and Comparative Samples ...... 138

Table 6.1 Dental Crowding Frequencies in hunter-Gatherer and Agricultural Groups from Japan and North America ...... 141

Table 6.2 Dental Crowding Frequencies in Extant Horticulturalists from Brazil ...... 149

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LIST OF FIGURES

Figure 2.1 Occlusal Relationships of Central Incisors ...... 11

Figure 2.2 Craniofacial Reduction Over Time from a Dietary Shift to Softer More Processed Foods ...... 22

Figure 3.1 Map of Florida Archaic Site Locations ...... 28

Figure 3.2 Windover Site Map...... 30

Figure 3.3 Cross Sectional View of Warm Mineral Springs ...... 31

Figure 3.4 Little Salt Springs Site Map ...... 33

Figure 3.5 Location of the Republic Groves Site in Hardee County, FL, Showing Area Excavated ...... 35

Figure 3.6 Map of Tick Island and Location of Archaeological Sites ...... 36

Figure 3.7 Bay West Site Map ...... 38

Figure 3.8 Map of the Upper Basin of the St. Johns River ...... 40

Figure 3.9 directional Terms and Tooth Identifications ...... 43

Figure 3.10 Little’s Irregularity Index ...... 44

Figure 3.11 Arch Depth Calculated as Median Height of a Triangle from Arch Chords AC, BC, and AB ...... 47

Figure 3.12 Dental Wear Scoring Systems ...... 48

Figure 4.1 Severe Dental Crowding Case in Windover Individual (Burial 99) ...... 57

Figure 4.2 Severe Dental Crowding Case in Windover Individual (Burial 86) ...... 57

Figure 4.3 Dental Crowding Plotted for Windover Pond ...... 62

Figure 4.4 Scatter Plot of Crowding Severity in Windover Pond ...... 64

Figure 4.5 Age Cohorts Plotted in Windover Pond ...... 67 xiv

Figure 4.6 Windover Maxillary Mesiodistal Means for Anterior Teeth ...... 69

Figure 4.7 Windover Maxillary Mesiodistal Means for Posterior Teeth ...... 70

Figure 4.8 Windover Mandibular Mesiodistal Means for Anterior Teeth ...... 70

Figure 4.9 Windover Mandibular Mesiodistal Means for Posterior Teeth ...... 71

Figure 4.10 Windover Maxillary Mesiodistal Means for Anterior Permanent Teeth in Individuals with Good Occlusion and Dental Crowding ...... 73

Figure 4.11 Example of Severe Dental Crowing ...... 73

Figure 4.12 Windover Mandibular Mesiodistal Means for Anterior Permanent Teeth in Individuals with Good Occlusion and Dental Crowding ...... 75

Figure 4.13 Windover Mandibular Mesiodistal Means for Posterior Permanent Teeth in Individuals with Good Occlusion and Dental Crowding ...... 76

Figure 4.14 Windover Mesiodistal Means for Maxillary Anterior Teeth in individuals with Dental Crowding ...... 78

Figure 4.15 Windover Mesiodistal Means for Maxillary Posterior Teeth in Individuals with Dental Crowding ...... 78

Figure 4.16 Windover Mesiodistal Means for Mandibular Anterior Teeth in Individuals with Dental Crowding ...... 80

Figure 4.17 Windover Mesiodistal Means for Mandibular Posterior Teeth in Individuals with Dental Crowding ...... 81

Figure 4.18 Maxillary and Mandibular Mean Arch Depth Differences ...... 82

Figure 4.19 Windover Female Arch Depth Means for Individuals with Good Occlusion and Dental Crowding ...... 83

Figure 4.20 Windover Males Arch Depth Means for Individuals with Good ...... 84

Figure 4.21 Windover Mean Arch Width for Females and Males ...... 85

Figure 4.22 Windover Female Arch Width in Groups with Good ...... 87

Figure 4.23 Windover Males Arch Width in Individuals with Good Occlusion and Dental Crowding...... 88

Figure 4.24 Significant Cranial Metric Differences Between Males and Females in the Windover Sample...... 92

Figure 4.25 Significant Cranial Metric Differences Between Males and Females in the Windover Sample Continued ...... 93

Figure 4.26 Windover Attrition Comparison Between Age Cohorts in the Maxilla and Mandible ...... 95

Figure 4.27 Maxillary Attrition Means Between Age and Sex Cohorts ...... 96

Figure 4.28 Mandibular Attrition Means Between Sex and Occlusion Cohorts ...... 97

Figure 4.29 Windover Maxillary ...... 98

Figure 4.30 Windover Mandibular Attrition Means Comparison Between Occlusion Cohorts ...99

Figure 4.31 Interproximal Grooved Wear Pattern Plotted within Windover ...... 101

Figure 4.32 Example of Lingual Root Wear Pattern on Right Maxillary Third Molar from the Windover Sample ...... 103

Figure 4.33 Lingual Root Wear Pattern Plotted within Windover Pond ...... 104

Figure 4.34 Lingual Wear Pattern on Incisors Plotted within Windover Pond ...... 106

Figure 5.1 Total Crowding Based on LII Scores ...... 108

Figure 5.2 Severe Crowding Based on LII Scores ...... 108

Figure 5.3 Unerupted Unilateral Uto-Aztecan Premolar From Windover (Burial 265) ...... 111

Figure 5.4 Tick Island UAP Drawings ...... 111

Figure 5.5 UM2 Enamel Extension and UAP Frequencies for Windover, Gauthier, and Tick Island ...... 112

Figure 5.6 Lingual Talon Cusp on a Permanent Maxillary Left I2 from Windover (Specimen 75) ...... 113

Figure 5.7 Gemination of Deciduous Mandibular Incisors in the Windover Population (Burial 421) ...... 115

xvi Figure 5.8 Germination of Permanent Mandibular Incisors in the Windover Population (Burial 98) ...... 115

Figure 5.9 Concrescence of a Mandibular Third Molar in the Windover Population (Burial 123) ...... 116

Figure 5.10 Example of an Incomplete Asymmetrical Bipartite Inca Bone Variant from the Windover Sample (Burial 69B) ...... 117

Figure 5.11 Comparative Aggregate Sample of Mesiodistal Mean Dimensions Maxillary Anterior Teeth in Individuals Exhibiting Good Occlusion and Dental Crowding ...... 121

Figure 5.12 Aggregate of Maxillary Mesiodistal Averages of Posterior Permanent Teeth in Individuals with Good Occlusion and Dental Crowding ...... 121

Figure 5.13 Aggregate Mesiodistal Mean for Mandibular Anterior Teeth in Individuals Exhibiting Good Occlusion and Malocclusion ...... 123

Figure 5.14 Aggregate of Mandibular Mesiodistal Averages of Posterior Permanent Teeth in Individuals with Good Occlusion and Dental Crowding ...... 123

Figure 5.15 Mean Maxillary Arch Depth Between Windover and the Comparative Samples ...... 125

Figure 5.16 Mean Mandibular Arch Depth Between Windover and the Comparative Samples ...... 126

Figure 5.17 Mean Maxillary Arch Width Between Windover and the Comparative Samples ...... 128

Figure 5.18 Mandibular Arch Width Means Between Windover and the Comparative Samples ...... 129

Figure 5.19 Cranial Metrics: Variables with Significant Differences Between Gauthier and Windover...... 133

Figure 5.20 Cranial Metrics: Significant Differences Between Bay West and Windover ...... 135

Figure 6.1 Scatter Plot of Severity Ranks within the Northeaster Subsection of Windover Pond ...... 145

Figure 6.2 Example of Good Occlusion (Functional Occlusion) in the Gauthier and Republic Groves Samples ...... 147

Figure 6.3 Example of Severe Dental Crowding in the Windover and Gauthier Samples ...... 147

xvii

ABSTRACT

Anterior dental crowding is a condition that is more prevalent in agricultural populations than foraging societies. Though the origin of dental crowding has been debated for years, earlier studies have tied the development of this malocclusion primarily to environmental factors with minimal genetic influence. The masticatory function hypothesis (Carlson and van Gerven 1977) and disuse theory (Price 1934) both describe craniofacial changes that relate to relaxed masticatory stress resulting from dietary shifts of hard-textured foods to preprocessed foods. These changes result in the underdevelopment of the maxillae and mandible, commonly leading to inadequate space in the jaws for genetically determined tooth dimensions. This dissertation investigates the origins of the high rate (47 percent) of dental crowding in the Early Archaic Floridian Windover population (8BR246). Windover exhibits an anomalously high dental crowding rate for a prehistoric hunter-gatherer population.

To compare Windover’s dental crowding rates to populations throughout the Florida Archaic, six comparative population samples were chosen based on their comparable subsistence practices to Windover (aquatic foraging) as well as temporal and special similarities to this site. It was possible to control for possible environmental factors that may have an impact on dental use and attrition by using groups that relied on similar subsistence strategies. This research includes a comprehensive investigation of dental crowding and its relationship to tooth size, arch size, dental wear and craniofacial measurements. To analyze genetic relatedness between the samples, I examined cranial and dental non-metric traits.

The results of this study demonstrate that dental crowding development within Florida Archaic populations had different primary influences based on crowding severity. For instance, the mild/moderate crowding seen in Windover and the comparative samples is similar to rates recorded in other prehistoric foraging societies. Mild/moderate crowding in these populations might represent a normal occlusal variant and is not the product of a discrepancy between dental arch size and tooth width. Conversely, severe dental crowding at Windover is unique amongst hunter-gatherer societies (including Early and Middle Archaic groups from Florida) and appears

xviii to have a predominantly environmental etiology. Windover exhibits a much greater frequency of dental crowding (particularly severe dental crowding) than the comparative samples as well as distinctions such as rare genetic anomalies and unique dental wear patterns. It is possible that the environmental influence on severe dental crowding development is the result of cultural and sociological peculiarities of an isolated society. In particular, non-masticatory cultural practices (e.g., using teeth as tools) might alter craniofacial formation differently than is discussed in the disuse theory and masticatory function hypothesis.

My findings at Windover pond contradict the assumption that hunter-gather societies have low levels of dental crowding. I argue that dental crowding development at Windover has a predominately environmental origin and is the product of mostly non-masticatory practices. Comparative analyses of other Archaic populations demonstrate that Windover, as a population, practiced exclusive cultural techniques that lead to hunter-gatherer attrition levels and agricultural-like malocclusions. On a broader scale, information from this study has the potential to advance our knowledge of dental crowding etiology and the relationship between the human dentition and environmental factors. Thus this project has implications for understanding the underlying development of modern human cranial and dental function.

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CHAPTER 1 INTRODUCTION

The study of malocclusion prevalence in extant and past populations has shown a clear dichotomy between modern industrial societies (high prevalence) and peoples who utilize more traditional subsistence practices such as hunter-gatherer populations (low prevalence) (Begg 1954, Corruccini 1984, Larsen 1995, Price 1936, Rose and Roblee 2009). Genetic and environmental factors both play a role in malocclusion etiology, however the predominant influence on the development of malocclusion is poorly understood. The term malocclusion consists of a wide variety of dental malpositions, many with distinct developmental origins. This study focuses on anterior dental crowding. A condition found to be primarily the result of environmental factors (Corruccini and Potter 1980, von Cramon-Taubadel 2011). Dental crowding has been described as a “disease of civilization” due to its high prevalence in urban societies (Klatsky and Fisher 1953). The origin of dental crowding has raised heated debates. Discrepancies in the literature demonstrate differential researcher results: some conclude that dental crowding has a predominantly genetic etiology (Lundström 1949, Mossey et al. 1999) while others state that environmental influences are the strongest factors in dental crowding development (Begg 1954, Corruccini and Potter 1980, Harris 2008, Harris and Smith 1980, Lauc et al. 2003, Price 1934, von Cramon-Taubadel 2011). Despite disputes of dental crowding etiology, much is known about this malocclusion. For instance, dental crowding is widespread affecting most modern societies with Westernized diets (Corruccini 1984, Hunt 1966). Dental crowding is highly associated with inadequate arch sizes (Corruccini and Potter 1980; Harris and Johnson 1991, Harris and Smith 1980). Hunter-gatherer societies have a universally low incidence of dental crowding (Begg 1954, Larsen 1999, Lombardi and Bailit 1972, Price 1936). My research explores the origins of the anomalously high rate of malocclusions in the Florida Early Archaic Windover population (8BR246). The rate of malocclusions in this population (47 percent) is anomalous when compared to other hunter-gatherer populations- both prehistoric and modern (Brace 1977, Corruccini 1984, Hanihara et al. 1981, Larsen 1995, Sciulli 1997). The heavy attrition rate recorded at Windover (Wentz 2006) is, however, similar to what is observed in other hunter-gatherer populations. This makes the Windover malocclusion rate 1 appear even more unusual because it contrasts with a long-held concept of dental evolution: the disuse theory (the reduction in chewing stress leads to under-development of maxillary and mandibular bone, creating insufficient space for proper tooth positioning) (Price 1934). The frequency of malocclusions shows a trend of increasing incidence throughout recent history. This trend has been interpreted in various ways but particular attention has been paid to the correlation between higher malocclusal rates and the adoption of agriculture. Studies of the effects of subsistence changes on orofacial adaptation in prehistoric and modern Japan have confirmed the connection between diet change and malocclusion (in particular dental crowding). Three major subsistence transitionary periods in Japan demonstrated nearly a two-fold increase in malocclusion rates per period. For instance a 20 percent malocclusion frequency was documented in Jomon period (1000-500 BC) hunter-gatherers, which increased to 45.5 percent in Kofun period (AD 50-350) emergent agriculturalists and further increased to the current rate of 76.2 percent in modern industrial Japanese populations (Hanihara et al. 1981). The increases in malocclusion frequency with agriculture adoption is explained through increasing craniofacial gracialization due to less masticatory stimulus from the softer pre-processed foods found in agricultural subsistence rather than the tougher, less processed and fibrous foods consumed by hunter-gatherer societies. This concept of dietary relationship with craniofacial change is the primary focus of two long-held concepts of cranial shape and malocclusion etiology: the masticatory function hypothesis (the relationship between craniofacial shape and size and the functional mechanics of mastication) (Carlson and Van Gerven 1977) and the disuse theory (Price 1936). To further test the role of subsistence practices on malocclusion development Corruccini (1984) collected cross- cultural occlusal data from extant and past aboriginal groups (“sociotechnologically preindustrial”) who utilized traditional subsistence techniques, prehistoric Amerindians, and modern industrial agricultural groups. The prehistoric Amerinindian samples in his study were from two North American sites. The first sample consisted of skeletal remains from Dickson Mounds, Illinois ranging from Woodland period horticulturalists to Mississippian period agriculturalists. The second sample was from a population that subsisted primarily on animal protein but practiced some horticulture (including maize production) from the Libben Late Wodland site in Northern Ohio. Corruccini utilized tooth rotation and displacement scores to

2 investigate anterior dental crowding. In his study sample means and 95 percent confidence limits were compared for each group. Other than the precolonial aboriginal New Britain group from Papua New Guinea, the prehistoric Amerindian samples had the lowest rotation and displacement scores while the industrial agricultural groups exhibited markedly higher scores. Comparison of means between the Amerindian groups also showed differences reflecting subsistence influence on malocclusion: the Libben site Late Woodland sample had a recorded range of 1.0-2.0, while the Dickson Mound sample (ranging from horticulturalists through agriculturalists) showed a higher range of rotation and displacement scores of 2.0-3.0. Coupled with low malocclusion scores, hunter-gatherer populations universally exhibit more extreme rates of dental attrition than agricultural populations (Begg 1954, Larsen 1995 and 1999, Kaifu et al. 2003). Differences in dental wear severity between agriculturalists and foraging groups are attributed to diet since agricultural populations generally subsist on more soft cereals and cooked foods than the tougher meats and minimally processed foods eaten by hunter-gatherer societies (Keith 1916, Larsen 1995). Studies of dental crowding prevalence in past populations are necessary to further our understanding of the roles subsistence plays in dental crowding development as well as the obscure nature of this malocclusion. Despite numerous studies conducted on dental crowding in hunter-gatherer societies (Begg 1954, Corruccini 1999, Larsen 1995, Price 1936) including prehistoric Native American populations from the Ohio Valley (Sciulli 1997), Dickson Mounds, IL, (Corruccini 1984), and Pickwick Basin, AL (Newman and Snow 1942), to my knowledge no study of dental crowding has been conducted on Florida prehistoric populations. Windover exhibits an unusual combination of agricultural-like dental crowding rates coupled with heavy attrition common to hunter-gatherers. In lieu of the high rates of dental crowding found in the Windover collection, comparative analyses on other Florida Archaic skeletal collections were conducted to determine if the rate of malocclusion at Windover is an isolated occurrence or if other Florida Archaic populations demonstrate similar crowding.

Hypothesis I propose that the high incidence of dental crowding in the Windover population is anomalous compared to other Florida Archaic populations. I hypothesize that these high rates are

3 the result of two possibilities: 1) predominant genetic variants or 2) unique cultural practices predisposing this Archaic population to malocclusion.

I. Genetic Sub-hypothesis: The malocclusions present in the population are the result of strong genetic factors that, despite heavy attrition and robust morphology, prevent functional occlusion (naturally occurring harmonious occlusion). It is then possible that malocclusions in this population have a polygenic inheritance and the Windover population was more endogamous than other Florida Archaic populations. The increased practice of endogamy may have increased the phenotypic emergence of rare and recessive traits. Inbreeding can lead to a change in the mean value of occlusal traits if the traits were recessive or partly recessive variants. This hypothesis can be tested through genetic relatedness via non-metric analyses. This would suggest that environmental factors are not the primary forces in achieving harmonious occlusion, contradicting the attritional occlusion (Begg 1954) and disuse hypotheses (Price 1934). If the proposed hypothesis is not refuted, it may be relevant to the current malocclusion rates in modern human populations that are frequently attributed to low masticatory loading from a diet of soft processed foods rather than genetic factors.

II. Environmental Sub-hypothesis: The Windover population does not exhibit typical hunter-gatherer craniofacial robusticity due to unique cultural practices (including food preparation, diet, and tool-use) that altered cranial-facial formation. The lack of space in a more gracile jaw and face could lead to dental crowding. The degree of severe attrition in the Windover population suggests heavy masticatory loading, but it is possible that this severe attrition is the result of using teeth as tools rather than a dietary etiology. Using teeth as tools (i.e., the production of baskets, fibers, or cordage) would utilize different muscles than mastication and thus may alter craniofacial formation differently than heavy masticatory use. This may lead to different craniofacial robustiticty/gracilization than is discussed in Carlson and Van Gerven’s (1977) masticatory function hypothesis, resulting in the odd combination of heavy attrition and dental crowding. It is also possible that using the teeth as tools or habitual cultural practices are the direct causal factor of

4 malocclusions in the Windover population. This may be a phenomenon similar to ethnographic findings of the Yanomami population. However, it should be taken into consideration that the Yanomami practice a slash-and-burn subsistence and are not hunter-gatherers. Pereira and Evans (1975) recorded high rates of malocclusions among the Yanomami Indians and attributed the cultural practice of placing tobacco wads between the lips and incisors as a main environmental factor of Yanomami dental crowding. Therefore it is possible that differences in environmental exposures due to cultural practices may predispose the Windover population to malocclusion. This hypothesis can be tested using cranial robusticity measurements and tooth wear analyses.

To test the environmental and genetic components of this hypothesis, I conducted analyses of anterior dental crowding on Windover (8BR246) as well as Early and Middle Archaic comparative samples from Florida, including Little Salt Springs (8SO18), Gauthier (8BR193), Harris Creek at Tick Island (8VO24), Warm Mineral Springs (8SO19), and Bay West (8CR200). Anterior crowding is considered to be the most common form of dental crowding (Gelgör et al. 2007). While all rotations, displacements, impactions, ectopic eruptions, agenesis, and supernumerary teeth were recorded, the primary focus of this study is on anterior crowding. In this study, dental crowding is defined as insufficient maxillary or mandibular arch dimensions for erupting dentition. Inadequate arch depth to tooth size ratios is a condition greatly influenced by environmental factors (Carlson and Van Gerven 1977, Corruccini 1999, Corruccini and Potter 1980, Harris and Johnson 1991, Harris and Smith 1980, Kaifu et al. 2003, Larsen 1995, Lauc et al. 2003, Price 1936, Rose and Roblee 2009, Weiland et al. 1997). This definition was utilized to control for genetic influences on dental crowding since it mostly excludes rotations, displacements, and ectopic eruptions of anterior teeth with genetic origins. My analyses utilize the Little (1975) Irregularity Index (1975), as well as measurements of tooth size, arch depth and width, and craniofacial features. To analyze genetic relatedness between the samples, I examined cranial and dental non-metric traits. Dental attrition among these populations was scored utilizing traditional surface wear scoring systems (Smith 1984, Scott 1979).

5 Exploration of malocclusions within the Archaic Florida populations may assist with the understanding of dental crowding etiology and distribution. Information from this study will advance our understanding of malocclusion etiology and the relationship between human dentition and environmental factors. Malocclusion information from this project has the potential to reveal causal factors for the condition and, therefore, may be pertinent to orthodontic research. Thus this project further investigates the underlying development of modern human cranial and dental function.

Chapter Overview This dissertation is organized in the following manner. Chapter 2 consists of a literature review on dental crowding etiology including biological processes, and genetic and environmental factors. This information is necessary for understanding dental crowding development in Florida Archaic period populations. Chapter 3 provides an overview and description of the skeletal collections analyzed in this study and the methodologies employed to investigate dental crowding etiology. The methodologies utilized in this study include: the Little Irregularity Index, nonmetric trait frequencies (dental and cranial), dental wear analyses (attrition scores and non-masticatory wear patterns), and measurements of dental width, arch depth, arch width, and cranial shape. These methodologies provide a comprehensive system for investigating the primary factors causing the development of dental crowding in the Windover population. Chapters 4 and 5 present the results of this dissertation. Chapter 4 consists of the analyses conducted within the Windover population, and Chapter 5 presents the analyses between Windover and the comparative samples. Chapter 6 provides an overview and interpretation of the results. This chapter is organized in a similar format to the Results chapter -- first discussing dental crowding results within the Windover sample and secondly, results recorded between Windover and the comparative samples. Chapter 7 provides an overview of this study and discusses areas for future research.

CHAPTER 2 DENTAL CROWDING

Dental crowding is one of the most frequent dental anomalies in modern societies, with world population prevalences of 40-80 percent (Evensen and Øgaard 2005, Rose and Roblee 2009). Numerous studies have investigated and confirmed the importance of both genetic and environmental influences on occlusal traits in humans and other mammals. Occlusal traits are generally dependent upon variation in tooth size and position as well as facial development. Tooth size is generally accepted as being more genetically influenced (Conceição and Cardoso 2010, Greene 1967), while tooth position is more dependent on environmental factors (Harris 2008, Lauc et al. 2003). Facial growth and development, to a greater extent, is genetically determined (Harris 2008, Moyers and Wainwright 1977) although it is greatly influenced by environmental factors (Beecher and Corruccini 1981, Brown and Maeda 2004, Carlson and Van Gerven 1977, Cassidy et al. 2010, Conceição and Cardoso 2010, Rose and Roblee 2009, Schoenau and Fricke 2008, von Cramon-Taubadel 2011). Occlusion depicts the alignment between the chewing surfaces of the maxillary and mandibular teeth. The ideal alignment of the teeth is known as normal occlusion (i.e., harmonious occlusion or ideal occlusion). Malocclusion consists of any dental anomalies and deviations that lead to a misalignment of the dentition resulting in structural disharmony of the teeth and dental arcades (Lombardi 1986). These dental anomalies include teeth malpositions (i.e., rotation, displacement, tipping, impaction, hyperdontia [supernumerary teeth] and agenesis), tooth deformation, as well as sagittal, transverse and vertical dental arch malrelations (Lombardi 1986). Dental crowding, in particular anterior dental crowding, refers to tooth malpositions resulting from inadequate arch space for the size of the dental arcade. Many of these distinct conditions are correlated with one another while others are independent traits. Angle’s classes are the primary and oldest recording method of malocclusion. Edward Angle coined the term malocclusion in the early 1900s. Angle was the first to define malocclusion as well as construct categories of the defect: Class I: normal first molar alignment, Class II: overjet (overbite), Class III: underbite. The categories are based on the position of the first molars. Normal occlusion is defined as the normal anteroposterior relationship of the 7 mandible to the maxillae; the mesiobuccal cusp of the permanent maxillary first molar occludes in the buccal groove of the permanent mandibular first molar and all other teeth align without rotation, impaction or displacement (Hillson 1996). Normal occlusion ultimately manifests as tooth-to-tooth contact or slight overbite and overjet. Angle’s categories also include three subdivisions further defining the severity of the malocclusion, the last of the subdivisions for each category include crowding of the anterior teeth. The term normal occlusion is misleading since it is an idealized occlusion or perfect occlusion. Besides tooth position, normal occlusion in modern dentistry usually includes little to no attrition (tooth wear) of the cusps. However this combination of good occlusion without wear is rarely obtained in populations. This is the primary motive behind Begg’s (1954) and Brace’s (1977) arguments that non-attritional occlusion is an idealized form while attritional occlusion is the norm. Some degree of variation among individuals of a species/population is always present, so in many ways virtually everyone has some sort of malocclusion (Begg 1954, Harris and Corruccini 2008). In modern industrial societies, malocclusion occurs at rates much higher than ideal occlusal rates (Larsen 1995). This is particularly true for anterior dental crowding.

Growth and Developmental Factors Affecting Dental Crowding Despite being anchored to the alveolar bone via cementum, teeth have a tendency to migrate and shift throughout an individual’s lifetime. The four main non-pathological processes leading to tooth movement are mesial drift, anterior lingual tipping, continuous eruption and third molar eruption. In normal dentitions many of these process act as checks and balances to keep the teeth aligned. However, some of these processes can also contribute to malocclusion and crowding (particularly if there is inadequate jaw space).

Deciduous and Permanent Teeth and the Formation of Occlusion The sequential eruption of teeth is an adaptive process in mammals that allows for close correspondence between occlusal surfaces of teeth within the dental arcades (Lombardi 1986). Enamel is a static material that cannot undergo regeneration or modification except through wear, while bone matrix is dynamic and consistently remodeling throughout life. Sequential

8 tooth eruption during childhood and adolescence allows teeth to continually “fit” within the dental arcade during craniofacial growth and changes in dietary intake (Lombardi 1986). The formation of human deciduous teeth begins in utero at approximately the sixth week of development. At birth, deciduous teeth have already emerged from the crypts within the alveolar bone, however they are still covered by the gums and do not erupt until the root is mostly formed. As the permanent teeth form they push against the deciduous tooth roots, essentially breaking down the root that is then resorbed by the body. Once deciduous roots have been completely resorbed the tooth crown sheds, providing space for the eruption of the permanent tooth already fully formed with the exception of the root apex. Deciduous teeth are systematically and sequentially shed and replaced by permanent teeth throughout adolescence (Baker et al. 2005, Scheuer and Black 2000, White et al. 2011). The sequential eruption of teeth occurs whether there is ample space in the arcade or not. Inadequate space in the dental arcade for proper tooth eruption is known as tooth size arch size discrepancy (TSASD) and is the most common cause of dental crowding (Alt and Türp 1998b).

Mesial Drift Mesial drift refers to the process of dental migration within the alveolar bone toward the midline of the dental arcade (Kaifu et al. 2003). Mesial drift is a skeletal growth process that acts as a natural force to reduce interproximal space between teeth. The continuous movement of teeth toward the midline of the dental arcade contributes to the tight contact between teeth that diminishes interproximal spaces. This migratory process occurs both in juvenile and adult dentitions (Kaifu et al. 2003, Yilmaz et al. 1980). Prehistoric peoples, in particular hunter- gatherer societies, and hominin fossils exhibit high rates of interproximal attrition (Wolpoff 1971) but rarely are interproximal spaces observed. The lack of interproximal spaces in these populations is attributed to mesial drift (Fishman 1976, Hinton 1982, Hylander 1977, Nara et al. 1998). Movement of the posterior teeth in particular seems to be affected by mesial drift. In good occlusion, movement of the teeth maintains a Class I bite between the maxilla and mandible; however, in individuals with inadequate dental arcades this migration leads to pressure on the teeth that can cause crowding of the anterior dentition including impaction of erupting teeth (Lombardi 1982, Richardson 1982).

9 Despite numerous descriptions of mesial drift in contemporary and prehistoric populations, the forces behind mesial drift and questions regarding individual tooth movement and the extent of the movements are still poorly understood. However, macaque studies by Moss and Picton (1974, 1982) suggest that traction from transplantal fibers shift teeth toward the midline of the dental arcade. Although mesial drift is a developmental feature, this process is greatly reduced if the forces of mastication are inactive (Picton and Moss 1980).

Lingual Tipping Mesial drift, however, is not solely responsible for interproximal space closure. Lingual tipping of the anterior teeth resulting from dental attrition also seems to play a role in reducing space between teeth (Kaifu et al. 2003). Lingual tipping refers to the lingual movement of the anterior teeth, resulting in an upright position and tooth-on-tooth occlusion. The mechanism behind lingual tipping is poorly understood, but the counteraction of lip tension to tongue pressures is thought to be the primary driving force (Hylander 1977). Kaifu (2000b) documented lingual tipping in a Japanese hunter-gatherer skeletal sample and Hylander (1977) recorded significant lingual tipping of the maxillary and mandibular incisors (particularly the maxillary) in the prehistoric North American sample from Indian Knoll, Kentucky. In both studies, juveniles with mixed dentition exhibited scissor occlusion (a combination of overbite and overjet) between the maxillary and mandibular anterior teeth. As attrition became more advanced, the incisors and canines tilted creating a tooth-on-tooth (i.e., edge-to-edge or attritional occlusion) bite. Similar to mesial drift, lingual tipping is a process that reduces space between the teeth. In particular, this process allows for space closure within the anterior arch. However, lingual tipping occurs more prominently in individuals with progressive tooth wear and has only a minimal effect on dentitions with mild attrition (d’Incau et al. 2011). Therefore, significant lingual tipping in both jaws can be observed more clearly in hunter-gatherer societies (Kaifu et a. 2003) due to the heavy attrition common among these societies. Skeletal samples from populations practicing agricultural subsistence in Japan, Sweden and North America exhibited slight or no lingual tipping in the maxillae and no cases were recorded in the mandible (Hasund 1965, Fishman 1976, Kaifu 2000b, Lundström and Lysell 1953, Lysell and Fillipson 1958, Mohlin et al. 1978).

10 Tooth-on-tooth occlusionon (attritional occlusion) is common across hunterter-gatherer populations and throughout humaman evolution. It was believed humans have underdergone a rapid occlusal transition in the last thouousand years from tooth-on-tooth occlusion to scissorsci occlusion (Brace 1977, 1986). This transitioition was directly linked to decrease in tooth wearar in populations changing from hunter-gatherer susubsistence to agriculture. More recent investigatationi (Kaifu 2000b), based on hunter-gathererrer societies, suggests that newly erupted permanennent teeth tend to be inclined labially (toward thee lilips), assuming a natural scissor occlusion. Withth age and advanced attrition development,t, ooverbite decreases as anterior tooth wear advancnces and overjet reduces as a result of lingual tippipping (Kaifu et al. 2003), creating attritional occclusionlu (Figure 2.1). With decreased attrition, the conversion from scissor occlusion to tooth-on-tooth occlusion reduces, resulting in the retentionion of overbite and overjet. Besides attritional levels,ls, there are craniofacial developmental differencecess between hunter-gatherer and agriculturall ssocieties that result from variations in masticatortory stresses among populations. These skeletaletal differences also contribute to the occlusal transitionran seen in hunter-gatherers. This is discussessed in detail in the Environmental Influences sectctioni below.

Figure 2.1: Occlusall rerelationships of central incisors from Kaifu et al. (2003): 52.5 (A) Scissors Occlusion (B) Edge-to-Edge Occlusion (C) Tooth-on-Tooth Contact or Attritionitional Occlusion.

11 Third Molar Eruption Another biological process responsible for tooth movement within the alveolar bone is third molar eruption. The third molar is the last permanent tooth to erupt, coming into full occlusion almost a decade after the eruption of the second molar. If there is inadequate space in the jaws for third molar occlusion, the force generated during the eruption has been recorded to intensify existing incisor crowding and affect buccal or lingual inclination of the second molars (Niedzielska 2005). Richardson (1979) recorded movement of the first molar after a change in space condition in the mandible resulting from third molar eruption. She also has recorded positive correlations between third molar impaction and increased anterior crowding (Richardson 1982). Third molars are an often-cited culprit of increased anterior crowding with age, yet studies have shown mixed results regarding the role of the third molar in dental crowding development (Harris 1997, Niedzielska 2005). Most hunter-gatherer samples and early hominins exhibit ample jaw space for third molar eruption and functional occlusion. Thus the third molars do not commonly alter occlusion or lead to dental crowding in these populations. It has been suggested that increased incisor crowding with age is predominately caused by occlusal and physiologic forces that alter the arcades regardless of third molar eruption (Harris 1997). Despite negative associations between crowding and third molar retention (Kaplan 1974, Little et al. 1981, Björk and Skieller 1972), it cannot be denied that the third molar does play some role in anterior crowding and overall tooth movement in certain cases. Third molars appear to have the most effect on individuals with TSASD, in particular, individuals with large tooth dimensions. Richardson (1982) recorded a higher third molar impaction rate in individuals with significantly larger teeth. Increases in third molar agenesis also have been recorded as a more recent trend in human dental evolution. Third molar agenesis may be linked to the overall decrease in tooth dimensions recorded throughout human evolution and in particular in Post- Pleistocene agricultural societies (Carlson and Van Gerven 1977, Pinhasi et al. 2008). This may be suggestive of relaxed selective pressures on larger teeth and full dental arcades, or it may indicate that large tooth size and third molars are now maladaptive (Lombardi 1982).

12 Continuous Eruption The concept of continuous eruption refers to the active eruption of teeth throughout one’s life. However, the mechanisms behind this biological process are poorly understood (d’Incau et al. 2011). Similar to lingual tipping, continuous eruption is more commonly seen in populations that undergo heavy dental wear or antemortem tooth loss (Kaifu et al. 2003). Though more commonly recorded in permanent dentition, continuous eruption also occurs in deciduous teeth as well (Bimstein et al. 1990). Continuous eruption has been recorded as an increase of the cementoenamel junction- alveolar crest (CEJ-AC), acting as a compensatory process that retains tooth height. Despite this CEJ-AC increase the alveolar bone shows no significant change in height (Kaifu et al. 2003), therefore this is a process primarily affecting the tooth. This increase of the CEJ-AC with wear has been documented in Australian aborigines (Murphy 1959), prehistoric Native Americans from Indian Knoll (Hylander 1977), Romano-British samples (Whittaker et al 1982) and preindustrial Irish populations (Glass 1991).

Genetic Factors Dental crowding has a multifactorial genetic etiology since many of the orofacial structures involved are produced by a combination of genes interacting with the environment (Harris 2008, Mossey et al. 1999). There seems to be different degrees of genetic influence on different malocclusion traits. For instance, deep overjet and underbite, arch size, and individual tooth displacements or individual tooth rotations seem to have strong genetic origins. Genetic influence is considerably less, however, on the development of overbite, dental arch shape, crowding and general tooth rotation and displacement (Harris and Corruccini 2008, Lauc et al. 2003). In general, variables pertaining to the position and crowding of teeth have a stronger environmental basis (Corruccini and Potter 1980, Harris and Smith 1982). However, studies of familial similarity in the shape of the dental arc and in tooth position have recorded contradictory results. In particular, familial similarities exhibited strong genetic influences on the position of the second premolar and the first molar, but positions of the maxillary and mandibular incisors exhibited the least familial similarities (Hu et al. 1992).

13 The genetic factors influencing malocclusion are believed to be polygenically determined (many genes at different loci interacting with environmental factors leading to phenotypic variations). Polygenetic inheritance is most commonly associated with traits that manifest along a continuum such as human stature. This mode of inheritance differs from simple Mendelian inheritance since expression of the extreme range of a trait actually implies a greater frequency of affected relatives rather than a weak expression (Lauc et al. 2003). This finding is attributed to the additive effect of genes on polygenetic phenotypes and the typically large number of genes that contribute to the overall manifestation of that trait. Harris (2008) describes the effect as being analogous to a large array of light switches. Each switch operates independently but the more switches that are in the “on” position, the greater the expression of the trait.

Studies of Heredity Teeth are unique and useful objects for genetic investigation since each tooth is a single morphological unit that presents itself twice in each individual (Lundström 1977). Each of these morphological units is governed by additive genes, non-additive genes, and to a lesser extent environmental influences. As discussed previously, the high mineralization of enamel creates a fossilization effect of the tissue thus inhibiting further development after the crown is fully formed. While teeth are separate morphological units, they are inter-correlated with one another. In particular, adjacent teeth are more highly correlated to each other than to more distant teeth within the arcade. Twin, familial, endogamy, population and cross-cultural studies have demonstrated significant heritability of tooth size and morphology, arch length, Angle malocclusions, and individual tooth malpositions.

Dental Size and Morphology The dentofacial traits shown to be predominately influenced by genetic factors are tooth size and tooth morphology (Calcagno 1989, Greene 1970). Tooth size is usually determined by three measurements: mesiodistal (MD) and buccal-lingual (BL) dimensions and crown height (CH). In particular, MD and BL dimensions are highly correlated with one another (Dempsey and Townsend 2001). In a study by Dempsey and Townsend (2001), additive genes were found to mostly account for overall tooth size and morphology than non-additive genes. Additive gene

14 pairs have an equal chance of expression. Trait expression effected by additive genes is based on the sum of the effects of individual genes. Non-additive gene pairs are not equally expressed since one will dominate in the phenotype. Non-additive traits are more susceptible to natural selection (Kacser and Burns 1981). In the Dempsey and Townsend (2001) study, they found that non-additive genetic variation mostly affected MD diameters of teeth. Conversely, environmental influence on tooth size and shape was extremely low, especially in the permanent dentition. In their study, the only teeth shown to have a moderate amount of exogenous influence (approximately 25 percent) were the first molars. This may be due to the early development of the first molars in utero (being the first permanent teeth to erupt) and, therefore, are more susceptible to prenatal and early postnatal environmental influences. Tooth size is highly hereditary although its influence on dental crowding has shown contradictory results. In a study comparing tooth size and malocclusion between two groups of young adult females from the United States, Peck and Peck (1972) found overall larger MD widths of the anterior teeth in the sample with gross crowding and lower means in groups with good alignment. Doris et al. (1981) compared MD tooth widths of North American male and female dental patients with severe dental crowding to individuals with little to no crowding. Crowding ranges were determined using Little’s Irregularity Index (LII). They found overall significantly larger anterior tooth width in the severe malocclusion group. Bernabé and Flores- Mir (2006 b) analyzed MD anterior crown widths in children from Lima, Peru. Crowding metrics were conducted utilizing the LII scoring system. The authors recorded significantly larger MD widths in children (both male and female) with severe crowding than in the good alignment groups. Contradictory results were found in other studies. Mills (1964), in his study of adult males from the US Naval Academy, found no correlation between incisor MD width and crowding. Radnzic (1988) analyzed the relationship between MD crown dimensions and dental crowding in a randomized selection of British and Pakistani boys in England between 13 and 16 years of age. His results were similar to Mills (1964), with no significant association between MD width and dental crowding.

15 Arch Dimensions Arch length, width, and shape are crucial developmental factors in the etiology of dental crowding. The disproportion between the size of the jaws and the teeth (TSASD) is the most common recorded cause of dental crowding (Alt and Türp 1998b). Arch shape has been recorded as having a high degree of heritability within families. Harris and Smith (1980) derived arch length estimates from three different types of first-degree relative pairs: sibling-sibling, mother-offspring, and father-offspring. They found that arch dimensions, including arch length, width and shape, involved few sources of environmental covariation leading them to conclude that the influence of genetics on arch dimensions is moderately high. The most heritable similarities were between siblings, followed by mother- offspring and then father-offspring pairs. Corruccini and Potter (1980) used monozygotic and dizygotic twins to investigate the heritability of occlusal variables. Like Harris and Smith (1980), they reported arch length and width as exhibiting significant intrapair differences in the dizygotic twins. This result suggests that these arch dimensions are a highly heritable component of the dentofacial complex. In both of the aforementioned studies, dental crowding was not found to be under significant genetic influence and to be predominately the result of environmental factors.

There are many studies that have shown similarities between occlusal traits and dentofacial morphology within families. However, as Harris and Potter discuss (1980), there are two general factors that need to be taken into consideration: the sharing of common genes and the sharing of common environments. In many heredity studies, the latter situation is often overlooked and resemblances between family members are concluded to be due to genetic influence rather than shared environmental factors. While all dental arch dimensions are recorded to have strong heritable components, these traits are also highly influenced by exogenous factors (Beecher and Corruccini 1981, Calcagno 1989, Carlson and Van Gerven 1977, Cassidy et al. 2010, Rose and Roblee 2009, von Cramon- Taubadel 2011). Arch dimensions are discussed further in the Environmental Factors section of this chapter.

16 Angle Malocclusion Classes As discussed previously, Angle’s malocclusion classes categorize occlusion by the relationship and position between the maxillary and mandibular first molars. Angle’s categorizations for malocclusion are not necessarily indicative of dental crowding since many of these classes can be present without dental crowding. However, Angle’s sub-classifications for the classes include subdivisions that incorporate dental crowding. Severe Class II and Class III cases have been consistently shown to be highly heritable occlusal traits. Probably the most famous genetic occlusal variant is the overt Class III malocclusion in the Hapsburg family. The Hapsburgs were Hungarian/Austrian royals and were famous for their mandibular prognathism that manifested in several generations. Studies of the Hapsburgs by Strohmayr (1937) and Wolff et al. (1993) concluded that the Class III malocclusions within the family were the result of an autosomal dominant trait. However, despite the independent occurrence and prevalence of this trait within European noble families (Wolff et al. 1993), the autosomal dominant genes responsible for these cases could be regarded as exceptional and not a typical mode of underbite inheritance (Mossey et al. 1999). In an endogamy study of subadults from the Island of Hvar, Croatia, Lauc et al. (2003) concluded that the Angle Class II and Class III traits exhibited strong hereditary tendencies and are most likely the product of rare and recessive genetic variants, not autosomal dominant traits; the researchers, however, did not discount the role of polygenetic additive effects as well. They proposed that inbreeding could predispose individuals to these malocclusions. They found this especially true for severe manifestations of the Class II and Class III malocclusions. Lauc et al. (2003) also found that the inbreeding effect observed in these populations was not recorded in all traits, only those with a significant genetic basis (i.e., overjet); there was only a negligible effect on dental crowding.

Dental Traits Affecting Malocclusion Individual tooth rotations, displacements, agenesis, supernumerary development, impaction and congenital malformations are the most notable genetic traits that can affect malocclusion and dental crowding in individuals.

17 Individual Rotations and Displacements Unlike general rotations and displacements, individual cases routinely have adequate arch space to erupt in normal occlusal position. This is not to say, however, that individual tooth rotations and displacements do not also occur along with general dental crowding; individual and general dental malpositions are not mutually exclusive but they have different etiologies. The most notable individual tooth rotations and displacements with genetic origins are incisor winging (the bilateral rotation of the central incisors) (Escobar 1976, Iizuka 1976), palatally displaced canines (bilateral displacement of the canines toward the palate) (Peck et al. 1994, Pirinen et al. 1996), and rotated premolars (Hu et al. 1992, Rougier et al. 2006). The genetic etiology for these tooth malpositions is still poorly understood. There appears to be a familial segregation patterning of these traits that is not as simple as Mendelian inheritance, nor is it as complex as polygenetic traits with additive effects. Scott and Turner (1997) therefore suggest the quasicontinuous inheritance model for many of the morphological dental variants and some individual tooth malpositions (in particular bilateral winging). Quasicontinuous inheritance is a polygenetic model, first proposed by Grüenberg (1952). It describes certain traits as being phenotypically discontinuous while having continuous genotypic distributions; the continuous genotype has both an underlying component and one that is phenotypically expressed, producing many genes at subsequent loci interacting with one another (Harris 2008, Scott and Turner 1997).

Supernumerary Teeth, Impaction, and Agenesis Disorders of tooth number and eruption can be divided into four categories: supernumerary teeth, supplementary dental formations, missing teeth, and impacted (ectopic) teeth. Impacted teeth may occur as a result of complications from any of the other traits (as well as deciduous tooth retention). Supernumerary teeth (also known as hyperodontia, polydontia, and polygenesis) are excess teeth of the normal deciduous or permanent dentitions. These teeth can either take the form of separate tooth formations or they may be fused with existing teeth (gemination) (Alt and Türp 1998a). Supernumerary teeth (polygenic teeth) is a term used for excess teeth that mimic the normal shape of a tooth, while accessory teeth describes excess teeth that do not resemble the

18 normal form such as peg teeth (Bhaskar 1973). These teeth are often impacted. Supernumerary teeth have a strong genetic component to their etiology (in particular, expressed as the simultaneous occurrence of multiple supernumerary teeth), although their etiology is still poorly understood (Wang and Fan 2011). Supernumerary teeth can occur anywhere in either dental arcade but most commonly develop in the anterior and molar portions of the maxilla, and the premolar portion of the mandible. The most common manifestations of supernumerary teeth (in order of frequency) are mesiodens, paramolars, and distomolars (Lavelle and Moore 1973). Mesiodens account for approximately 50 percent of all polydontic cases, and are located in the anterior portion of the maxilla (Alt and Türp 1998b). In particular, premaxillary conical mesiodens located between the maxillary incisors are the most frequent supernumerary teeth reported (Mossey et al. 1999) primarily occurring in the anterior dentition. Supplementary dental formations take the form of accessory microteeth (abnormally small teeth), interradicular teeth (separate teeth germs fused to the roots of existing teeth), enamel pearls, and enamel extensions (Alt and Türp 1998b). Similar to supernumerary teeth, accessory microteeth may lead to crowding of the affected dental region. There is an association between microdontia and the congenital absence of teeth (tooth agenesis, hypodontia, aplasia). However, supernumerary/accessory teeth appear to have a separate genetic origin while general microdontia and hypodontia have been recorded to share a genetic link (Baccetti 1998). Hypodont teeth and microteeth are considered to be different manifestations of aplasia along a continuum of expression. In other words, these traits are all considered microsymptoms from the same etiology hindering tooth germ development; it’s most extreme manifestation results in aplasia (Alt and Türp 1998b). Bjerklin et al. (1992) reported associations between hypdontia and first maxillary permanent molar impaction in their familial study. Specifically, the congenital absence of premolars and ectopic eruption of maxillary first permanent molars were found to be significantly associated showing a common hereditary etiology. Hypodontia, microdontia and associated impaction are believed to have a multifactorial polygenetic inheritance. Harris (2008) advises that the term “heredity” should be used with caution especially with regard to continuous traits, since control of environmental and genetic confounders is often impossible even in monozygotic twin studies. This is especially true as there are numerous sources of environmental covariation that often lead to a convergence of phenotypes resulting in

19 an inflated estimate of heredity. However, Harris (2008) further states that these environmental sources can be accounted for as confounding factors as long as the sources can be recognized and they can be measured. In summary, there is strong evidence for genetic etiologies for individual dental traits and dentofacial structures. However, environmental factors are believed to play a predominant role in general anterior dental crowding. Numerous studies have shown negligible to no genetic influence on general dental crowding; overall anterior tooth-based issues such as malpositions, rotations and displacement seem to predominately be a consequence of environmental factors (Corruccini and Potter 1980, Harris 2008, Harris and Smith 1980, Lauc et al. 2003).

Environmental Etiology Experimental, osteological and cross-cultural comparative studies have been conducted to assess the environmental factors involved in dental crowding. The current epidemic of dental crowding has been attributed to a dietary shift in recent history involving cooked foods and “soft cereals replacing tough meats” and minimally processed foods (Keith 1916:198). An excellent example of environmental influences on malocclusion was a study conducted by Weiland et al. (1997) that showed changes in malocclusion (including dental crowding) in Austrian men from the 1880s to the 1990s that corresponded to dietary changes. Similar findings of elevated malocclusion rates have been recorded in societies that have undergone subsistence shifts from hunter-gatherer lifestyles to agricultural style subsistence. Changes in the oral environment have been recorded to have a strong influence on the apposition, shape, and resorption of maxillary and mandibular bone (Beecher and Corruccini 1981, Calcagno 1989, Harvlod et al. 1973). Differences in masticatory stress have also been observed to result in craniofacial changes over time, including more gracile and less prognathic maxillomandibular complexes (Carlson and Van Gerven 1977). These craniofacial changes are believed to be a major driving force in arch size tooth/size discrepancies, a predominant aspect of dental crowding.

Masticatory Function Hypothesis The shift from hard to soft textured foods is believed to be the primary force in the recent human evolutionary trend toward craniofacial gracialization consisting of more posteriorly

20 placed faces and a general reduction in size and robusticity of faces, jaws and teeth (Carlson and Van Gerven 1977). This reduction in craniofacial robusticity has been well documented in populations transitioning from foraging to farming in both the Old and New Worlds (Brown and Maeda 2004, Carlson and Van Gerven 1977, Corruccini 1999, Larsen 1995, Rose and Roblee 2009). Carlson and Van Gerven (1977) developed the masticatory function hypothesis based on skeletal analyses of the Nubian transition from hunter-gatherer to agricultural subsistence. They traced morphological change in Nubian skulls from the Mesolithic through the Christian periods (approximately 10,000 years). They argued that craniofacial gracilization was not a result of genetic factors but a direct result of a reduction in chewing stress during development. Reported craniofacial changes included an increase in height of the cranial vault and a decrease in overall cranial length, an inferoposterior shift of the midface and lower face relative to the anterior portion of the cranium, and a decrease in overall robusticity of the craniofacial complex (most notably a decrease in the growth of the maxillomandibular complex) leading to more gracile maxillae and mandibles and decreased prognathism (Figure 2.2). Tooth rows were also noted to shift distally in relation to the cranium. Carlson and Van Gerven (1977) also found that these changes covaried together and in the same direction. In particular, the size of the mandibular body, height of the mandibular ramus, length of the cranial vault, as well as the size and attachment of masseter muscle all changed together in directionally similar ways. They also found inverse relationships among mandibular size and shape, masseter attachment, and cranial length. For instance, individuals with large mandibles also had large masseter muscles and longer/lower cranial vaults, while individuals with smaller mandibles had smaller masseter muscles and shorter/higher cranial vaults.

Figure 2.2: Craniofacial reduction over time from dietary shift to softer more processed foods. Arrows represent a reduction in mastication and maxillomandibular complex and cranial length. Uppermost arrow represents the heightening of the cranial vault. Dotted lines outline these changes. Image adapted from Carlson and Van Gerven (1977): 502.

Other studies have shown similar results to Carlson and Van Gerven, including a chronological study by Rose and Roblee (2009). Analyzing human skeletal remains from the last 10,000 years in Egypt, they demonstrated increased gracilization associated with the development of agriculture (Rose and Roblee 2009). These results were also supported by numerous studies that recorded correlations of diminished use of jaw muscles and subsequent growth reduction in the maxilla and mandible (Beecher and Corruccini 1981, Brown and Maeda 2004, Kaifu 1997). Carlson and Van Gerven argued that these craniofacial changes are attributable to reduced chewing stress during development and that they are not the product of genetic changes. These responses to environmental changes result in the underdevelopment of the maxillae and mandible, commonly leading to inadequate alveolar space for genetically determined tooth dimensions. Studies have also recorded an overall reduction in dental size in conjunction with dietary changes (Calcagno 1989, Carlson and Van Gerven 1977, Greene 1970); more recent dental changes are attributed to adaptations resulting from natural selection for morphologically 22 less complex caries-resistant teeth (Larsen 1999). Reduction of dental complexity is considered to have a predominately genetic origin and is an independent process from craniofacial changes with a primary environmental etiology (Carlson and Van Gerven 1977). With the exception of attrition, environmental effects have little influence on the size and morphology of teeth (Calcagno 1989). It also has been noted that although human teeth have become smaller, this has not led to excess room in the jaws; conversely, there is often inadequate space for teeth to erupt in proper alignment. Thus dental crowding may be considered an alveolar bone deficiency (Rose and Roblee 2009).

Disuse Theory Based on observations of Inuits and other native and rural populations from around the world, Price (1936) developed the disuse theory that attributes dental disease and malocclusion to diets based on processed foods. In general, Price observed low incidences of dental disease and malocclusion in populations who ate traditional, minimally processed foods. In the Inuit populations, he observed widespread good occlusion until the incorporation of industrialized highly processed foods in their diet; this shift was correlated with a 50 percent increase of malocclusions (Price 1936). Corruccini (1999) built upon the disuse theory and the masticatory function hypothesis by conducting extensive cross-cultural comparisons testing the hypothesis that reduction in chewing stress leads to under-development of maxillary and mandibular bone and creates insufficient space for proper tooth positioning. He also observed malocclusion in several extant populations including rural Kentucky, Punjabi communities from northwest India, and Chinese immigrants to Liverpool. In all of these populations Corruccini noticed a rapid increase in the incidence of malocclusions after the introduction of an industrialized diet. The changes were observed after only one generation (Corruccini 1999). This is compelling evidence for environmental influences since many of these populations consisted of isolated breeding groups with little or no gene flow.

Dental Wear and Attritional Occlusion Paralleling craniofacial reduction is a decrease in tooth wear in modern populations. Tooth wear (dental attrition) is a direct result and indication of masticatory use and diet (Larsen

23 1995, Smith 1984). Attrition occurs from a variety of processes, including tooth-to-tooth contact (i.e., mastication, bruxism), abrasion (friction resulting from exogenous material during mastication or using the teeth as tools), and erosion (chemical dissolution of tooth surfaces, mostly as a result of highly acidic diets) (Kaifu et al. 2003). Dental wear rates are highly influenced by the consistency and texture of food, which is either a direct characteristic of that food, how heavily it is processed before mastication, or the amount of grit present. Tooth wear is highly confounded by age since older individuals have been exposed to the processes of attrition for a longer period of time (Larsen 1995). Wear severe enough to obliterate cusp morphology, create interproximal space, and flatten occlusal surfaces was ubiquitous in almost every prehistoric society and among fossil hominins (Brace 1979, Kaifu et al. 2003, Rose and Roblee 2009, Wolpoff 1971). Relatively recent hunter-gatherers, such as Australian Aborigines and Inuits, experienced heavy dental wear until adopting a more Westernized lifestyle (Begg 1954, Kaifu et al. 2003, Molnar et al. 1983). Heavy occlusal wear was also ubiquitous and normal among ancient homininds. A study by Kaifu (2000 a) demonstrated the universality of wear rendering the occlusal surface flat in hominid fossils from the genus Homo in the course of the past two million years. These observations support the notion that human dental traits have been selected for the inevitability that extreme wear will occur. This seems only logical since humans have evolved, until recently, in “heavy-wear environments” (Kaifu 2003 47). Thus, dental problems such as malocclusions in contemporary societies seem to be related to the discrepancy between our dental design and the modern environment, which does not allow for the same masticatory loading and dental wear (Kaifu et al. 2003, Rose and Roblee 2009). This discrepancy is the basis of Begg’s (1954) concept of attritional occlusion. Attritional occlusion is described as harmonious occlusion resulting from heavy attrition. The concept is based on three primary assumptions. First, ancient populations in heavy-wear environments undergo continuous and dynamic dental changes as a direct result of attritional tooth reduction throughout their lifetime. This loss in tooth mass creates space that is filled by compensatory tooth migration. The second assumption is that contemporary peoples have inherited these compensatory mechanisms but the lack of wear has resulted in the failure to develop attritional occlusion. Lastly, this failure has led to the increase in dental anomalies seen today such as dental crowding and other malocclusions.

24 As discussed above, anthropological and dental research suggests that there are three kinds of physiological tooth migration processes: mesial drift, continuous eruption and lingual tipping. Most researchers agree that mesial drift of the posterior teeth and continuous eruption of all the teeth are universal processes found in all human populations. Data for lingual tipping of the anterior teeth has been documented primarily in prehistoric peoples with heavy attrition (Hylander 1977, Kaifu et al. 2003). These three processes support Begg’s assumption that dental wear is a factor in achieving attritional occlusion. Although attrition does create space for tooth migration and eruption, it is only a single component of the functional occlusion found in pre-agricultural societies. Heavy attrition is not necessary for harmonious occlusion because functional occlusion can occur without severe dental wear (Corruccini 1999, Harris and Corruccini 2008, Kaifu et al. 2003). Heavy chewing stress usually results in heavy attrition, thus dental wear severity is a good indicator of the degree of chewing stress and dietary coarseness. The chewing stress required to masticate minimally processed foods during development increases the size and position of the masseter and pterygoid muscles, which in turn alters the position and size of the mandible; the jaws become more robust, the alveolar bone becomes more dense, and third molars have room to erupt and function normally. Therefore the masticatory function and disuse hypotheses better delineate the major environmental influences of dental crowding than Begg’s (1954) notion of attritional occlusion. Robust features, heavy attrition, and functional occlusion are the norm in the available literature on hunter-gatherer populations; therefore the high rate of dental crowding and other malocclusions found in the Windover population appears to be atypical. This Archaic population demonstrates malocclusion rates comparable to those of agricultural societies. The Windover skeletal population appears to contradict these long-standing theories of the environmental origins of malocclusions. Dental and cranial analyses from this collection and comparative samples were conducted to further explore the significance of these observations.

CHPATER 3 MATERIALS AND METHODS

This chapter discusses the skeletal samples, osteological techniques and statistical methodology used to investigate dental crowding and the potential environmental and genetic factors involved in its etiology. Samples consisting of adult and subadult skeletal remains were analyzed from seven Florida Archaic collections dating from the Archaic period. Only crania, maxillae (if separate from crania) and mandibles were included in the samples. Isolated teeth were not studied and individuals exhibiting pathologies that confound dental crowding were excluded. The populations sampled exhibit large degrees of sexual dimorphism; therefore confident sex estimations of the crania could be made without the postcrania. Sex and age data were considered for each individual to assess developmental differences in dental crowding, and to evaluate any evidence for sexual divisions of labor and activities affecting the dentition. The Little Irregularity Index (Little 1975) was used to compare dental crowding between the samples. Genetic relatedness and population similarities were studied using presence/absence frequencies of dental and cranial non-metric traits. Measurements of mesiodistal tooth size and arch dimensions were used to investigate arch/tooth size discrepancies. Potential environmental factors, masticatory effects, and non-masticatory uses of the dentition were analyzed utilizing cranial metrics, dental attrition, and tooth wear patterns.

Primary and Comparative Skeletal Collections

The osteological samples in this study are from seven Florida Archaic populations and include my primary sample (Windover, Early Archaic period) and six comparative samples (Table 3.1). These collections are from the Early to Middle Archaic periods in Florida (including some individuals from the Late Archaic period in the Gauthier collection). I chose Floridian Archaic populations as my direct comparative samples to reduce the biological and genetic variability between these samples and Windover, and to give an overview of attrition and malocclusion during the Florida Archaic.

26 The Archaic period in Florida covers a vast time range, spanning a period of approximately 10,000 years. Early Archaic samples are ideal for comparison with Windover, however there are few skeletal collections with good preservation from this period. North American skeletal collections dating prior to 7,000 BP are rare and 60 percent of the total individuals from these collections are from Florida sites (n = 194), with the majority of those individuals belonging to Windover (n = 168) (Doran 2002). The only Florida Early Archaic comparative collection with good dental and cranial preservation is Warm Mineral Springs. Middle Archaic populations are the next best collections available for comparison, and these comprise the majority of my comparative samples. Due to the temporal distance between the Early and Late Archaic periods, samples from the Late Archaic were kept to a minimum. The six comparative populations were also chosen because of their primarily riverine or marine subsistence practices. Samples with similar subsistence practices were selected to control for possible environmental factors that may have an impact on dental use and attrition.

Table 3.1- Skeletal Samples Temporal Range Site 14 Subsistence N Reference C yrs. BP

Windover Early Archaic Doran and Dickel 1988: Riverine 89 (8BR246) 8,120-6,990 367 Warm Mineral Springs Early Archaic Clausen et al. 1975: 191, Riverine 5 (8SO19) 11,950-9,950 Lien 1983: 106

Little Salt Springs Middle Archaic Clausen et al. 1979: 611, Riverine 6 (8S018) 6,900-6,085 Pääbo et al. 1988: 9776

Bay West Middle Archaic Marine 9 Beriault et al. 1981: 39, 50 (8CR200) 6,710-6,550

Republic Groves Middle Archaic Riverine 6 Purdy 1991: 173 (8HR4) 6,585-5,640

Harris Creek Middle Archaic at Tick Island Riverine 34 Jahn and Bullen 1978: 22 5,640-5,030 BP (8VO24) Middle-Late Gauthier Archaic Riverine 20 Maples 1987: 2 (8BR193) 3,300- 1,600 BP

Skeletal collections analyzed in this study were Windover (8BR246), Warm Mineral Springs (8SO19), Little Salt Springs (8SO18), Harris Creek (8VO24), Republic Groves (8HR4), Bay West (8CR200) and Gauthier (8BR193). As indicated in Fig. 3.1, they are broadly distributed across the central portion of Florida and along water systems linked to the Gulf and Atlantic coasts.

Figure 3.1: Map of Florida Archaic Site Locations

Windover (8BR246) Windover is a collection of over 160 well-preserved Early Archaic skeletons. Located in east-central Florida near present-day Titusville, Windover was a large charnel (mortuary) pond where the dead were buried in shallow water underlain with peat (Figure 3.2). The anaerobic conditions of the peat environment led to remarkable preservation of textiles, wood, bottle gourds, bone and soft tissue. Notably, 91 individuals have preserved brain matter (Doran 2002). Fabric and wooden stakes were found in association with the burials. It is possible that individuals were wrapped in fabric that was fastened to the pond floor using small stakes. Larger stakes were also recovered in association with the burials. These were embedded vertically, suggesting that they were initially erected to stand out of the water to demarcate individual graves (Doran 2002, Purdy 1991). This form of water burial, characterized as a “mortuary pond”, is unique to the Florida Archaic periods (Doran 2002, Wentz and Gifford 2007) and was also employed by a number of the comparative populations. The population buried at Windover was a semi-sedentary hunter-gatherer society whose subsistence consisted mostly of riverine resources. Paleodietary analyses, based on abdominal samples taken from the area of the sacrum and nitrogen carbon isotopic analyses of bone collagen, affirm the focus on inland river, marsh and pond resources (Tuross 1994). In addition to aquatic fauna, some terrestrial mammals (deer and rabbit) as well as botanicals such as fruits, nuts, greens, seeds and tubers were also consumed, indicating a diverse subsistence strategy (Newsom 2002). Previous health assessments of the Windover sample (Wentz 2006) show the rate and severity of attrition in the population is comparable to other hunter-gatherer populations. Attrition rates were recorded to be high among adults, with subadults demonstrating moderate attrition of the deciduous dentition. The Windover population has few carious lesions (Wentz 2006); this is generally attributed to rapid loss of occlusal features, in conjunction with a low carbohydrate diet. The condition of the Windover collection necessitated some cranial reconstruction, but overall Windover exhibits excellent preservation. This study includes 89 of the crania.

Figuree 3.3.2: Windover Site Map, from Doran (2002): 5.

Warm Mineral Springs (8SO1919) Warm Mineral Springs is an Early Archaic site located in and around a 70-meter deep collapsed sinkhole in Sarasota CoCounty (Figure 3.3). The skeletal remains were recoveredre both in and around the spring-fed sinkholhole and they arguably represent some of the oldesdest humans recovered in Florida (Clausen et aal. 1975, Royal and Clark 1960). Similar to Winindover, the anaerobic conditions of the sinkhkhole allowed for some remarkable preservationn includingin the first recovery in Florida of prehistoricric brain tissue. There are no clear burial pits or markersma at Warm Mineral Springs and it is debateded whether or not the interments were intentionalal (Clausen( et al. 1975, Purdy 1991). Excavation frfrom the 1950s through the 1970s exhumed skeleeletal material representing approximately 30 inindividuals (Purdy 1991). Portions of seven humaman skulls were recovered (six adults and one sububadult). Of those seven skulls, one was completeete and only three retained enough elements for partartial reconstruction. Provenience of many of thee individualsi was poorly recorded and their temporaoral associations with one another are difficult to assess.a Despite

30 provenience issues, the early dateate, the location of the site, and the good preservvationat make Warm Mineral Springs an appropriate cocomparative sample for Windover. Unfortunatelytely, only three crania were suitably preserved fofor this study.

Figure 3.3: Cross sectional view of WWarm Mineral Springs. Map indicates the location of the excavation site during the Clausen et al. (1975)75) excavation. The human crania were recovered from the 13m ledge. Figurere ffrom Purdy (1991): 182. Figure not to scale.

31 Little Salt Springs (8SO18) Little Salt Springs is a Middle Archaic Period habitation and underwater burial site consisting of a large flooded sinkhole located in Sarasota County (Figure 3.4). A slough, pond basin and midden constitute the primary area of the Middle Archaic cemetery and this received a different state site number (8SO79) (Wentz and Gifford 2007). For the sake of simplicity only the predominate state site number (8SO18) is used in this study. Numerous hearths (now underwater) and associated artifacts were recorded around the perimeter of the sinkhole reflecting multiple occupations during the Early and Middle Archaic periods (Clausen et al. 1979). The site represents several thousands of years of alternating occupation and abandonment, but burials were only recovered from the Middle Archaic period. Underwater mortuary practices, similar to Windover, were found at this site. The dead were interred in the peat-laden shallows of the slough, and as water levels dropped burials extended into the pond basin (Purdy 1991). The full extent of the cemetery, however, is unknown. Consistent water temperature and an anaerobic environment resulted in good preservation of artifacts and skeletal remains. Brain material was also recovered from Little Salt Springs. Plant fiber shrouds made from grasses, wax myrtle and grape vines were found in associated with the burials. Purdy (1991) suggested that the bodies were wrapped with the grass shrouds and buried on biers of wax myrtle or covered with grape vines. Post molds were also found in association with one of the slough burials; these possibly represent an overlain structure, effigy poles (Purdy 1991, Wentz and Gifford 2007), or burial markers. A retired air force colonel and sport diver, William Royal, discovered both Warm Mineral Springs and Little Salt Springs during the 1950s. Similar to the situation for Warm Mineral Springs, there is little reliable provenience information for the skeletal remains from Little Salt Springs. The remains consist primarily of femurs, reflecting Royal’s preference for collecting exposed remains rather than utilizing systematic excavation techniques (Purdy 1991, Wentz and Gifford 2007). Only six of the Little Salt Springs crania have dental arcades (maxillary or mandibular) suitable for dental crowding analyses.

Figure 3.4: Littletle SSalt Springs site map from Clausen et al. (1979): 610.

Republic Groves (8HR4) Located in east-central HaHardee County just southeast of Zolfo Springs, the Republic Groves site is a multiple componeonent Middle Archaic mortuary pond (Figure 3.5).5). Radiocarbon analyses and projectile point styletyles indicate dates from approximately 5,000-7,00,000 BP (Purdy 1991). The Republic Groves sitete is comprised of the bayhead cemetery and an adjacentad habitation zone. Prior to citrus agriculturalral development in 1968, the area was originallyy swampsw and marshlands with three associateded springs. During drainage canal development humanhu remains

33 were unearthed. Construction was halted and excavation of the site began. Human remains and artifacts were found to be exclusively in the peat layers of the site; the anaerobic conditions of the peat-laden marsh allowed for good preservation of skeletal remains and wooden artifacts. However, disturbance from construction and vegetation resulted in heavy postmortem damage to the human remains (Wharton et al. 1981). A total of 33 wood stakes were recovered from Republic Groves. Most of the stakes were in direct association with burials. The stake tops exhibited battering marks suggesting they were hammered into place (Purdy 1991). Comparative analyses between these Archaic mortuary ponds suggest these stakes were used to both inter the dead and to demarcate the grave. Many of the crania in the Republic Groves collection needed reconstruction. A total of six individuals are included in this study.

Figure 3.5: Location of the Republicic GGroves site in Hardee County, FL, showing area excavatedated. Figure adapted from Purdy (1991): 168.

Harris Creek at Tick Island (8V(8VO24) Harris Creek at Tick Islanland (this study refers to this site simply as Tick Island)Is is a Middle Archaic shell midden siteite in Volusia County, Florida adjacent to the St.t. John’sJ River (Figure 3.6). Graves were dug dirdirectly into the shell and sand of the midden (sandand burial mound) (Russo 1996). Many of these gravraves contained multiple individuals (up to elevenen) (Aten 1999). A total of 149 individuals were exhuxhumed. Isotopic and archaeological analyses showhow these people lived a semi-sedentary existencece cclose to the river. Zooarchaeological and isotopopici analyses by Quinn et al. (2008) infer a primarary subsistence on freshwater and terrestrial resousources, with an emphasis on freshwater foodstuffuffs. Turtle, white-tailed deer, and alligator seemeded to comprise a majority of the fauna consumed.d. HHowever Quinn et al. (2008) suggest that fish andan shellfish are 35 greatly underestimated in the zooooarchaeological analyses due to a lack of screeninning during the initial excavation. As a direct result of shellll midden interment, the Tick Island remains exhibhibit large amounts of taphonomic damagee ((including crushing and concretions). The leechiching of calcium carbonate from shells surroundinging the inhumations resulted in heavy concretionsns. The majority of the skeletons also exhibit crushushing from the weight of the overlying midden sediment.se The Tick Island skeletal sample consinsists of 32 individuals, but the concretions and otheroth taphonomic damage prevented cranial robustisticity and cranial nonmetric analyses of this sampmple.

Figure 3.6: Map of Tick IslaIsland and location of archaeological sites from Aten (1999999): 133.

36 Bay West (8CR200) Bay West is a mortuary pond burial located along the Florida Gulf Coast in Collier County (Figure 3.7). Bay West consists of flexed burials affixed with wooden stakes in a shallow peat laden cypress pond (Beriault et al. 1981). The Bay West population was a Middle Archaic hunter-gatherer society consuming large amounts of marine resources (Quinn et al. 2008). This site was chosen as a comparative sample because its marine subsistence more closely parallels the riverine foraging practiced by the Windover population as compared to terrestrial hunter- gatherer populations. The anaerobic environment of the Bay West burials allowed for good preservation of wooden artifacts and decent preservation of skeletal material (dental remains in particular were well preserved). Wooden stakes similar to those at Windover, Little Salt Springs and Republic Groves were recovered in association with the burials (Beriault et al. 1981). The skeletal remains recovered from Bay West were the result of a salvage excavation after they were discovered during a dredging operation. This expedited excavation was mostly uncontrolled and little to no provenience information or associated grave goods are documented (Purdy 1991, Wentz and Gifford 2007). All crania and isolated dental arcades from this site are included in the sample (n = 10).

1 2

Figure 3.7: Bay West Site Map. Numbers one and two within the mortuary pond (circled in yellow) represent primary locations of skeletal exhumation. Figure adapted from Beriault et al. (1981): 41.

Gauthier (8BR193) Gauthier is a multicomponent site composed of a prehistoric midden and cemetery. The site is located near the Upper Basin of the St. Johns River to the east side of Lake Poinsett in Brevard County (Figure 3.8). It contains a mixed skeletal assemblage from the Middle and Late

38 Archaic Periods. The midden itself is not a shell midden but a sheet midden of bone refuse (Russo 1986). The burials were not interred within the midden but rather were interred along an elevated sandy ridge. In many instances the graves had multiple individuals (mostly two or three) (Jones and Carr 1981). Zooarchaeological studies of Gauthier material show this population was a hunter- gatherer society that subsisted primarily on riverine resources. Fish, duck, and turtles (with fluctuating frequencies throughout the periods) were the most abundant resources recovered from the midden (Russo 1986). Similar to the Bay West excavation, Gauthier was a salvage project that took place over the course of 20 days. The excavation report and analyses were never published. Therefore there is little provenience information for the burials. Materials from the site were also highly mineralized, confounding the radiocarbon dating results. In the absence of radiocarbon dates, associated projectile points were used to determine the dates of the burials (Jones and Carr 1981). The skeletal material (the dentition in particular) from the site is well preserved. Twenty individuals were included in this study.

Figure 3.8: Map of the Upperer BBasin of the St. Johns River. Map highlighting the Gauthiethier site location and assocssociated lakes. Map adapted from Russo (1986): 2.

Methods

Age Estimation Dental and skeletal ageinging techniques included in this study were dentall development,d dental wear severity, and cranial suture closure. To ensure that age estimates arere independent of dental wear, other criteria were alalso used to assess wear. This is particularly truuee for Windover, whose skeletal collection has beeeen extensively analyzed. Dental eruption and formormation was based on Ubelaker’s (1989) sequequence of dental development and eruption amongng Amerindian

40 populations. Dental wear ageing assessments were based on Lovejoy’s (1985) modal tooth-wear patterns of prehistoric Native American populations. Ectocranial suture scores were modeled after Standards for Data Collection for human Skeletal Remains (Buikstra and Ubelaker 1994) cranial suture closure scoring system, which is a composite model of studies by Baker (1984), Mann et al. (1987), Meindl and Lovejoy (1985), and Todd and Lyon (1924, 1925a, 1925b, 1925c).

Age Cohorts To investigate the onset of dental crowding, in particular when crowding can first be observed macroscopically, this study includes subadult remains as well as adult. Age is divided into six categories: Infant, Child, Juvenile, Young Adult, Mid-Adult, and Older Adult (Table 3.2). These age cohorts are based on the categorizations of Scheuer and Black (2000) and the Arizona State Museum Burial Inventory Form. For some analyses the broader groupings of subadult (cohorts 1-3) and adult (cohorts 4-6) were used.

Table 3.2 Age Cohorts Statistical Code Age 1 Infant 0-1 2 Child 1-9 3 Juvenile 10-20 4 Y. Adult 21-34 5 M. Adult 35-49 6 O. Adult 50+

Sex Estimation Florida Archaic peoples were highly sexual dimorphic. Therefore confident sex estimates were made employing the methods in Standards for Data Collection for Human Skeletal Remains (Buikstra and Ubelaker 1994) for cranial features including the nuchal crest, mastoid processes, supraorbital margins, supraorbital ridge/glabella, and mental eminence. Frontal and parietal bossing as well as ascending ramus width and angle were also noted.

41 Sex categories were originally recorded based on five sex cohorts (Table 3.3). For the statistical analyses and to increase sample sizes, these categories were reduced from five ordinal scores to three (see Table 3.3) as ‘probable’ individuals were added to their appropriate cohort.

Table 3.3 Sex Cohorts Score Sex 1 Female 2 Probable female 3 Indeterminate 4 Probably male 5 Male

Subadult skeletal material usually received an indeterminate score for sex estimation since sex-specific characteristics are ambiguous until well after puberty (Patil and Mody 2005). Indeterminate scores were predominately assigned to juveniles. One-quarter of the Windover sample was juveniles.

Dental Crowding Malocclusion encompasses a wide range of dental maladies including rotation, displacement, impaction, ectopic eruption, agenesis, and supernumerary teeth (see Chapter 1). While all the aforementioned aspects of malocclusion were recorded during this study, my research focuses on dental crowding of the anterior teeth as this is the most common form of dental crowding (Alt and Türp 1998b). I define dental crowding as a situation where there are insufficient maxillary or mandibular arch dimensions for erupting dentition. Corruccini and Potter (1980) found that anterior tooth malpositions were primarily the product of tooth-size arch-size discrepancy (TSASD) while posterior tooth malpositions (e.g., premolar rotation) showed greater genetic influence. By focusing on anterior crowding this study investigates malocclusions of the teeth resulting primarily from inadequate arch depth to tooth size ratios; a condition recorded to be greatly influenced by environmental factors (Carlson and Van Gerven 1977, Corruccini 1999, Corruccini and Potter 1980, Harris and Johnson 1991, Harris and Smith 1980, Kaifu et al. 2003, Larsen 1995, Lauc et al. 2003, Price 1936, Rose and Roblee 2009, Weiland et al. 1997). 42

Table 3.4 Dental Terminology and Definitions Type Term Definition Abbreviation Toward the midline of the arcade (toward Mesial M the central incisors) Away from the midline, toward the back of Distal D the arcade Toward the cheek Directional Buccal B Toward the tongue Lingual Li Toward the lips Labial La Toward the palate Palatal - Chewing surface of the tooth Occlusal - Crown surface area between teeth Interproximal - Measurement of maximum tooth width from Metric Mesiodistal MD mesial and distal crown surfaces Measurement of maximum tooth width from Buccolingual BL buccal and lingual crown surfaces Incisors and canines Tooth classes Anterior Teeth - Premolars and molars Posterior Teeth -

Figure 3.9: Directional terms and tooth identifications 43

Little’s (1975) Irregularity Index The Little Irregularity Index (LII) is a dental crowding metric and scoring system found to have a direct correlation with arch depth (Agenter et al. 2009, Janson et al. 2011). This system is the primary method used to record dental crowding severity. The index is a sum of the contact point displacements of the anterior teeth (Figure 3.10). LII only involves the anterior dentition and therefore does not include posterior teeth irregularities such as premolar rotation.

Figure 3.10: Little's (1975) Irregularity Index. Black lines indicate displacement between contact points. Image from Harris and Corruccini (2008): 6.

Contact points are the interproximal surfaces where two teeth come in contact with one another. An ideal contact alignment would result in a measurement of zero. The Irregularity Index is the total of five measurements between the six anterior teeth and includes ten measurements per dentition (five mandibular and five maxillary). These measurements were taken in millimeters using a dental caliper. The sum of these measurements for the mandibular and the maxillary arcades were scored separately and ranked in severity categories (Table 3.5).

Table 3.5 LII scores Score Severity Rank 0 Perfect alignment 1-3 Minimum irregularity 4-6 Moderate irregularity 7-9 Severe irregularity 10 Very severe irregularity

For many of the dental crowding analyses within and between populations, severe through very severe scores were combined for statistical purposes. It is common practice to combine these two categories into a single score (Little 1975). Therefore the aggregate of severe and very severe scores in this study make these frequencies directly comparable to other dental crowding studies. Individuals who exhibited abscess lesions of the anterior teeth and those whose dental crowding could not be assessed due to trauma and/or antemortem tooth loss resulting in alveolar bone resorption were excluded. LII was not assessed on teeth exhibiting little to no enamel (7.5-8 attrition score, see Figure). It was necessary to employ qualitative dental crowding assessments for two individuals from the Tick Island sample. As a result of their interment in a shell midden, the remains have severe post-mortem damage including crushing and concretions. Therefore, LII measurements could not be taken. Qualitative assessment of the interproximal facets were used to gauge the presence and severity of dental crowding. Individuals with observable crowding but could not be measured were compared to other individuals within the same sample who received an LII score. Severity ranks were recoded for these individuals based on these similarities (these individuals were not given LII scores).

Tooth and Arch Dimensions in Relation to Dental Crowding Tooth-size arch-size discrepancies (TSASD) are arguably the most common cause of anterior dental crowding (Alt and Türp 1998b). To assess TSASD measurements of tooth size, arch depth, and arch width were collected.

Dental Metrics Mesiodistal measurements have been found to correlate more with dental crowding than buccolingual dimensions (Bernabé and Flores-Mir 2006 (a), Doris et al. 1981, Peck and Peck 1972). Mesiodistal breadths were recorded for all maxillary and mandibular anterior teeth by measuring the maximum dimension along the mesiodistal plane. This particular technique was chosen, rather than the practice of measuring interproximal contact points, due to the high

45 attrition rates of the Archaic populations under study. Measuring the maximum width of the tooth in these populations allows for more data collection opportunities since hunter-gatherer populations are more prone to interproximal attrition (Wolpoff 1971). Mesiodistal calculated means were compared between the samples. Investigations of tooth size and anterior crowding correlations were conducted by comparing the averages of each sample’s mesiodistal means to their dental crowding frequency. To control for sexual dimorphism tooth width averages were also compared between sexes and occlusal groups (dental crowding and good occlusion groups) within Windover.

Arch Metrics Arch depth (length), width, and shape are crucial developmental factors in the etiology of dental crowding (Beecher and Corruccini 1981, Calcagno 1989, Harris and Smith 1980). To investigate arch size and dental crowding in Florida Archaic populations these following measurements of arch depth and arch width were analyzed. Arch depth and arch width means were compared between the samples. To investigate sex factors on dental crowding etiology within the Windover sample, arch depth and arch width means were compared between sexes and occlusal groups (dental crowding and good occlusion groups).

Arch Depth and Arch Width. Arch depth is the distance from the central contact point between the first incisors (Point C) to the central contact point between the first and second molars (Points A and B) (Harris and Corruccini 2008). Arch depth is calculated as the median height of a triangle formed from three measured cords using these landmarks (Figure 3.11) (DeKock 1972, Harris and Corruccini 2008, Knott 1961) and the formula:

Figure 3.11: Arch depth (D) calculated as median height of a triangle from arch chords AC, BC and AB. Image from Harris and Corruccini (2008): 7.

AC2+ BC 2 AB2 D = − 2 4

In this study arch width is simply the isolated AB chord from the arch depth metrics. This technique was chosen so arch depth and arch width could be directly comparable to each other. Arch depth and width measurements were not taken on partial dental arcades, however estimations were made on complete jaws that exhibited interproximal attrition.

Investigation of Environmental Factors Relating to Dental Crowding Cranial metrics and dental wear methodologies were employed to investigate the possible environmental factors effecting dental crowding. Possible environmental factors analyzed in this study include masticatory adaptations affecting cranial shape, dental attrition relating to masticatory use, and patterns of dental wear resulting from non-masticatory dental uses (extramasticatory use).

Attrition Heavy chewing stress usually results in heavy attrition, thus dental wear severity is a good indicator of the degree of chewing stress and dietary coarseness (Bonfiglioli et al. 2004, Kaifu et al. 2003). Dental attrition of permanent and deciduous teeth was scored as the degree of

47 exposed dentine. Two different scoring systems were used to record wear: Smith’s (1984) dental surface wear scoring system and Scott’s (1979) quadrant scoring system. The Smith scoring system was used to assess wear on incisors, canines and premolars while the Scott scoring system was used for molar wear assessment. Smith’s scoring system is an eight-point scale based on dentin exposure (Figure 3.12). Teeth were matched to the drawings below as well as to descriptions of the attrition stages. Stage one represents no wear (no dentin exposure), while Stage eight represents severe wear with no enamel present where the tooth takes on the shape of the roots (total dentin exposure).

A. B.

Figure 3.12: Dental Wear Scoring Systems. A.) Smith (1984): 45-46 dental surface wear scoring system. B.) Scott (1979): 214 molar surface wear scoring system. Images from Buikstra and Ubelaker (1994): 52-53.

The Scott (1979) scoring system was used specifically for molars. In this system each occlusal surface is divided into quadrants. The amount of enamel present is scored on a scale of zero to ten. The final score per tooth is a sum of all four quadrants. Each tooth can therefore yield a possible score from zero through forty. A score of zero represents no wear, while a score

48 of ten represents complete absence of enamel on any part of the quadrant of the molar (See Figure 3.12). For each individual the sums of the anterior and posterior teeth scores were calculated for the maxillary and mandibular dentition separately. The total maxillary scores and mandibular scores (adult and subadult) per sample were analyzed separately. The mean attrition scores for each jaw, standardized by age, were analyzed between samples to investigate wear severity differences and possible correlations between attrition and dental crowding.

Dental Wear Patterning Many non-masticatory behaviors practiced over long periods of time can be identified in unique dental wear patterns (Scott and Turner 1988). It has been suggested that non-masticatory and habitual practices can lead to anterior dental crowding (Oppenheimer 1964, Pereira and Evan 1975, Singh 2007). Wear patterns were recorded to investigate the relationship between dental crowding and the cultural practice of using teeth as tools. In particular, grooved wear patterns were described, recorded and analyzed. Patterns observed in this study were interproximal grooving, occlusal groove wear, and a previously undocumented pattern described in this study as “lingual root wear”. The frequencies of these patterns were analyzed between population samples. The spatial distribution of dental wear patterns within the Windover site (based on burial plots) was compared to the distribution of dental crowding.

Cranial Metrics Cranial shape and form, though governed by many genetic factors, is highly influenced by exogenous forces (Brown and Maeda 2004, Carlson and Van Gerven 1977, Corruccini 1999, Larsen 1995, Rose and Roblee 2009). In particular, subsistence and diet variability relate to masticatory stresses that are known to influence cranial robusticity (see Chapter 2). Low masticatory stress has been found to correlate with dental crowding (Corruccini 1999, Price 1936, Rose and Roblee 2009). Therefore, cranial metrics were employed to investigate the relationship between craniofacial size and dental crowding in the Florida Archaic populations under study.

49 The craniofacial measurements, listed in Table 3.6, were conducted following the methodology of Buikstra and Ubelaker (1994) and Moore-Jansen et al. (1994). To control for developmental differences measurements were made on adult crania only. Individuals with gross pathologies were excluded. Crania are more susceptible to taphonomic damage than dentition due to the durability of enamel (White et al. 2011). Therefore, to address statistical issues caused by incomplete data Missing Value Analyses were conducted on all cranial metric data for each sample. Measurements with missing data greater than 50 percent were excluded from further analyses. Mann-Whitney U nonparametric tests of significance were then conducted comparing each sample to Windover.

Table 3.6 Craniometric (CM) Abbreviations Maximum Cranial Length (MXCL) Interorbital Breadth (IB) Maximum Cranial Breadth (MXCB) Frontal Chord (FC) Bizygomatic Diameter (BD) Parietal Chord (PC) Basion-Bregma Height (BBH) Occipital Chord (OC) Cranial Base Length (CBL) Foramen Magnum Length (FML) Basion-Prosthion Length (BPL) Foramen Magnum Breadth (FMB) Maxillo-Alveolar Breadth (MAB) Mastoid Length (MastL) Maxillo-Alveolar Length (MAB) Chin Height (CM) Biauricular Breadth (BiaBr) Height of Mandibular Body (HMB) Upper Facial Height (UFH) Breadth of Mandibular Body (BMB) Minimum Frontal Breadth (MFB) Bigonial Width (BW) Upper Facial Breadth (UFB) Bicondylar Breadth (BB) Nasal Height (NH) Minimum Ramus Breadth (MRB) Nasal Breadth (NB) Maximum Ramus Breadth (MXRB) Orbital Height (OH) Maximum Ramus Height (MXRH) Orbital Breadth (OB) Mandibular Length (MandL) Biorbital Breadth (BioBr) Mandibular Angle (MA)

50 Investigation of Genetic Relatedness Using Nonmetric Traits Nonmetric traits have been shown to possess a high genetic component in expression (Larsen 1999, Rightmire 1999, Scott and Turner 1997) and are commonly used to test genetic relatedness within and between populations (Buikstra and Ubelaker 1994, Irish 2005, Saunders and Popovich 1978, Scott and Turner 1997, Stojanowski and Schillaci 2006). Therefore nonmetric traits were used to investigate genetic relatedness within and between the samples to evaluate how these patterns correlate with dental crowding. This study employs the dental nonmetric Arizona State University Dental Anthropology System and scores cranial nonmetirc traits using Standards (Buikstra and Ubelaker 1994) to test for significant differences in nonmetric frequencies between samples. Dental crowding frequencies were then analyzed in relation to nonmetric significance levels to investigate possible genetic factors in dental crowding etiology.

Dental Nonmetrics Nonmetric traits were scored on all teeth (R and L) in the samples under study. Arizona State University Dental Anthropology System (ASUDAS) standard ranges were used to record 34 traits and their variance. Once all antimeres were scored, the highest expression per individual was used to determine prevalence in the sample (Turner and Scott 1977). Like many dental and cranial nonmetric trait studies, this project converts the traits to a presence/absence score for further statistical analyses (Table 3.7). Defining a nonmetric trait as present or absent essentially divides a continuous distribution into two discontinuous pieces (Fraser 1998). Ranges used to convert the dental nonmetric ordinal scores into presence/absence scores were based on Irish (2005). Converting these traits to presence/absence scoring makes these frequencies directly comparable to other dental nonmetric studies.

Table 3.7 Dental Nonmetric Traits and Presence Ranges Trait Range for Trait Trait Range for Trait Scored as Present Scored as Present Bilateral winging UI1 + = ASUDAS 1 Carabelli’s cusp UdM2 + = ASUDAS 2-7 Shoveling UI1 + = ASUDAS 2-6 Enamel extensions UM1 + = ASUDAS 1-3 Double Shoveling UI1 + = ASUDAS 2-6 Enamel extensions UM2 + = ASUDAS 1-3 Peg-reduced UI2 + = ASUDAS P or A Enamel extensions LM1 + = ASUDAS 1-3 Tuberculum dentale UI1 + = ASUDAS 2-6 Enamel extensions LM2 + = ASUDAS 1-3 Tuberculum dentale UI2 + = ASUDAS 2-6 Parastyle UM1 + = ASUDAS 1-5 Tuberculum dentale UC + = ASUDAS 2-6 Parastyle UM2 + = ASUDAS 1-5 Tuberculum dentale LC + = ASUDAS 2-6 Deflecting wrinkle UM1 + = ASUDAS 2-3 Distal accessory ridge UC + = ASUDAS 2-5 Deflecting wrinkle LM1 + = ASUDAS 2-3 Distal accessory ridge LC + = ASUDAS 2-5 Deflecting wrinkle Ldm2 + = ASUDAS 2-3 UtoAztecan Premolar UP3 + = ASUDAS P or A Cusp 7 LM1 + = ASUDAS 2-4 C3 UM1 + = ASUDAS 2-5 Cusp 7 LM2 + = ASUDAS 2-4 C3 UM2 + = ASUDAS 2-5 Protostylid LM1 + = ASUDAS 2-6 C5 UM1 + = ASUDAS 2-5 Protostylid LM2 + = ASUDAS 2-6 C5 UM2 + = ASUDAS 2-5 Anterior fovea LM1 + = ASUDAS 2-4 Carabelli’s cusp UM1 + = ASUDAS 2-7 Anterior fovea LM2 + = ASUDAS 2-4 Carabelli’s cusp UM2 + = ASUDAS 2-7 Anterior fovea Ldm2 + = ASUDAS 2-4 Plus signs represent presence of a trait; capital U and L indicate upper (maxillary) and lower (mandibular) dentition; a lower case d differentiates deciduous dentition

Cranial Nonmetric Traits Cranial nonmetric traits were scored using the Standards (Buikstra and Ubelaker 1994) scoring system, which is based primarily on studies by Falconer (1965, 1967) and Hauser and De Stefano (1989). For the statistical analyses, the scores were converted to presence/absence scores (Table 3.8). In order to convert cranial nonmetric traits to dichotomous variables, the maximum expression of the trait for bilateral traits was used, while continuous categorical traits were rescored as dichotomous variables by converting and combining all non-absence categories to a present group (Carson 2006). Previous studies were used as a basis for certain expression rescoring. For instance, incomplete metopic sutures were considered absent, and only complete metopic sutures were considered to be present following Berry and Berry (1967). Hauser and De Stefano (1989) determined the important feature of the condylar canal was its patency, therefore in this study condylar canals were considered to be present only if patent. Divided hypoglossal canals, 52 however, were considered to be present if a partial to complete division within the canal was observable. The Foramen ovale and foramen spinosum traits were recorded based on incompleteness of the foramen (Buikstra and Ubelaker 1994); therefore, these traits were considered present only if there was incomplete or a lack of bone formation around the foramen. A fully enclosed foramen (a foramen with complete definition) was considered to be an absence of that trait. Inca bones were considered present whether they were singular, bipartite or tripartite (Perizonious 1979). If bipartite or tripartite manifestations were observed their presence was noted. Because few crania in this study exhibited complete absence of mastoid and mental foramen these traits were redefined as a measure of additional foramina; this redefinition is based on Carson (2006) who, in his study, redefined the lesser palatine, mastoid, and zygomatico-facial foramina scores in this same way. The superior sagittal sulcus flexure, mylohyoid bridge location, and mastoid foramen location could not be parsed as present/absent and were left as multilevel (Carson 2006); the frequency of each trait expression was examined separately. Subadults were included in cranial nonmetric analyses as many of the traits manifest early in development. However, the tympanic dehiscence was excluded from subadult assessments aged 5 years or younger since this trait occurs as a developmental trait in children and can only be discerned as a nonmetric variant in those over the age of five (Lacout et al. 2005).

Table 3.8 Cranial Nonmetric Traits and Their Presence Ranges Trait Range for Trait Trait Range for Trait Scored as Present Scored as Present Metopic Suture + = B&U 2 L. Flexure Superior Sagittal Suture + = B&U 2 Supraorbital Notch + = B&U 1-4 Bi. Flexure Superior Sagittal Suture + = B&U 3 Supraorbital Foramen + = B&U 1-2 Foramen Ovale Incomplete + = B&U 1-2 Infraorbital Suture + = B&U 1-2 Foramen Spinosum Incomplete + = B&U 1-2 Multiple Infraorbital Foramina + = B&U 1-3 Pterygo-spinous Bridge + = B&U 1-3 Zygomatico-facial Foramina + = B&U 1-6 Pterygo-alar Bridge + = B&U 1-3 Parietal Foramen + = B&U 1-2 Tympanic Dehiscence (+ = B&U 1-2) Epiteric Bone + = B&U 1 Auditory Exostosis (+ = B&U 1-3) Coronal Ossicle + = B&U 1 Temporal Mastoid Foramen (+ = B&U 1) Bregmatic Bone + = B&U 1 Sutural Mastoid Foramen (+ = B&U 2) Sagittal Ossicle + = B&U 1 Occipital Mastoid Foramen (+ = B&U 3) Apical Bone + = B&U 1 Sutural & Temporal Mastoid Foramen (+ = B&U 4) Lambdoid Ossicle + = B&U 1 Occipital & Temporal Mastoid Foramen (+ = B&U 5) Asterionic Bone + = B&U 1 Mental Foramen (+ = B&U 2-3) Occipitomastoid Ossicle + = B&U 1 Mandibular Torus (+ = B&U 1-3) Parietal Notch Bone + = B&U 1 Mylohyoid Bridge Mand. Foramen (+ = B&U 1) Inca Bone + = B&U 1-4 Mylohyoid Bridge Center of Groove (+ = B&U 2) Condylar Canal + = B&U 1 Double Mylohyoid Bridge Hiatus (+ = B&U 3) Divided Hypoglossal Canal + = B&U 1-4 Double Mylohyoid Bridge No Hiatus (+ = B&U 4) R. Flexure Superior Sagittal Suture + = B&U 1

Statistical Analyses A Fisher’s Exact test was used to compute the exact P-value for analyses of discrete/nominal variables (e.g., ILL severity analyses, nonmetric frequencies, sex and age frequencies). A Fisher’s Exact test was chosen due to expected small sample sizes for the aforementioned tests. P-values from Fisher’s Exact tests are considered more accurate than chi- squared tests (including the application of Yate’s Correction) or G-tests if the expected numbers are small (McDonald 2009). Contingency tables (2x2) were employed to test for differences and similarities between the samples and two-tailed P-values were used to determine the significance of the results; p < 0.05 indicated statistical significance.

54 Mann-Whitney U tests (nonparametric tests of significance) were employed to analyze differences of continuous variables (e.g., cranial metrics, dental metrics, arch depth, and arch width) between two samples. In cases where more than two samples were being compared for significant differences, a Kruskal-Wallis nonparametric analysis of variance was utilized to compare sample distributions. If test results exhibited significant differences then Mann-Whitney U tests were utilized to analyze specific sample pairs. Comparisons of means and standard deviations were used to investigate differences between Windover and the comparative samples, and for analyses within Windover for attrition assessments (e.g., arch depth, arch width, mesiodistal tooth widths, and cranial metrics). The Statistical Package for Social Sciences (SPSS Inc., Chicago, Illinois, USA) version 19.0.1 was used to run the descriptive statistics, Mann-Whitney U nonparametric analyses, Kruskal-Wallis non-parametric analysis of variance, and the Missing Values analyses. Fisher’s Exact tests, 2x2 contingency tables were calculated using Graphpad Software.

55 CHAPTER 4 WINDOVER ANALYSES

This chapter presents the results for analyses conducted on the Windover sample. Measurements of dental crowding were employed to investigate the prevalence and distribution of malocclusion at Windover. Dental crowding has a multifactorial etiology. To assess the primary influences on dental crowding development in the Windover sample, analyses of cranial and jaw shape, tooth size, and dental alterations (masticatory and non-masticatory) were examined.

Dental Crowding Analyses The Little Irregularity Index (LII) was used to assess dental crowding severity. The Windover sample exhibits a total dental crowding incidence of 47 percent. Severe crowding accounts for over half of the crowding total (25 percent), while there is a higher frequency of mild crowding (13 percent) over moderate crowding (9 percent). Figures 4.1-4.2 represent two cases of severe dental crowding in the Windover sample.

Dental Crowding Frequencies By Age Cohorts The Windover sample has a good representation of most age groups, ranging from children through older adults. Dental crowding frequencies were investigated for age cohorts to identify correlations between age crowding (Table 4.1). As was expected, because there is limited dental development in infants (aged 0-1 year), dental crowding was not detected. Of the 47 percent total dental crowding frequency in Windover, young adults comprise the majority of dental crowding (17 percent), followed closely by juveniles (13 percent). Young adults also had the highest rate of severe dental crowding (10 percent). Juveniles exhibited the next highest severe crowding rate (8 percent). Crowding rates increased with age from the child (4 percent) through young adult groups (17 percent) but mild and moderate crowding scores were more similar between all age groups, ranging from 0-4 percent. Dental crowding frequencies declined with age in the adult groups from the high in the young adults down to 7 percent in the older adults.

Figure 4.1: Severe dental crowding case in Windover individual (Burial 99). Occlusal view (left), anterior view (right). Note the unique dental morphology of the left lateral incisor.

Figure 4.2: Severe dental crowding case in Windover individual (Burial 86). Occlusal view (left), anterior view (right). Note the retention of the right second deciduous molar as well as premolar rotations and displacements.

Dental crowding was also analyzed within each age cohort (Table 4.1). A staggering 83 percent of juveniles exhibit dental crowding, which is comprised mostly of severe crowding (58 percent). In the young adult group 75 percent have dental crowding and 45 percent of that is severe. Older adults show a 33 percent rate of dental crowding with the majority of that expressed as mild crowding (17 percent). Children have a 30 percent crowding rate with an equal distribution of moderate and severe crowding (15 percent). Middle-aged adults have the lowest

57 rate of dental crowding (28 percent), with the majority of mild crowding cases (16 percent). Interestingly, severe crowding frequencies reduce dramatically from the young adult group (45 percent) to the middle-aged adult group (8 percent).

Table 4.1 Windover Dental Crowding Within Age Cohorts Good Total Mild Moderate Severe Site Sample Occlusion Crowding Crowding Crowding Crowding Child % 70% 30% 0 15% 15% (1-9yrs) n = 13 # 9 4 0 2 2 Juvenile % 17% 83% 17% 8% 58% (10-20yrs) n = 12 # 2 10 2 1 7 Y. Adult % 25% 75% 15% 15% 45% (21-34yrs) n = 20 # 5 15 3 3 9 M. Adult % 82% 28% 16% 4% 8% (35-49yrs) n = 25 # 18 7 4 1 2 O. Adult % 67% 33% 17% 5% 11% (50+ yrs) n = 18 # 12 6 3 1 2 The # symbolizes the number of dental crowding cases. Percentages are rounded to the nearest tenth of a percent.

The assessments of dental crowding frequency within and between age cohorts show a trend of reduced dental crowding with age in the adult groups as well as increased crowding frequencies with age in the child through young adult groups. These trends are a product of three factors: the initial onset of dental crowding within this sample, the reduction of dental crowding with age due to increased attrition and antemortem tooth loss, and improved tooth alignment due to heavy interproximal and occlusal attrition. Attrition increases with age, therefore the lower rates of dental crowding in the middle and older adult cohorts are most probably a combination of dental crowding assessment complications (due to tooth surface reduction from tooth wear and antemortem tooth loss), and/or because of the interproximal space created by heavy attrition that allows teeth (via mesial drift and lingual tipping) to shift into better alignment (Begg 1954, Wolpoff 1971). Therefore the high rates of crowding seen in the juvenile and young adult groups

58 in this sample are a reflection of dental crowding unaltered by attrition and other age-related confounders.

Onset of Dental Crowding The Windover sample has the largest (n = 26) and most diverse representation of subadults in this study, with individuals ranging from infancy to adolescence. Given limited dental eruption in the youngest individuals, only child (n = 13) and juvenile (n = 12) cohorts were assessed for dental crowding. Of the 25 individuals in this subadult sample, 14 have visible dental crowding. Severe crowding constitutes the majority of the subadult dental crowding cases (n = 9). Mild (n = 2) and moderate (n = 3) crowding cases comprise less than half of the total dental crowding in the subadult group. The youngest individual who exhibits observable anterior dental crowding in the Windover sample is a child aged 7-8 years. The next youngest subadults exhibiting crowding are two individuals aged 8-9 years. There are a total of eight individuals under the age of seven in this sample and none of them exhibited dental crowding. Unlike the anterior crowding recorded in a modern sample of 3-4 year-olds from India (Prabhakar et al. 2008), crowding was not recorded in individuals under the age of seven in this sample and therefore is only found in individuals with mixed dentition (having a combination of deciduous and permanent teeth). Crowding appears to manifest only with the eruption of the anterior permanent dentition and does not affect the deciduous dental arcade in the Windover subadults. The sample of individuals younger than seven years old was small (n = 8). Therefore cases of dental crowding may be present in subadults younger than seven years but crowding is not observable due to small sample size. However, dental crowding rates were high in the children aged seven years and older. In the child cohort (n = 13, including those under seven), five were seven years and older and four of them (80 percent) exhibit crowding. This high rate of crowding is also true of the juvenile cohort. Within the juvenile group, ten individuals out of 12 exhibit dental crowding and seven of those individuals have severe crowding. These data support the observation that dental crowding at Windover is a result of permanent tooth eruption.

59 Dental Crowding Frequencies Between Sex Cohorts Both sexes are well represented in the Windover sample (female = 29, male = 34). Sex was estimated for all adult crania. Individuals with indeterminate sex scores belong solely to the subadult cohorts and also comprise a large portion of the sample (n = 26). Table 4.2 presents the distribution of crowding cases by sex cohorts for the entire sample. For example, females with crowding represent 13 of the total 89 individuals, or approximately 15 percent.

Table 4.2 Windover Dental Crowding Between Sex Cohorts Good Total Mild Moderate Severe Sex Cohort Sample Occlusion Crowding Crowding Crowding Crowding Female % 85% 15% 5% 5% 6% n = 29 # 16 13 4 4 5 Male % 84% 16% 7% 1% 8% n = 34 # 20 14 6 1 7 The # symbolizes the number of cases of dental crowding. Percentages are rounded.

Slight differences were observed in dental crowding distribution between the sexes. Males comprised the majority of the dental crowding cases (16 percent) in the Windover sample, followed closely by the females (15 percent). Female adults exhibit a higher rate of moderate dental crowding (5 percent) and male adults exhibited higher rates of mild (7 percent) and severe (8 percent) dental crowding. The largest difference in dental crowding rates between the sexes is a 4 percent difference within the moderate crowding category. Comparing the distribution within the 42 crowding cases, 13 females constitute 31 percent of the total cases, and 14 males constitute 33 percent; this illustrates that crowding is roughly evenly distributed among males, females and subadults. Four females and six males have mild dental crowding. Another four females and one male exhibit moderate crowding. Lastly, five females and seven males exhibit severe crowding; these represent approximately 23 percent and 32 percent respectively of the severe crowding cases with the remaining 45 percent made up of the subadults. Among the adult males, 50 percent of those with crowding have severe crowding, while only 39 percent of the females with crowding have severe expressions of the condition. This

60 indicates that size of the dentition may be a factor in dental crowding development (see tooth size section).

Spatial Distribution of Crowding at Windover There are four excavation sites within the Windover pond. The archaeological investigations were initially carried out in discontinuous pond subsections (northwest, northeast, southwest, and southeast) (Dickel 2002). The southeast subsection was sterile of burials, yielding only three isolated skeletal elements (Dickel 2002). The northwest, northeast and southwest subsections can be compared using scatterplots. As can be observed in Figure 4.3 the northeast subsection contained the greatest burial density. The location of individuals with dental crowding was plotted with the north and east axes are burial coordinates. The scatterplot shows a broad distribution of dental crowding throughout Windover pond; it occurs in all three subsections.

NW Subsection

NE Subsection

SW Subsection

Figure 4.3: Dental crowding plotted for Windover pond. North and east axes are burial coordinates measured in meters. Circles indicate absence of dental crowding; squares indicate presence of dental crowding.

Spatial Distribution of Windover Dental Crowding Severity Ranks To investigate dental crowding severity throughout Windover the individual severity levels were plotted (Figure 4.4). There are no obvious patterns for levels of dental crowding in the northwest and southwest quadrants of the pond. However, the small samples sizes for these subsections compromise the distribution analyses. In the northeast subsection, the area with the greatest burial density, severe crowding cases cluster in the eastern portion of the section.

62 In their intra-cemetery biodistance analysis of Windover, Stojanowski and Schillaci (2006) hypothesized a subpopulation divide within the northeast subsection of the site (see line in Figure 4.4); dental crowding was not a component of their analyses. Interestingly, severe dental crowding clusters according to this hypothesized division. Between the east and west portions severe dental crowding show highly disparate levels (Table 4.3). The east exhibits a severe crowding frequency of 67 percent, more than three times the severe crowding rate in the west (14 percent).

Table 4.3 Windover Dental Crowding Between East and West Portions in NE Pond Subsection Good Total Mild Moderate Severe Sample Occlusion Crowding Crowding Crowding Crowding East Division % 32% 68% 6% 0 67% n = 19 # 6 13 1 0 12 West Division % 60% 40% 10% 14% 14% n = 48 # 29 19 5 7 7 The # symbolizes the number of cases of dental crowding. Percentages are rounded.

Two-tailed Fisher’s Exact tests were conducted on the dental crowding frequencies in the east and west portions of the NE subsection. There was a highly significant difference in the severe crowding frequency between the two sections (p = 0.0002). However tests of significance for total crowding (p = 0.056), mild (p = 0.1679), and moderate (p = 1.00) crowding scores exhibit non-significant differences between the east and west NE subsection.

Figure 4.4: Scatter plot of crowding severity in Windover pond. Circles indicate no dental crowding, squares represent mild crowding, triangles represent moderate crowding and diamonds represent severe crowding. The dividing line shows the hypothesized division of the NE subsection into two groups.

64 To account for possible age factors in the analyses of these dental crowding cluster patterns, age cohorts were plotted to show the age distribution within the cemetery (Figure 4.5). Overall there was a higher rate of subadults and young adults in the eastern portion and a higher rate of middle-aged and older adults in the western division (Table 4.6). In particular, five juveniles and six young adults constitute 50 percent of the east division’s sample size collectively. In the eastern portion, 32 percent of the sample consists of five older adults and two middle-aged adults. The remaining 18 percent of the burials are children, all under the age of six. In the western division 15 middle-aged adults and 16 older adults comprised 60 percent of the total sample in the western division; six juveniles and eight young adults constituted 28 percent of the western portion demographics. The remaining 12 percent consist of six children, two of which are under the age of seven.

Table 4.4 Windover Demographics for East and West Portions in the NE Pond Subsection

Sample Child Juvenile Y. Adult M. Adult O. Adult

East Division % 18% 23% 27% 9% 23% n = 22 # 4 5 6 2 5 West Division % 12% 12% 16% 29% 31% n = 51 # 6 6 8 15 16 The # symbolizes the number of cases of dental crowding. Percentages are rounded.

In the previous age cohort dental crowding analysis, 58 percent of all juveniles and 43 percent of young adults exhibited severe dental crowding. In the child cohort 40 percent of all children over the age of seven showed severe crowding. In contrast, only 8 percent of middle- aged adults and 11 percent of older adults exhibited severe crowding. Therefore, demographic factors may be influencing differential severe crowding ratios between the east and west portions of the northeast subsection. Fisher’s Exact tests were conducted for age frequencies between the east and west divide. Each of the five age cohorts exhibited non-significant differences between the east and west- child (p = 0.4763), juvenile (p = 0.2889), young adult (p = 0.3321), middle- aged adult (p = 0.0744), and older adult (p = 0.5774), To mitigate complications from small sample sizes in the middle-aged adult group (east division has only two individuals from this age

65 cohort), aggregates based on severe dental crowding frequencies were analyzed; juveniles and young adults were combined because of there comparable high severe dental crowding frequency, while middle-aged adults and older adults were combined based on their similar low severe crowding frequencies. Fisher’s Exact tests were conducted for these two aggregates between the east and west portions. The juvenile/young adult group exhibited non-significant differences between the east and west (p = 0.1053), however the middle-aged/older adult group showed significant differences between the portions (p = 0.0399). Therefore it appears that age is a confounding factor in the significantly higher severe dental crowding frequencies from the east portion of the northeast subsection.

Figure 4.5: Age cohorts plotted in Windover pond. Stars indicate infants, circles represent children, squares represent juveniles, triangles represent young adults, diamonds represent middle-aged adults, and pentagons represent older adults.

Windover Tooth Size and Arch Size Relationships to Dental Crowding Environmental influences on the cranium that reduce chewing stresses may result in the underdevelopment of the maxillae and mandible, commonly leading to inadequate jaw space for 67 genetically determined tooth dimensions (Calcagno 1989, Carlson and Van Gerven 1977, Greene 1970). Tooth size, arch depth and arch width were analyzed to investigate the role of tooth-size arch-size discrepancy (TSASD) in dental crowding at Windover.

Windover Tooth Size Analyses Mesiodistal tooth widths have been documented as having a direct correlation with dental crowding (Bernabé and Flores-Mir 2006 (a), Doris et al. 1981, Peck and Peck 1972). In this study dental crowding was shown to only affect permanent dentition (see Onset of Dental Crowding section above). Therefore only permanent teeth were compared. Individuals exhibiting marked interproximal attrition were excluded from the analysis.

Windover MD Metrics Between Sex Cohorts Windover males and females have similar levels of dental crowding, but males have proportionally more severe crowding. To investigate how tooth size contributes to this pattern, MD measurements were analyzed by sex cohort. As discussed previously sex was only determined for adults, therefore subadults were not included in this analysis. Overall, the male MD widths exhibit slightly higher values than the females. The exceptions, showing higher means for females, were the maxillary third premolars (P3), mandibular incisors (I1 and I2), and mandibular premolars (P3 and P4) (Table 4.5 and Table 4.6). The greatest difference between the sexes is in the maxillary anterior dentition, particularly between the lateral incisors (0.52 mm difference) and the canines (0.5 mm difference), and the mandibular lateral incisors (0.74 mm difference). Despite the large degrees of skeletal sexual dimorphism in the Windover sample, the maxillary posterior tooth sizes (Figures 4.6 and 4.8) show minor differences and the mandibular posterior teeth are only slightly more variable in size (Figures 4.7 and 4.9). Mann-Whitney U tests of significance for MD tooth widths were conducted between the sex cohorts in the Windover sample (Tables 4.7 - 4.8). Only the mandibular lateral incisor (p = 0.002) and the maxillary canine (p = 0.033) were significantly different between the two groups. The differences in the male and female canines are to be expected, as these are typically the most dimorphic teeth in sexually dimorphic primates. The mandibular lateral incisor shows the largest mean difference between the sexes (0.74 mm).

68 Interestingly, this tooth was exceptional in that the mean value was larger in the female group than the males. Despite the visible differences in maxillary later incisor MD means between males and females (Figure 4.6), I2 tooth size is not significantly different between the sexes due to the high variability in the female sample and the low sample sizes for the males. The differences in the other anterior teeth, combined with the high levels of variation (reflected in the high standard deviations) for many of the teeth, may indicate that the size of specific teeth, rather than the dentition in general, is contributing to dental crowding at Windover. The following analysis addresses that issue.

8.4 8.2 8 7.8 7.6

7.4 Female 7.2 Male MD Means MD Means (mm) 7 6.8 6.6 I1 I2 C Maxillary Anterior Teeth

Figure 4.6: Windover maxillary mesiodistal means for anterior teeth.

69 12

6 Female 4 Male MD Means MD Means (mm) 2

0 P3 P4 M1 M2 M3 Maxillary Posterior Teeth

Figure 4.7: Windover maxillary mesiodistal means for posterior teeth.

8 7 6 5 4 Female 3 Male MD Means MD Means (mm) 2 1 0 I1 I2 C Mandibular Anterior Teeth

Figure 4.8: Windover mandibular mesiodistal means for anterior teeth.

70 14

6 Female Male MD Means MD Means (mm) 4

0 P3 P4 M1 M2 M3 Mandibular Posterior Teeth

Figure 4.9: Windover mandibular mesiodistal means for posterior teeth.

Table 4.5 Windover Maxillary MD Descriptive Statistics by Sex Cohorts Female Male Tooth Type N Mean Std. Deviation N Mean Std. Deviation I1 12 7.97 1.311 5 8 0.707 I2 15 7.2 1.014 10 7.7 0.675 C 19 7.79 0.631 13 8.31 0.48 P3 19 6.84 0.688 15 6.73 0.884 P4 19 6.47 0.513 20 6.7 0.733 M1 20 10.45 0.826 18 10.61 0.85 M2 23 9.83 0.937 20 10.15 0.745 M3 22 9.5 1.225 22 9.59 0.959

Table 4.6 Windover Mandibular MD Descriptive Statistics by Sex Cohorts Female Male Tooth Type N Mean Std. Deviation N Mean Std. Deviation I1 16 5.38 0.619 7 5.29 0.756 I2 18 6.17 0.618 7 5.43 0.535 C 16 6.81 0.911 13 7 0.408 P3 15 7.07 0.594 15 6.8 0.414 P4 14 7.07 0.829 15 6.87 0.516 M1 17 11.47 0.624 12 11.75 0.622 M2 20 11 1.076 19 11.37 0.955 M3 18 10.89 0.9 23 11.35 0.982

Table. Table 4.7 Windover Maxillary MD Metrics: Mann-Whitney U Test Results Between Sex Comparisons Dental MD Metric P-value, Two-Tailed I1 0.657 I2 0.212 C 0.033 P3 0.736 P4 0.238 M1 0.594 M2 0.226 M3 0.481 Bold entries are statistically significant.

Table. Table 4.8 Windover Mandibular MD Metrics: Mann-Whitney U Test Results Between Sex Comparisons Dental MD Metric P-value, Two-Tailed I1 0.824 I2 0.002 C 0.536 P3 0.172 P4 0.438 M1 0.206 M2 0.210 M3 0.125 Bold entries are statistically significant.

Windover Maxillary MD Metrics Between Occlusion Cohorts Variation in tooth size and its relationship to dental crowding was evaluated by stratifying the sample by crowding levels. These comparisons include subadults as well as adults. From Figures 4.10 - 4.11, it appears that the mean dimensions of the maxillary lateral incisors are smaller in individuals with good occlusion, and overall individuals with crowding appear to have larger teeth. For the maxilla, the central incisors show little difference between the dental crowding and good occlusion groups, while the lateral incisors exhibit the greatest difference. Again, most of these variables are characterized by high degrees of variation (Tables 4.9-4.10). Mann-Whitney U tests were conducted to test whether the teeth of individuals with crowding are significantly larger (Table 4.11). All teeth exhibit non-significant differences

72 between the occlusion cohorts. Therefore, despite slightly larger means in the dental crowding group, maxillary tooth width does not correlate with dental crowding in this sample.

8.4 8.2 8 7.8 7.6

7.4 Good Occlusion 7.2 Dental Crowding MD Means MD Means (mm) 7 6.8 6.6 I1 I2 C Maxillary Anterior Teeth

Figure 4.10: Windover maxillary mesiodistal means for anterior permanent teeth in individuals with good occlusion and dental crowding.

6 Good Occlusion 4 Dental Crowding MD Means MD Means (mm)

0 P3 P4 M1 M2 M3 Maxillary Posterior Teeth

Figure 4.11: Windover maxillary mesiodistal means for posterior permanent teeth in individuals with good occlusion and dental crowding.

Table 4.9 Windover Maxillary MD Descriptive Statistics of Permanent Anterior Teeth with Good Occlusion Tooth N Mean Std. Deviation I1 8 8.25 0.707 I2 11 7.27 1.104 C 10 8.10 0.568 P3 11 6.82 0.874 P4 14 6.57 0.514 M1 18 10.56 0.922 M2 18 9.94 0.725 M3 20 9.70 1.174

Table 4.10 Windover Maxillary MD Descriptive Statistics of Permanent Anterior Teeth with Dental Crowding Tooth N Mean Std. Deviation I1 21 8.24 1.179 I2 25 7.52 0.714 C 30 8.03 0.669 P3 31 6.97 0.795 P4 33 6.79 0.740 M1 37 10.76 0.723 M2 35 10.09 0.919 M3 25 9.24 1.332

Table. Table 4.11 Windover Maxillary MD Metrics: Mann-Whitney U Test Results Between Occlusion Cohorts Dental MD Metric P-value, Two-Tailed I1 0.675 I2 0.588 C 0.865 P3 0.532 P4 0.313 M1 0.390 M2 0.507 M3 0.260 Bold entries are statistically significant.

74 Windover Mandibular MD Metrics Between Occlusion Cohorts Similarly, with regard to the mandibular dentition, most of the means were higher for the crowding group (Tables 4.12-4.13 and Figures 4.12-4.13). Mann-Whitney U tests of significance for these variables produced different results (Table 4.14): interestingly, the central (p = 0.002) and lateral incisors (p = 0.005) show highly significant difference. This indicates a correlation between mandibular incisor MD width and dental crowding in the Windover sample. This further supports the idea that the size of specific teeth, rather than the dentition in general, is contributing to dental crowding at Windover.

8 7 6 5 4 Good Occlusion 3 Dental Crowding MD Means MD Means (mm) 2 1 0 I1 I2 C Mandibular Anterior Teeth

Figure 4.12: Windover mandibular mesiodistal means for anterior permanent teeth in individuals with good occlusion and dental crowding.

75 14 12 10 8

6 Good Occlusion 4 Dental Crowding MD Means MD Means (mm) 2 0 P3 P4 M1 M2 M3 Mandibular Posterior Teeth

Figure 4.13: Windover mandibular mesiodistal means for posterior permanent teeth in individuals with good occlusion and dental crowding.

Table 4.12 Windover Mandibular MD Descriptive Statistics of Permanent Teeth with Good Occlusion Tooth N Mean Std. Deviation I1 6 4.67 0.516 I2 7 5.43 0.535 C 8 6.62 0.518 P3 10 6.8 0.422 P4 10 6.90 0.568 M1 10 11.5 0.527 M2 13 11.38 0.650 M3 18 11.0 0.97

Table 4.13 Windover Mandibular MD Descriptive Statistics of Permanent Teeth with Dental Crowding Tooth N Mean Std. Deviation I1 27 5.59 0.501 I2 29 6.28 0.649 C 28 7.0 0.770 P3 27 7.15 0.602 P4 26 7.04 0.720 M1 31 11.71 0.588 M2 32 11.19 1.120 M3 23 11.35 0.935

76 Table. Table 4.14 Windover Mandibular MD Metrics: Mann-Whitney U Test Results Between Occlusion Cohorts Dental MD Metric P-value, Two-Tailed I1 0.002 I2 0.005 C 0.151 P3 0.098 P4 0.582 M1 0.340 M2 0.547 M3 0.183 Bold entries are statistically significant.

Windover Maxillary MD Metrics Between Crowding Severity Ranks To further isolate the main sources of variation within the dental crowding group, tooth sizes of individuals with mild/moderate and severe crowding were compared (Tables 4.15-4.16). Dividing the crowding sample greatly reduced the sample size per tooth. Only two individuals could be assessed for the maxillary central and lateral incisor in the severe crowding sample. Overall, the severe crowding group has larger maxillary teeth where comparisons can be made (Figures 4.14-4.15). Sample sizes were too small in the severe crowding group to test for significant differences between occlusion cohorts.

77 10 9 8 7 6 5 Mild/Moderate Crowding 4 Severe Crowding MD Means MD Means (mm) 3 2 1 0 I1 I2 C Maxillary Anterior Teeth

Figure 4.14: Windover mesiodistal means for maxillary anterior teeth in individuals with dental crowding. Dental crowding divided by severity.

6 Mild/Moderate Crowding

4 Severe Crowding MD Means MD Means (mm)

0 P3 P4 M1 M2 M3 Maxillary Posterior Teeth

Figure 4.15: Windover mesiodistal menas for maxillary posterior teeth in individuals with dental crowding. Dental crowding divided by severity.

78 Table 4.15 Windover Maxillary MD Descriptive Statistics of Permanent Teeth with Mild/Moderate Crowding Tooth N Mean Std. Deviation I1 7 8 0.816 I2 10 7.4 0.699 C 13 7.85 0.689 P3 12 7 0.739 P4 15 6.6 0.737 M1 16 10.5 0.73 M2 16 10.06 0.772 M3 12 9.08 1.676

Table 4.16 Windover Maxillary MD Descriptive Statistics of Permanent Teeth with Severe Crowding Tooth N Mean Std. Deviation I1 2 9 - I2 2 7 - C 4 8.25 0.5 P3 4 7.25 0.5 P4 4 7.25 0.5 M1 4 10.75 0.5 M2 4 19.75 0.5 M3 3 10 1

Windover Crowding Severity Mandibular MD Metrics Similar to the maxillary tooth size comparisons, tooth size in the severe crowding group is slightly higher than the mild/moderate crowding occlusion cohort (Tables 4.17-4.18). The mandibular second molars showed the most diversity in both dental crowding groups in the Windover population. The mandibular molars exhibit a standard deviation of 1.0. However, mandibular canines within the mild/moderate crowding group show the most diversity with a standard deviation of 1.14. The difference between the mandibular means in the mild/moderate and severe dental crowding groups in Windover mimic the pattern in the maxilla (Figures 4.16-4.17). With the exception of the lateral incisors, the means are higher for all other tooth types for the severe crowding group. Mann-Whitney U tests were conducted to evaluate whether the mandibular teeth of individuals with severe crowding are significantly larger. Due to sample size

79 considerations only incisors and canines were assessed. Only the canine was significantly different between the dental crowding groups (p = 0.022). This indicates that mandibular canine width correlates with severe dental crowding in the Windover sample. Previously, Windover Mesiodistal Metrics Between Sex Cohorts section, significant differences were found between males and females for lateral mandibular incisors. However, the majority of the MD widths in this sample are from subadult samples, therefore sex cannot be evaluated in these occlusion cohorts.

8 7 6 5 4 Mild/Moderate Crowding 3 Severe Crowding MD Means MD Means (mm) 2 1 0 I1 I2 C Mandibular Anterior Teeth

Figure 4.16: Windover mesiodistal means for mandibular anterior teeth in individuals with dental crowding. Dental crowding divided by severity.

80 14

8 Mild/Moderate 6 Crowding Severe Crowding

MD Menas MD Menas (mm) 4

0 P3 P4 M1 M2 M3 Mandibular Posterior Teeth

Figure 4.17: Windover mesiodistal means for mandibular posterior teeth in individuals with dental crowding. Dental crowding divided by severity.

Table 4.17 Windover Mandibular MD Descriptive Statistics of Permanent Teeth with Mild/Moderate Crowding Tooth N Mean Std. Deviation I1 7 5.29 0.488 I2 6 6.33 0.516 C 5 6.6 1.14 P3 4 6.75 0.5 P4 4 7 0.816 M1 6 11.67 0.516 M2 5 11 1 M3 4 11.25 0.957

Table 4.18 Windover Mandibular MD Descriptive Statistics of Permanent Teeth with Severe Crowding Tooth N Mean Std. Deviation I1 16 5.75 0.447 I2 19 6.26 0.733 C 19 7.26 0.562 P3 19 7.21 0.631 P4 19 7.16 0.688 M1 22 11.77 0.612 M2 20 11.35 1.137 M3 13 11.54 0.519

Windover Arch Depth Analyses

Windover Arch Depth Comparisons Between Sexes Males and females show differences in both maxillary and mandibular arch width in the Windover population (Table 4.19). Overall, males show a greater arch depth than females (Figure 4.18). However, Mann-Whitney U tests indicate there are non-significant differences in maxillary (p = 0.322) and mandibular (p = 0.866) arch depth between the sexes.

29 Female 28 Male

Mean ArchMean Depth 27

25 Maxilla Mandible

Figure 4.18: Maxillary and mandibular mean arch depth differences.

Table 4.19 Windover Arch Depth Means by Sex Cohorts Female Male Maxillary N 25 29 Arch Depth Mean 30.59 31.31 Std. Deviation 3.67 2.12 Mandibular N 28 30 Arch Depth Mean 27.50 28.13 Std. Deviation 3.76 2.99

Windover Arch Depth Comparisons Between Sex and Occlusion Cohorts Arch depth was then analyzed in the good occlusion and combined dental crowding groups for the female sample from Windover (Table 4.20). Unexpectedly, the mean arch depth is greater for the dental crowding group in the maxilla and mandible (Figure 4.19). Mann-Whitney U tests of significance were conducted to determine whether the differences are significant between the dental crowding and good occlusion group in the female sample. Differences in arch depth between the dental crowding and good occlusion females were highly significant; P-values = 0.002 (maxilla) and 0.001 (mandible).

34 33 32 31 30 29 Good Occlusion 28 27 Dental Crowding 26

Female ArchFemale Depth Means 25 24 23 Maxilla Mandible

Figure 4.19: Windover female arch depth means for individuals with good occlusion and dental crowding. 83

Table 4.20 Windover Female Arch Depth in Groups with Good Occlusion and Dental Crowding Good Occlusion Dental Crowding Maxillary N 11 13 Arch Depth Mean 28.18 32.83 Std. Deviation 2.23 3.38 Mandibular N 14 11 Arch Depth Mean 25.29 29.54 Std. Deviation 3.89 1.92

Mean differences between good occlusion and dental crowding males exhibit a different pattern from that characterizing the females (Table 4.21). The mean maxillary arch depth was slightly larger in the dental crowding group than in the good occlusion group (Figure 4.20). Interestingly, male arch depth differences between the occlusion groups are not significant in either the maxilla (p = 0.611) or the mandible (p = 0.202). This indicates that arch depth correlates with dental crowding in the female sample but not for males within Windover .

29 Good Occlusion Dental Crowding 28

Male Arch Male Depth Means 27

26 Maxilla Mandible

Figure 4.20: Windover male arch depth means for individuals with good occlusion and dental crowding.

Table 4.21 Windover Male Arch Depth in Groups with Good Occlusion and Dental Crowding Good Occlusion Dental Crowding Maxillary N 13 12 Arch Depth Mean 30.69 31.17 Std. Deviation 1.75 1.64 Mandibular N 15 11 Arch Depth Mean 27.80 29.09 Std. Deviation 2.83 3.27

Windover Arch Width Analyses

Windover Arch Width Comparisons Between Sex Cohorts Males exhibit greater mean arch width than females for both the maxilla and the mandible (Table 4.22 and Figure 4.21), but these differences are not significant (Mann-Whitney U test, p = 0.134). Therefore, similar to arch depth, sexual dimorphism alone does not contribute to arch width variation in the Windover sample.

25.5

24.5 Female 24 Male

Arch Width Means WidthArch Means (mm) 23.5

23 Maxilla Mandible

Figure 4.21: Windover mean arch width for females and males.

Table 4.22 Windover Arch Width Descriptive Statistics by Sex Cohort Female Male Maxillary N 27 29 Arch Width Mean 24.39 25.11 Std. Deviation 1.27 1.94 Mandibular N 28 31 Arch Width Mean 23.72 25.10 Std. Deviation 1.17 1.03

Windover Arch Width Comparisons Between Sex and Occlusion Cohorts Differences within the female group were observed between the good occlusion group and the dental crowding group (Figure 4.22), particularly in the mandible. Overall the female dental crowding group has a smaller mean arch width (Table 4.23). Comparisons of the maxillary arch width means in both groups only showed a slightly mean arch width in the dental crowding group compared to the good occlusal group (difference of 0.06mm). Mann-Whitney U tests of significance showed non-significant P-values between the two groups for maxillary arch width (p = 0.742) within the female sample, however significant values were recorded for mandibular arch width (p = 0.040).

86 25

24.5

23.5 Good Occlusion 23 Dental Crowding

22.5 Arch Width Means WidthArch Means (mm) 22

21.5 Maxilla Mandible

Figure 4.22: Windover female arch width in groups with good occlusion and dental crowding.

Table 4.23 Windover Female Arch Width in Groups with Good Occlusion and Dental Crowding Good Occlusion Dental Crowding Maxillary N 12 13 Arch Width Mean 24.51 24.45 Std. Deviation 1.30 1.30 Mandibular N 13 16 Arch Width Mean 23.77 22.56 Std. Deviation 1.17 1.03

Male arch width also showed differences between the good occlusion and dental crowding groups (Figure 4.23). Similar to the females, mean arch widths (maxillary and mandibular) in the dental crowding group are smaller than the good occlusion group (Table 4.24). Mandibular arch widths showed a greater difference between the occlusion groups than the maxillary measurements. However, the maxillary arch widths in the males were more disparate between the dental crowding and good occlusion groups (difference of 0.54) than the female maxillary width averages. This may indicate a greater variance of arch width for males in the Windover population than for the females.

87 Similar to the female group, the male sample exhibited significantly different values between the good occlusion and dental crowding groups for mandibular arch width. However, male maxillary arch width was not significant between the occlusion groups. This suggests arch width correlates with dental crowding for both sex groups in the mandible but not in the maxilla. There appears to be similar environmental effects on arch width between males and females, which is influencing dental crowding development.

25.5

25 Good Occlusion 24.5 Dental Crowding

Arch Width Means WidthArch Means (mm) 24

23.5 Maxilla Mandible

Figure 4.23: Windover male arch width in individuals with good occlusion and dental crowding.

Table 4.24 Windover Male Arch Width in Groups with Good Occlusion and Dental Crowding Good Occlusion Dental Crowding Maxillary N 15 12 Arch Width Mean 25.35 24.81 Std. Deviation 1.88 2.16 Mandibular N 16 11 Arch Width Mean 25.50 24.27 Std. Deviation 1.09 1.42

88 Windover Cranial Metrics Analyses

Windover Cranial Metrics Comparisons Between Sex Cohorts To investigate sexual dimorphism differences in the cranial shape, cranial metric comparisons were conducted between the female and male samples (Table 4.25). To control for developmental differences subadults were excluded from the analysis. Almost all of the cranial metrics (22 out of 27) were found to be significantly different between males and females in the Windover sample (Table 4.26). As expected, all means are larger in the males than the females. This affirms the high degree of skeletal sexual dimorphism observed at Windover. Those variables that are significantly different between males and females are shown graphically in Figures 4.24 - 4.25. There is no region of the crania that is clearly male or female. There was no clear pattern in aspects of the cranial vault, splanchnocranium or mandible for variables that were not significantly differences versus those that were.

Table 4.25 Windover Cranial Metrics: Between Sex Cohorts Female Male Cranial Metric N Mean Std. Deviation N Mean Std. Deviation MXCL 32 178.09 4.53 32 187.06 4.16 MXCB 34 130.85 5.29 31 134.4 4.00 BD 15 119.00 6.06 22 134.40 3.99 BBH 19 137.32 6.72 15 140.43 4.80 CBL 19 102.74 5.84 15 108.60 4.67 BPL 15 94.35 12.18 14 103.63 5.01 MAB 25 59.79 3.51 26 64.15 3.34 MAL 28 51.94 4.31 27 55.81 3.36 BiaBr 32 119.19 5.00 28 126.02 4.71 UFH 16 65.91 4.31 23 70.88 8.07 MFB 33 91.20 4.19 32 94.89 4.07 UFB 33 101.92 3.23 31 107.21 3.42 NH 17 49.45 2.89 24 52.79 2.70 NB 18 24.52 1.81 22 25.72 1.73 OB 25 36.12 2.24 27 37.76 1.91 OH 17 33.71 2.11 24 34.09 2.20 BioBr 32 93.80 3.64 32 98.20 3.29 CH 28 31.178 3.68 33 34.26 3.55 HMB 30 30.31 2.82 32 33.34 3.06 BMB 31 12.78 3.64 32 15.21 10.7 BW 27 92.41 7.42 31 104.97 5.84 BB 20 120.73 6.39 19 129.92 5.57 MRB 30 35.49 2.89 31 36.98 2.61 MXRB 27 48.27 3.28 28 48.95 3.08 MXRH 27 58.33 3.41 27 63.02 7.70 MandL 28 78.34 5.42 30 83.38 4.73 MA 26 27.21 5.50 29 24.45 5.24

Table 4.26 Windover Cranial Metrics: Mann-Whitney U Test Results Between Sex Comparisons Cranial Metric P-value, Two-Tailed MXCL 0.000 MXCB 0.003 BD 0.000 BBH 0.095 CBL 0.005 BPL 0.003 MAB 0.000 MAL 0.004 BiaBr 0.000 UFH 0.009 MFB 0.002 UFB 0.000 NH 0.001 NB 0.040 OB 0.006 OH 1.000 BioBr 0.000 CH 0.001 HMB 0.001 BMB 0.003 BW 0.000 BB 0.000 MRB 0.059 MXRB 0.405 MXRH 0.000 MandL 0.001 MA 0.068 Bold entries are statistically significant.

Figure 4.24: Significant craniall memetric differences between males and females in the Windindover sample.

Figure 4.25: Significant cranial metricric differences between males and females in the Windovverer sample continued.

Windover Cranial Metrics Comomparisons Between Sex and Occlusion Cohortorts Mann-Whitney U tests wewere conducted to evaluate whether cranial metrictric differences were significantly different betwetween males and females with or without dental crowding.cro In the female sample only the mandibulular angle (p = 0.042) was found to be statisticallyally different between the good occlusion andd ddental crowding groups. In the male sample, craranial base length 93 (p = 0.043) and basion-prosthion length (p = 0.014) exhibit significant differences between the dental crowding and good occlusion groups (Table 4.24).

Table 4.27 Windover Cranial Metrics: Mann-Whiney U Test Results Between Occlusion Cohorts in Females and Males Female Male Cranial Metric P-value, Two-Tailed P-value, Two-Tailed MXCL 0.870 0.130 MXCB 0.614 0.310 BD 0.245 0.643 BBH 0.530 0.604 CBL 0.092 0.038 BPL 0.179 0.018 MAB 0.763 0.642 MAL 0.878 0.568 BiaBr 0.762 0.083 UFH 0.908 0.143 MFB 0.262 0.155 UFB 0.415 0.266 NH 0.791 0.393 NB 0.491 0.537 OB 0.215 0.612 OH 0.560 0.194 BioBr 0.330 0.054 CH 0.667 0.421 HMB 0.844 0.981 BMB 0.622 0.323 BW 0.725 0.438 BB 0.424 0.916 MRB 0.337 0.063 MXRB 0.349 0.511 MXRH 0.638 0.555 MandL 0.218 0.093 MA 0.042 0.330 Bold entries are statistically significant.

Windover Attrition Analyses As expected, attrition in the Windover sample increased with age (Figure 4.26). Interestingly, the attrition score (both maxillary and mandibular) increased at a steady rate in each age cohort by approximately 50 points (Tables 4.28–4.29). These results suggest dental

94 wear was constant throughout life. Overall, mandibular attrition means are higher (approximately 10 points) than those in the maxilla.

250

200

150

Maxilla 100 Mandible Attrition Attrition Score Mean 50

0 Juvenile Y. Adult M. Adult O. Adult

Figure 4.26: Windover attrition comparison between age cohorts in the maxilla and mandible.

Table 4.28 Windover Maxillary Attrition Scores: Descriptive Statistics for Age Cohorts Age Cohort N Mean Std. Deviation Juvenile 12 66.25 14.542 Y. Adult 20 118.20 31.550 M. Adult 27 170.22 26.510 O. Adult 21 221.81 45.321

Table 4.29 Windover Mandibular Attrition Score: Descriptive Statistics for Age Cohorts Age Cohort N Mean Std. Deviation Juvenile 11 72.45 11.553 Y. Adult 18 125.0 36.710 M. Adult 26 187.92 43.994 O. Adult 22 229.77 49.653

95 Windover Attrition Mean Comparisons Between Age and Sex Cohorts To investigate differential dental use for males and females, attrition was analyzed between sex and age cohorts. Overall, the means between males and females are fairly similar (Tables 4.30–4.31 and Figures 4.27–4.28). Mann-Whitney U non-parametric T-tests were conducted to compare the attrition levels between the age and sex cohorts. Attrition levels (both maxillary and mandibular) in all three adult age groups are non-significant between the sexes (Table 4.32).

250

200

150 Female 100 Male

Maxillary Attrition Maxillary Attrition Means 50

0 Y. Adult M. Adult O. Adult

Figure 4.27: Maxillary attrition means between age and sex cohorts.

96 250

200

150

Female 100 Male

50 Mandibular Attrition Mandibular Attrition Means

0 Y. Adult M. Adult O. Adult

Figure 4.28: Mandibular attrition means between sex and occlusion cohorts.

Table 4.30 Windover Maxillary Attrition Scores: Descriptive Statistics for Age Cohorts by Sex Female Male Age Cohort N Mean Std. Deviation N Mean Std. Deviation Y. Adult 15 114.40 35.420 5 129.6 10.991 M. Adult 13 182.77 24.035 13 175.92 30.311 O. Adult 6 227.17 63.402 15 219.67 38.489

Table 4.31 Windover Mandibular Attrition Scores: Descriptive Statistics for Age Cohorts by Sex Female Male Age Cohort N Mean Std. Deviation N Mean Std. Deviation Y. Adult 13 124.08 42.472 5 127.40 17.487 M. Adult 12 189.58 52.394 12 184.75 39.095 O. Adult 6 225.83 83.038 16 231.25 33.832

Table 4.32 Windover Attrition Scores: Mann-Whitney U Test Results for Age Cohorts by Sex Maxilla Mandible Age Cohort P-value, Two-Tailed P-value, Two-Tailed Y. Adult 0.081 0.138 M. Adult 0.555 0.751 O. Adult 0.533 0.417

Windover Attrition Mean Comparisons Between Age and Occlusion Cohorts To analyze patterns between attrition and dental crowding comparisons were conducted between good occlusion and dental crowding groups within each age cohort (Figures 4.29-4.30). Small sample sizes in the juvenile age group meant that only adults were included in this analysis. Dental attrition mean scores were higher in the young adult maxillary good occlusion group by approximately 39 points. There was little variation between occlusion cohorts in the middle-aged and older adult groups (Tables 4.33-4.34). However, the maxillary dental crowding group in the middle-aged and older adult cohorts exhibited higher attrition means than the good occlusion group. It appears from the Windover LII scores that the group with the highest amount of dental crowding (young adult group) exhibits the greatest difference in attrition scores (individuals with dental crowding had lower attrition scores). However, Mann Whitney U tests were conducted on the young adult group to assess correlations between dental crowding and dental attrition and only non-significant differences were found in the maxilla (p = 0.130) and mandible (p = 0.243).

250

200

150 Good Occlusion 100 Dental Crowding

50 Attrition Attrition Score Mean

0 Y. Adult M. Adult O. Adult

Figure 4.29: Windover maxillary attrition comparison between occlusion cohorts.

98 300

250

200

150 Good Occlusion

100 Dental Crowding

Attrition Attrition Score Mean 50

0 Y. Adult M. Adult O. Adult

Figure 4.30: Windover mandibular attrition means comparison between occlusion cohorts.

Table 4.33 Windover Maxillary Attrition Scores: Descriptive Statistics for Age and Occlusion Cohorts Good Occlusion Dental Crowding Age Cohort N Mean Std. Deviation N Mean Std. Deviation Y. Adult 4 149.50 52.697 16 110.37 19.466 M. Adult 19 178.00 27.901 6 181.83 28.826 O. Adult 12 218.83 44.849 4 225.25 27.945

Table 4.34 Windover Mandibular Attrition Scores: Descriptive Statistics for Age and Occlusion Cohorts Good Occlusion Dental Crowding Age Cohort N Mean Std. Deviation N Mean Std. Deviation Y. Adult 4 155.25 65.744 14 116.36 20.071 M. Adult 17 189.00 37.578 6 159.00 48.982 O. Adult 12 243.50 41.252 4 214.00 37.982

Windover Non-Masticatory Dental Wear Patterns Wear patterns were used to investigate the relationship between dental crowding and cultural practices using teeth as tools and other non-masticatory factors. Patterns observed in the Windover sample include interproximal grooving, lingual wear on anterior teeth, and a previously undocumented pattern on posterior teeth termed lingual root wear. Lingual root wear was only present in Windover. The spatial distribution of the wear patterns were analyzed to

99 investigate associations of these dental wear patterns with dental crowding or particular levels of dental crowding severity.

Interproximal Grooved Dental Wear Patterns Interproximal grooves only occurred in adult dentitions. This is to be expected since these patterns are usually the result of the repetitive insertion of an instrument between the teeth over a long period of time (Ubelaker 1989). This wear pattern was found in the maxillary and mandibular dentition and was only observed between molars. More cases were recorded in the females (females (14 cases), males (10 cases). Individuals with interproximal grooved wear are clustered in the western portion of the northeast subsection (Figure 4.31). This corresponds with the subdivision proposed for dental crowding severity. The western group has an interproximal grooved wear frequency of 39 percent, while the eastern division exhibits a rate of 13 percent. However, individuals with interproximal grooved wear cluster on the opposite side of the subsection than those with the higher frequency of severe dental crowding. Two tailed Fisher’s Exact tests were conducted to compare the interproximal grooving frequencies between the east and west divisions of the NE subsection. The frequencies were not significantly different between the divisions (p = 0.0552). Therefore, the frequency of this dental wear pattern was not distinct between the east and west subpopulations.

100

Figure 4.31: Interproximal groovedd wwear pattern plotted within Windover pond. North andd eastea axes are burial coordinates measured in meters. Trianriangles indicate presence of grooved wear and squares representrep absence of grooved wear patterns. Thee divdividing line shows the hypothesized pond division intoo twotw groups.

101 Lingual Root Wear Dental Pattern An interesting dental wear pattern was present only at Windover. This wear is located on the lingual surface of the exposed root just below the crown, beginning at the CEJ and extending down to the root (Figure 4.32). There were well-defined areas of eburnation-like grooving (appearance of groove is shiny and smooth). It is primarily found on molars (maxillary and mandibular), in particular the first and second molars, and occasionally on P4s and M3s. I was careful not to confuse this wear pattern with similar patterns resulting from postmortem calculus removal, which appears as a discoloration around the same region rather than a groove. This pattern is found only in adults and is fairly evenly distributed between males (10 individuals) and females (7 individuals). This suggests the activity causing this wear pattern is not exclusively conducted by one sex. Lingual root wear patterns occurred in individuals with both good occlusion and those with dental crowding.

Figure 4.32 Example of lingual root wear pattern on right maxillary third molar from the Windover sample. This individual (Burial 87) exhibited lingual root wear, in varying degrees of severity, on all maxillary molars; the third molars in this individual showed the most extreme manifestation of the wear pattern. Note the calculus development around the wear patterns.

102 A scatterplot (Figure 4.33) of the spatial distribution of lingual root wear reveals an identical clustering pattern to that seen for interproximal grooved wear. Individuals exhibiting lingual root wear cluster on the same side of the northeastern subsection. The frequency of lingual root wear within the western division of the NE subsection is 26 percent, while the frequency in the eastern section is 13 percent. However, Fisher’s Exact tests indicate that these frequencies are not significantly different (p = 0.7632). However non-significant results for lingual root wear might be the product of the low frequency of this practice.

103

Figure 4.33: Lingual root wear pattern plotted within Windover pond. North and east axes are burial coordinates measured in meters. Stars indicate presence of LRW, squares represent absence of LRW. The dividing line shows the hypothesized pond division into two groups.

104 Incisor Lingual Wear Pattern Another wear pattern was recorded on the lingual portion of the crowns of mandibular incisors. This wear is oriented vertically rather than the horizontal orientation seen with lingual root wear on the posterior teeth. This type of wear is only found in adult dentitions (ranging from young to older adult) in the Windover sample and it is limited to 10 dentitions: males (4 cases) and females (6 cases). It was recorded separately from the rounded wear exhibited in anterior teeth with full dentine exposure. Unlike the grooved wear and lingual root wear spatial clusters, this type of incisor wear appears to be sparsely distributed throughout the charnel pond (Figure 4.34).

105

Figure 4.34: Lingual wear pattern on incisors plotted within Windover pond. North and east axes are burial coordinates measured in meters. Diamonds indicate presence of dental wear pattern; squares represent absence of the wear pattern.

106

CHAPTER 5 COMPARISONS OF WINDOVER TO OTHER FLORIDA ARCHAIC SAMPLES

In this chapter, the Windover results are compared to dental crowding analyses for the six Florida Archaic comparative samples. These data are presented to assess the uniqueness of the dental crowding incidence at Windover and to further investigate dental crowding etiology. Unfortunately, due to small sample sizes, analyses stratified by sex and age could not be conducted for these comparative samples.

Dental Crowding Frequencies From Figures 5.1-5.2, it is clear that Windover exhibits the highest total crowding (47 percent) and severe crowding rates (25 percent), while Little Salt Springs (8VO18) and Republic Groves (8HR4) exhibit no dental crowding. Tick Island (8VO24) has the second highest frequency of total crowding (32 percent) and the highest mild crowding percentage (15 percent). The Gauthier (8Br193) frequency for overall dental crowding is 20 percent; this is evenly divided between mild (10 percent) and severe crowding (10 percent). Bay West (8Cr200) has an overall dental crowding frequency of 11 percent comprised of mild crowding only (Table 5.1). Although one case of mild crowding is observed for Warm Mineral Springs, the small sample size (n = 3) compromised statistical analyses. Therefore Warm Mineral Springs was excluded from the following dental crowding analyses. Dental crowding frequencies were too small to conduct two tailed Fisher’s Exact tests. Heavy taphonomic damage complicated dental crowding assessments of Tick Island. Severe crowding was evident in the Tick Island sample despite this taphonomic damage, however crushing and calcification of the material made the identification of mild/moderate crowding problematic. Some cases of mild/moderate crowding may have been overlooked or could not be assessed.

107 50 45 40 35 30 25

Frequency 20 15 10 5 0

Figure 5.1: Total crowding based on LII scores.

15 Frequency 10

Figure 5.2: Severe Crowding based on LII scores.

108

Table 5.1 LII Dental Crowding Comparison of Samples Good Total Mild Moderate Severe Site Sample Occlusion Crowding Crowding Crowding Crowding Windover % 53% 47% 13% 9% 25% n = 89 # 47 42 12 8 22 Little Salt % 100% 0 0 0 0 Springs n = 6 # 6 0 0 0 0 Gauthier % 20% 10% 0 10% n = 20 # 16 4 2 0 2 Republic % 100% 0 0 0 0 Groves n = 6 # 6 0 0 0 0 Tick % 68% 32% 15% 12% 6% Island n = 34 # 23 11 5 4 3 Bay West % 89% 11% 11% 0 0 n = 9 # 8 1 1 0 0 The # symbolizes the number of individuals with dental crowding. Percentages are rounded.

Dental Nonmetric Analyses Small samples sizes proved particularly problematic for dental nonmetric comparisons since many of these traits manifest in low frequencies. Therefore an aggregate of samples including Little Salt Springs, Warm Mineral Springs and Bay West) was compared with Windover. Fisher’s Exact Tests were used to compare Windover with the other samples (Appendix I). Overall, nonmetric trait frequencies are not significantly different between Windover and the comparative samples. One exception is enamel extensions of the maxillary second molar, where the frequencies for Gauthier and Republic Groves are significantly different from Windover (Table 5.2). Tick Island is similar to Windover for this trait.

109

Table 5.2 Frequencies of Maxillary Second Molar Enamel Extensions Site being compared to Windover Enamel Extensions UM2 Gauthier p < 0.0001 Republic Groves p = 0.0399 Tick Island p = 0.1678 Aggregate p = 0.0162

Though not found to be significantly different in its distributions the presence of Uto-Aztecan premolars (UAPs) is notable due to the rarity of this trait (Figure 5.5). UAPs are characterized as a pronounced ridge extending from the apex of the buccal cusp to the distal occlusal border at or near the sagittal sulcus on a maxillary third premolar. A distal fossa is commonly situated between the distosagittal ridge and the distal occlusal border of the buccal cusp (Johnson et al. 2011, Turner et al. 1991). The frequencies range between 0-16.7 percent in Native American populations. The Uto-Aztecan premolar at Windover (Burial 265) is the oldest documented case of UAP in the world (Johnson et al. 2011). UAPs were only present at Windover (Figure 5.3) and Tick Island (Figure 5.4). Windover exhibited one case and there were three UAP cases observed at Tick Island (including one individual with two UAPs).

110

Figure 5.3: Unerupted unilateral Uto-Aztecan premolar from Windover (Burial 265)

Figure 5.4: Tick Island UAP drawings. A and B are bilateral manifestations of a UAP in a single individual (Burial 54 2003-49-54). C represents a unilateral UAP in an individual with moderate attrition (Burial 100/101 2003-49-95 box 3). Drawings courtesy of Eduardo Miyar.

Table 5.3 Nonmetric Frequencies

Site Sample UM2 Enamel Extensions UAP

Windover % 70% 4% n = 73 # 51 1 Gauthier % 13% 0 n = 20 # 4 0 Tick Island % 45% 6% n = 34 # 11 2 The # symbolizes the number of individuals with the trait. Percentages are rounded.

111 80

40 Enamel Extensions UM2 Uto-Aztecan Premolar UP3 30 Nonmetric Nonmetric Frequency

0 Windover Gauthier Tick Island

Figure 5.5: UM2 Enamel Extension and UAP frequencies for Windover, Gauthier, and Tick Island.

An extreme manifestation of the tuburculum dentale trait, known as talon cusps (Turner 1998), was noted separately in this study. Talon cusps, a rare trait, are documented for Windover (3 cases) and Tick Island (2 cases). Talon cusps consist of cingulum projections on either the labial or lingual surfaces of maxillary and mandibular incisors and canines. Those documented at Windover are the oldest recorded cases in the New World (Stojanowski et al. 2010). This demonstrates further dental similarities between Windover and Tick Island.

112

Figure 5.6: Lingual talon cusp on a permanent maxillary left I2 from Windover (specimen 75).

In addition to the dental traits of the ASUDAS system, six other dental traits having primarily genetic origins were noted. These variants include premolar rotation (both maxillary and mandibular), polydontia (supernumerary teeth), tooth agenesis, impaction, concrescence and gemination. Windover exhibited the greatest number of cases for all six variants - polydontia (n = 3), impaction (n = 4), agenesis (n = 8), concrescence (n = 1), gemination (n = 2) and premolar rotation (n = 12) (Table 5.4). However, Gauthier show the highest frequency of agenesis (10 percent), polydontia (5 percent), and premolar rotation (14 percent). Warm Mineral Springs exhibited the highest rate of impaction (20 percent) but this frequency is most likely inflated because of small sample size (n = 5). Republic Groves exhibited one case of agenesis (8 percent) and premolar rotation (8 percent). The Tick Island sample had one case of polydontia (3 percent) and impaction (3 percent). Little Salt Springs and Bay West had no observable cases of any of the variants. Overall Gauthier exhibits the most comparable frequencies to Windover. However nonmetric frequencies were too small to conduct two tailed Fisher’s Exact tests.

113

Table 5.4 Dental Nonmetric Frequencies

Premolar Site Sample Polydontia Impaction Agenesis Concrescence Gemination Rotation Windover % 3% 5% 9% 1% 2% 13% n = 89 # 3 4 8 1 2 12 Little Salt % 0 0 0 0 0 0 Springs n = 6 # 0 0 0 0 0 0 Gauthier % 5% 10% 10% 0 0 14% n = 21 # 1 2 2 0 0 3 Republic % 0 0 8% 0 0 8% Groves n = 10 # 0 0 1 0 0 1 Tick Island % 3% 3% 0 0 0 6% n = 34 # 1 1 0 0 0 2 Warm % 0 20% 0 0 0 0 Mineral Springs # 0 1 0 0 0 0 n = 5 Bay West % 0 0 0 0 0 0 n = 10 # 0 0 0 0 0 0

Windover is the only sample with examples of fusion and gemination. Gemination was recorded in both the permanent (Burial 98) and deciduous (Burial 421) dentition (Figures 5.7-5.8). Fusion of a fully formed supernumerary molar to the root of an erupted third molar was also recorded (Burial 123). These teeth were fused via cementum (concrescence) and not dentin. The incisor geminations, particularly in the permanent dentition, and the concrescence case are exceptionally rare dental anomalies (Nunes et al. 2002, Romito 2004) demonstrating the unique genetic makeup of the Windover population.

114

Figure 5.7. Gemination of deciduous mandibular incisors in the Windover population (Burial 421).

Figure 5.8: Gemination of permanent mandibular incisors in the Windover population (Burial 98)

115

Figure 5.9. Concrescence of a mandibular third molar in the Windover population (Burial 123).

Cranial Nonmetric Analyses Cranial nonmetric frequencies were compared between the Florida Archaic samples (Appendix II). Overall cranial postmortem damage in the comparative samples complicated nonmetric assessments; in particular cranial nonmetric traits could not be recorded for any of the Tick Island crania due to heavy taphonomic damage. Of the remaining five comparative samples, the frequencies were too low to conduct Mann- Whitney U tests of significance. However 15 of the 32 nonmetric traits were found in frequencies over 20 percent in the Windover population and/or comparative samples (Tables 5.5– 5.6). In particular high frequencies of the suparorbital notch, zygomatic facial foramina, and tympanic dehiscence were observed in all six populations. However, many of these frequencies are confounded by low sample sizes in the comparative samples. Based on the cranial variants presented in Tables 5.5-5.6 Gauthier exhibits the most comparable frequencies (with the exception of tympanic dehiscence) to Windover of the five comparative samples. These results contrast with the dental nonmetric data that showed Gauthier to be dissimilar to Windover. Since Gauthier had the largest sample size of the comparative samples, these cranial nonmetric frequencies may be a direct reflection of sample size discrepancies rather than genetic similarities. Though found in higher frequencies in the New World than other world populations, the inca bone (Os incae) was only observed in two individuals from this

116 study (both from Windover). Notably a unique variant of the inca bone, the partial asymmetrical bipartite variant (Os incae duplex asymmetricum), was present in a Windover individual (Figure 5.10). In their study of Os incae variant frequencies in various world populations, Hanihara and Ishida (2001) found the partial asymmetrical bipartite inca bone to occur very rarely in New World populations. To this author’s knowledge this inca bone variant has not been recorded previously in a skeletal sample from the southeast region of North America.

Figure 5.10 Example of an incomplete asymmetrical bipartite Inca bone variant from the Windover sample (Burial 69B). The transverse incision of the calvaria is from postmortem brain tissue removal.

117

Table 5.5 Cranial Nonmetric Frequencies Zygomatico- Supraorbital Supraorbital Infraorbital Parietal Lambdoid Condylar facial Notch Foramen Suture Foramen Ossicle Canal Foramina Windover Sample # 93 92 52 73 84 80 52 Present # 84 44 26 65 61 39 43 % 90% 48% 50% 89% 73% 49% 83% Little Salt Springs Sample # 6 6 6 6 1 2 1 Present # 6 3 3 4 0 2 1 % 100% 50% 50% 67% 0 100% 100% Gauthier Sample # 14 15 5 7 14 11 11 Present # 14 9 1 4 7 2 0 % 100% 60% 20% 57% 50% 18% 0 Republic Groves Sample # 4 1 0 4 4 3 0 Present # 4 1 0 3 3 0 0 % 100% 100% 0 75% 75% 0 0 Warm Mineral Springs Sample # 4 4 3 2 4 3 1 Present # 4 0 1 2 3 0 0 % 100% 0 33% 100% 75% 0 0 Bay West Sample # 7 7 7 5 6 5 6 Present # 7 3 1 4 0 1 5 % 100% 43% 14% 80% 0 20% 83% Sample # refers to the total number of individuals assessed for that trait; Present # refers to the number of individuals with traits present. Percentages were rounded.

118

Table 5.6 Cranial Nonmetric Frequencies Continued Right Bifurcate Foramen Foramen Pterygo- Temporal Mylohyoid Flexure Flexure Tympanic Ovale Spinosum spinous Mastoid Bridge Saggital Saggital Dehiscence Incomplete Incomplete Bridge Foramen (Center) Sulcus Sulcus Windover Sample # 82 82 61 65 46 52 91 88 Present # 20 55 13 21 20 76 46 20 % 24% 67% 21% 32% 43% 67% 51% 23% Little Salt Springs Sample # 1 1 3 3 1 2 2 4 Present # 0 0 0 1 0 2 1 1 % 0 0 0 33% 0 100% 50% 25% Gauthier Sample # 5 5 7 7 7 11 2 13 Present # 2 3 2 3 4 3 2 4 % 40% 60% 29% 43% 57% 17% 100% 31% Republic Groves Sample # 2 2 2 3 1 4 4 7 Present # 0 2 0 1 1 3 3 1 % 0 100% 0 33% 100% 75% 75% 14% Warm Mineral Springs Sample # 3 3 1 1 1 3 3 3 Present # 2 0 0 0 0 3 1 0 % 67% 0 0 0 0 100% 33% 0 Bay West Sample # 6 6 7 7 5 4 4 1 Present # 5 0 1 1 0 3 0 0 % 83% 0 14% 14% 0 75% 0 0 Sample # refers to the total number of individuals assessed for that trait; Present # refers to the number of individuals with traits present. Percentages were rounded.

Tooth Size and Arch Size Analyses As discussed previously, tooth size and arch size are strongly correlated with dental crowding (Calcagno 1989, Carlson and Van Gerven 1977, Greene 1970). In particular tooth-size arch-size discrepancies (TSASD) are the most common cause of anterior dental crowding (Alt and Türp 1998b). To investigate correlations between dental crowding and TSASD, comparisons of tooth-size, arch-depth and arch-width were made between Windover and the comparative samples.

119

Tooth Size Analyses Due to small samples sizes, an aggregate of the comparative samples was used as an MD comparison to Windover; this is justified by a Kruskal-Wallis non-parametric analysis of variance on all six comparative samples. All tooth types exhibited non- significant differences; therefore the aggregation of these measurements is justified. Due to sample size constraints, some comparisons were still not possible. To assess the effect of tooth size on dental crowding in this sample, comparisons of the good occlusion and dental crowding groups were made. Only permanent dentition was used in this study. Due to the low attrition rates in younger individuals, subadult permanent dentition was also included.

Maxillary MD Metrics Between Occlusion Cohorts Tooth sizes in the comparative aggregate sample are smaller than Windover, particularly for the dental crowding group (Tables 5.7-5.8). The maxillary teeth in the comparative show fewer differences between the dental crowding and good occlusal groups than what is seen for Windover (Figure 5.11 and Figure 5.12). Of the eight tooth types, only half of them are larger in the group with dental crowding: I1, C, M1, and M3. Mann-Whitney U tests for maxillary tooth differences between the occlusal cohorts found non-significant differences for all tooth types. Therefore, dental crowding does not appear to correlate with maxillary tooth width in the comparative aggregate sample. These results are similar to Windover.

120 10 9

8 7 6

5 Good Occlusion 4 Dental Crowding MD Mean (mm)MD Mean 3

2 1 0 I1 I2 C Maxillary Anterior Teeth

Figure 5.11: Comparative aggregate sample of mesiodistal mean dimensions maxillary anterior teeth in individuals exhibiting good occlusion and dental crowding.

6 Good Occlusion Dental Crowding MD Mean (mm)MD Mean 4

0 P3 P4 M1 M2 M3 Maxillary Posterior Teeth

Figure 5.12: Aggregate of maxillary mesiodistal averages of posterior permanent teeth in individuals with good occlusion and dental crowding.

121

Table 5.7 Aggregate Sample Maxillary MD Descriptive Statistics of Permanent Teeth with Good Occlusion Tooth N Mean Std. Deviation I1 14 8.64 0.745 I2 15 7.53 0.516 C 22 8.23 0.429 P3 22 7.73 0.767 P4 23 7.22 0.671 M1 25 10.96 0.869 M2 23 10.87 0.869 M3 19 9.42 0.692

Table 5.8 Aggregate Sample Maxillary MD Descriptive Statistics of Permanent Teeth with Dental Crowding Tooth N Mean Std. Deviation I1 4 8.75 0.5 I2 5 7.0 0.707 C 6 8.33 0.516 P3 7 7.43 0.535 P4 6 7.17 0.408 M1 7 11 0.577 M2 7 10.71 0.951 M3 4 9.75 0.957

Mandibular MD Metrics Between Occlusion Cohorts All of the mandibular teeth in the comparative sample, with the exception of the canines, show slightly larger tooth sizes in the dental crowding group than in individuals with good occlusion (Figures 5.13-5.14). The posterior dentition (in particular the three molars) showed the greatest mean difference between the occlusion cohorts (Tables 5.9- 5.10). This differs from the Windover results that exhibit significant differences between occlusion cohorts and mandibular incisors. Mann-Whitney U tests of significance were conducted for mandibular MD molar width (teeth with the greatest mean differences) between good occlusion and dental crowding groups. Similar to Windover, these teeth exhibited non-significant differences. Due to small sample sizes in the dental crowding group, tests of significance could not be conducted on anterior mandibular dentition in the comparative aggregate sample.

122 8

4 Good Occlusion 3 Dental Crowding MD Mean (mm)MD Mean 2

0 I1 I2 C Mandibular Anterior Teeth

Figure 5.13: Aggregate mesiodistal mean for mandibular anterior teeth in individuals exhibiting good occlusion and malocclusion.

6 Good Occlusion Dental Crowding MD Mean (mm)MD Mean 4

0 P3 P4 M1 M2 M3 Mandibular Posterior Teeth

Figure 5.14: Aggregate of mandibular mesiodistal averages of posterior permanent teeth in individuals with good occlusion and dental crowding.

123

Table 5.9 Aggregate Sample Mandibular MD Descriptive Statistics of Permanent Teeth with Good Occlusion Tooth N Mean Std. Deviation I1 7 5.57 0.535 I2 10 6.30 0.483 C 14 7.21 0.426 P3 13 7.15 0.689 P4 12 7.25 0.452 M1 15 11.67 0.816 M2 14 11.14 0.770 M3 16 11.06 0.772

Table 5.10 Aggregate Sample Mandibular MD Descriptive Statistics of Permanent Teeth with Dental Crowding Tooth N Mean Std. Deviation I1 3 5.67 0.577 I2 3 6.33 0.577 C 5 7.20 0.837 P3 5 7.20 0.447 P4 4 7.50 0.577 M1 5 12.20 0.447 M2 5 11.80 0.447 M3 3 12.33 0.577

Arch Depth Analyses

Maxillary Arch Depth Little Salt Springs has the largest mean maxillary arch depth, followed by Republic Groves and Gauthier. The smallest maxillary arch depths are for Windover and Bay West. (Figure 5.15 and Table 5.11). Only one individual could be assessed from Warm Mineral Springs for arch depth. Therefore, Warm Mineral Springs was excluded from this analysis. Tick Island was also excluded from the sample due to crushing and concretions from these individuals’ interment in a shell mound. The comparative samples utilized (Little Salt Springs, Gauthier, Republic Groves, and Bay West) consist of combined adult samples, therefore females and males could not be separated.

124 Kruskal-Wallis non-parametric analysis of variance was conducted for maxillary arch depth between all four samples. The results showed only non-significant differences between the groups (p = 0.170).

32.5

31.5

30.5

29.5 Maxillary Maxillary Arch WidthMeans 29

28.5

28 Little Salt Springs Gauthier Republic Groves Bay West

Figure 5.15: Mean maxillary arch depth between Windover and the comparative samples.

Table 5.11 Maxillary Arch Depth Descriptive Statistics Between Windover and Comparative Samples Site N Mean Std. Deviation Windover 54 30.98 2.93 Little Salt Springs 4 32.50 2.89 Gauthier 14 31.42 3.61 Republic Groves 5 31.43 3.61 Bay West 6 29.5 1.22

Mandibular Arch Depth Mandibular arch depth varies among the samples but not as greatly as the maxillary dimension (Table 5.12). Windover had the next largest, followed by Gauthier

125 and Republic Groves. These three sites also had the largest sample sizes for this measurement (Figure 5.16). Sample sizes for Warm Mineral Springs and Little Salt Springs were too small for statistical testing, and no data were available for Bay West. Due to heavy taphonomic damage, Tick Island was excluded from the sample. Kruskal-Wallis non-parametric analysis of variance was conducted for mandibular arch depth between all three samples. Similar to maxillary arch depth, the samples exhibit only non-significant results for mandibular arch depth (p = 0.161). However, based on the significant differences recorded for maxillary and mandibular arch depth in the Windover sample, it is possible that differences between the samples may exist between sex and occlusal cohorts. Unfortunately, small sample sizes prevent these analyses.

27.5

26.5

Mandibular Mandibular Arch Depth Means 25.5

25 Windover Gauthier Republic Groves

Figure 5.16: Mean mandibular arch depth between Windover and the comparative samples.

126

Table 5.12 Mandibular Arch Depth Descriptive Statistics Between Windover and Comparative Samples Site N Mean Std. Deviation Windover 60 27.81 3.32 Gauthier 13 26.69 2.46 Republic Groves 8 26.25 2.96

Arch Width Analyses

Maxillary Arch Width Analyses Descriptive analyses of maxillary arch width measurements were conducted between all seven populations (Table 5.13 and Figure 5.17). Little Salt Springs exhibited the highest arch width average (25.62) of the samples. Gauthier, Republic Groves and Windover were all within 0.15 mm of one other (ranging from 24.76 - 24.97). Due to low sample sizes Warm Mineral Springs was not included in these analyses. Tick Island was also excluded because of heavy taphonomic damage. Kruskal-Wallis non-parametric analysis of variance was conducted for maxillary arch width between all four samples. Maxillary arch width was found to be non- significant between the samples (p = 0.438).

127 25.2

24.8

24.6

24.4

24.2

23.8 Maxillary Maxillary Arch WidthMeans 23.6

23.4 Windover Gauthier Republic Warm Bay West Groves Mineral Springs

Figure 5.17: Mean maxillary arch width between Windover and the comparative samples.

Table 5.13 Maxillary Arch Width Descriptive Statistics for Windover and Comparative Samples Site N Mean Std. Deviation Windover 56 24.76 1.67 Gauthier 15 24.97 2.64 Republic Groves 5 24.81 3.09 Bay West 6 24.00 1.23

Mandibular Arch Width Analyses Mandibular arch width means were compared between the populations under study. The mandible showed more variability in arch width between the groups than maxillary arch width (Figure 5.18). The largest means were recorded in Republic Groves (25.11), Gauthier (24.69) and Windover (24.31) (Table 5.14). Sample sizes for Warm Mineral Springs and Little Salt Springs were too low for statistical confidence and were excluded from the analyses. Tick Island was also excluded due to taphonomic damage. Kruskal-Wallis non-parametric analysis of variance was conducted for mandibular arch width between all three samples. Mandibular arch width was found to be

128 non-significant between the samples (p = 0.058). Therefore there are no discernable differences between these groups for arch width. Nevertheless, as discussed previously arch size between sex and/or occlusal cohorts may differ between the samples; however small sample sizes prevented these assessments.

25.2

24.8

24.6

24.4

24.2 Mandibular Mandibular Arch Width Means 24

23.8 Windover Gauthier Republic Groves

Figure 5.18: Mandibular arch width means between Windover and the comparative samples.

129

Table 5.14 Mandibular Arch Width Descriptive Statistics for Windover and Comparative Samples Site N Mean Std. Deviation Windover 61 24.31 1.62 Gauthier 13 24.69 1.41 Republic Groves 9 25.11 1.45

Cranial Metrics Analyses Mann-Whiney U tests were employed to compare the samples to Windover (Tables 5.17-19). Only measurements with samples sizes of five or greater were included in the analysis. Bay West exhibited the largest number of significant differences (n = 5) with Windover, followed by Gauthier (n = 3), and Republic Groves (n = 0) (Table 5.16). Tests of significance could not be conducted on Warm Mineral Springs due to small sample sizes. Cranial measurements could not be taken on the Tick Island and Little Salt Springs skeletal collections due to heavy taphonomic damage. The high degree of sexual dimorphism (as inferred by the Windover cranial metric results discussed previously) in the Florida Archaic populations may confound cranial metric means if sex is not controlled for. However, due to low sample sizes, cranial metric differences between sex cohorts in the comparative samples could not be assessed.

130

Table 5.15 Cranial Metrics: Descriptive Statistics Between Comparative Samples Gauthier Republic Groves Bay West Cranial Metric N Mean Sd N Mean Sd N Mean Sd MXCL 13 182.15 7.10 4 183.25 6.34 5 176.20 4.03 MXCB 13 136/27 3.28 4 138.25 1.26 5 132.80 6.26 0 - - BD 3 120.70 3.62 3 135.00 5.29 BBH 13 135.50 6.70 4 131.75 13.87 3 134.67 5.69 CBL 12 99.75 3.67 4 98.75 9.36 4 104.00 7.39 3 91.98 6.10 BPL 7 97.93 5.19 3 89.67 15.54 MAB 10 63.65 2.43 4 60.25 4.50 7 60.35 2.37 MAL 13 53.13 3.19 3 53.33 4.93 5 55.01 2.56 BiaBr 13 116.12 31.43 4 124.50 2.65 1 - - UFH 7 67.44 4.76 3 76.00 1.00 5 71.49 2.34 MFB 12 93.71 5.02 4 92.00 7.12 6 87.83 3.31 6 100.83 3.86 UFB 12 105.42 4.42 3 105.33 3.79 NH 6 50.44 2.22 3 55.67 1.53 5 51.90 3.06 NB 6 25.26 1.93 3 24.67 0.58 6 24.50 1.16 OB 10 36.71 1.93 3 38.67 2.31 6 34.69 2.03 OH 7 35.82 2.75 3 36.33 1.53 4 33.40 3.27 6 91.13 2.92 BioBr 12 95.93 3.91 3 100.00 2.00 CH 11 31.23 2.83 6 32.81 2.96 1 - - - - HMB 15 30.73 2.06 7 32.72 1.73 1 1 - - BMB 16 12.92 1.34 8 12.18 1.38 BW 11 104.01 7.41 7 102.71 11.50 0 - - - - BB 8 124.48 9.75 7 129.60 10.46 0 1 - - MRB 14 36.96 2.24 8 37.00 1.88 MXRB 13 47.52 4.65 7 49.35 3.17 1 - - - - MXRH 12 62.33 5.76 8 63.00 1.83 1 MandL 13 82.31 4.14 8 81.62 2.83 1 - - MA 13 25.58 3.20 8 23.44 4.88 0 - -

Table 5.16 Cranial Metric Significant Differences Between Samples Comparative Sample # of Sig. Diff. Metrics of Sig. Diff. Republic Groves 0 - Gauthier 4 MXCB, CBL, FMB, CH Bay West 5 MXCL, MFB, UFB, OB, BioBr, Tests of significance for each site are compared to Windover.

131 Descriptive statistics for variables within Republic Groves are provided in Table 5.15. Mann-Whitney U tests of significance were conducted on traits with sample sizes of five or greater. All variables exhibited non-significant results (Table 5.17). Bizygomatic diameter and upper facial height exhibit the largest mean differences between the Republic Groves sample and Windover.

Table 5.17 Cranial Metrics: Mann-Whitney U Test Results Between Windover and Republic Groves Cranial Metric P-value, Two-Tailed CH 0.561 HMB 0.295 BMB 0.311 BW 0.525 BB 0.491 MRB 0.413 MXRB 0.636 MXRH 0.178 MandL 0.894 MA 0.181

Three cranial measures were found to differ significantly between Windover and Gauthier (Table 5.18). Maximum cranial breadth and cranial base length show the largest differences. Foramen magnum breadth and chin height have smaller mean differences but are still significantly different between Windover and Gauthier (Figure 5.19). The low number of statistically different cranial metrics between these populations may infer cultural and/or genetic similarities affecting cranial shape.

132

Cranial Metrics (mm)

Average

Figure 5.19: Cranial metrics: VarVariables with significant differences between Gauthier andan Windover.

133

Table 5.18 Cranial Metrics: Mann-Whitney U Test Results Between Windover and Gauthier Cranial Metric P-value, Two-Tailed MXCL 0.747 MXCB 0.008 BD 0.431 BBH 0.093 CBL 0.004 BPL 0.337 MAB 0.187 MAL 0.366 BiaBr 0.304 UFH 0.826 MFB 0.882 UFB 0.678 NH 0.407 NB 0.708 OB 0.833 OH 0.122 BioBr 0.690 CH 0.031 HMB 0.265 BMB 0.952 BW 0.325 BB 0.524 MRB 0.557 MXRB 0.618 MXRH 0.523 MandL 0.530 MA 0.904 Bold entries are statistically significant.

Of the five metrics found to be significantly different between Bay West and Windover, maximum cranial length, minimum frontal breadth, biorbital breadth, and the parietal chord exhibit the largest mean differences (Table 5.19 and Figure 5.20). These may reflect cultural or genetic differences between Bay West and Windover.

134

Average CranialAverage Metrics (mm)

Figure 5.20: Cranial metretrics: Significant differences between Bay West and Windindover

135

Table 5.19 Cranial Metrics: Mann-Whitney U Test Results Between Windover and Bay West Cranial Metric P-value, Two-Tailed MXCL 0.027 MXCB 0.732 MAB 0.287 MAL 0.723 UFH 0.112 MFB 0.010 UFB 0.027 NH 0.737 NB 0.613 OB 0.026 BioBr 0.006 Bold entries are statistically significant.

Dental Attrition Analyses Dental attrition rates were compared between the comparative samples and Windover to investigate exogeneous factors in dental crowding development. To control for age factors effecting attrition rates, only adults were used in this analysis. The frequencies of these patterns were analyzed between population samples and age cohorts (Tables 5.20 - 5.22). Due to small sample sizes Little Salt Springs, Republic Groves, Warm Mineral Springs and Bay West were combined into an aggregate sample. Kruskal- Wallas non-parametric analysis of variances was conducted for maxillary and mandibular attrition levels between these four samples to assess distribution differences. Tests for attrition levels in the maxilla (p = 0.129) and mandible (p = 0.244) showed non- significant results; therefore this aggregation is justified. Windover has the greatest attrition rate (maxillary and mandibular) in the young adult group (Table 5.). However Gauthier shows the highest attrition averages in the middle-aged and older adult groups (Tables 5. – 5.). Overall Windover and Gauthier exhibit higher attrition means than the other comparative samples. The high attrition levels in these two samples may suggest similar masticatory stressors between Windover and Gauthier. Mann-Whitney U tests of significance were conducted between Windover and the comparative samples per age cohort. In the young adult group all three sample showed

136 non-significant results- Gauthier (p = 0.914), Tick Island (p = 0.052), aggregate sample (p = 0.099). In the middle-aged adult group Gauthier exhibited non-significant results to Windover (p = 0.239) while the aggregate sample showed significantly different attrition levels (p = 0.037). Tick Island was excluded from this analysis due to its small sample size (n = 3). In the older adult group the aggregate sample also showed significantly different levels from Windover (p = 0.006). Gauthier and Tick Island could not be analyzed for statistical significance due to small sample sizes. Based on these results, it appears that Gauthier is the most similar to Windover for attrition levels in the young and middle-aged adult groups, while the aggregate sample is the most statistically different from Windover in the middle-aged and older adult groups.

Table 5.20 Young Adult Maxillary Attrition Descriptive Statistics Between Windover and Comparative Samples Site N Mean Std. Deviation Windover 20 118.20 31.550 Gauthier 10 115.5 37.756 Tick Island 13 92.92 41.317 Aggregate 8 80.88 47.828

Table 5.21 Middle-Aged Adult Maxillary Attrition Descriptive Statistics Between Windover and Comparative Samples Age Cohort N Mean Std. Deviation Windover 27 170.22 26.510 Gauthier 7 193.14 32.127 Tick Island 3 132.33 34.588 Aggregate 7 144.43 42.067

137

Table 5.22 Older Adult Maxillary Attrition Descriptive Statistics Between Windover and Comparative Samples Age Cohort N Mean Std. Deviation Windover 21 221.81 45.321 Gauthier 2 270.50 40.305 Tick Island 1 212.00 - Aggregate 5 125.20 69.615

138

CHAPTER 6 DISCUSSION

Previous studies provide evidence for increasing dental crowding as the result of dietary shifts from hunter-gatherer to agriculture subsistence practices (Begg 1954, Corruccini 1984, Price 1936). Dental crowding has become a prominent issue in modern societies, with rates of crowding at 60-80 percent in modern industrial populations (Rose and Roblee 2009). However, despite its current prominence in industrialized societies, comparatively little research has been conducted on the etiology of this malocclusion in conjunction with efforts that focus on its correction (Corruccini 1999, Garn 1961). In this dissertation, I developed and tested hypothetical models regarding the etiology of high dental crowding rates in the Florida Archaic Windover population based on cross-cultural dental crowding frequencies and dental anthropological and orthodontic studies, as well as physical anthropological methodologies and theory. This chapter discusses the results, including genetic and environmental etiological sub-hypothesis components:

Primary Hypothesis: The incidence of dental crowding in the Windover populations is anomalous compared to other hunter-gatherer populations including Florida Archaic peoples.

If dental crowding frequencies within the Windover are exceptional then: Genetic Sub-hypothesis: High dental crowding rates are the result of predominately genetic factors with high degrees of phenotypic expression despite heavy attrition and robust morphology Or Environmental Sub-hypothesis: High dental crowding rates are the product of unique cultural practices: either indirectly resulting in craniofacial robusticity differences leading to inadequate arch size or being a direct cause of dental irregularities.

Dental Crowding Within Windover The Windover population represents a hunter-gather society from the Early Archaic Period in Florida that primarily subsisted on riverine resources (Quinn et al. 2008). Little Irregularity Index results show that Windover people had a general dental crowding rate of 47 percent, of which 25 percent was considered severe. Compared to

139 other hunter-gatherer societies, both prehistoric and extant, the rate of dental crowding at Windover appears abnormally high (Table 6.1). In Japanese prehistoric and historic societies, Hanihara et al. (1981) recorded malocclusion rates for the Jomon aquatic hunter-gatherers (3000-2000 BP), emergent agriculturalists from the Kofun period (1800-1600 BP), Medieval agriculturalists (650- 580 BP) and Yedo period agriculturalists (400-100 BP) (these studies included anterior crowding in their malocclusion criteria). The Jomon period sample yielded malocclusion rates of 20 percent (2 percent of which consisted of severe dental crowding); a frequency less than half of Windover. Windover shows an even higher frequency than the Japanese emergent agriculturalists from the Kofun period that exhibit dental crowding rates of 45.5 percent (9 percent of which consisted of severe dental crowding). The frequency of dental crowding recorded at Windover is more equivalent to the Medieval and Yedo agriculturalists from Japan; the Medieval agriculturalists show a total dental crowding frequency of 52 percent (18 percent of which is severe crowding) and the Yedo agriculturalists (400-100 BP) have a dental crowding rate of 56 percent (25 percent of which is severe) (Hanihara et al. 1981). Newman and Snow (1942) conducted dental crowding analyses on Pickwick Basin, Alabama, hunter-gatherer (7,000-3,000 BP) and agricultural (1,200-15,00 AD) skeletal samples. Pickwick Basin hunter-gatherers showed a rate of 20 percent mild/moderate crowding (zero cases of severe crowding) and the agriculturalists exhibited 37 percent mild/moderate crowding (ten percent of which was severe). Again, Windover appears to be more comparable to the agriculturalists than hunter-gatherers. In fact, Windover appears to have crowding more similar to extant populations in the United States: Kelley and Harvey (1977) found a 40-60 percent total malocclusion rate for American youths. Windover is also comparable to severe malocclusion frequencies (ranging from 17 - 38 percent) identified in affluent Western groups (Lombardi and Bailit 1972). In sum, the rates of dental crowding and overall severity at Windover are more similar to agriculturalists than hunter-gatherers. This is extremely unusual given the fact that Windover predates the advent of agriculture in Florida by at least 5,500 years (Kelly et al. 2006). In fact, it appears that this extraordinarily high frequency of total dental

140 crowding and severe crowding found in the Windover has not been identified in any other prehistoric hunter-gatherer society.

Table 6.1 Dental Crowding Frequencies in Hunter-Gatherer and Agricultural Groups from Japan and North America

Date Range Good Total Dental Severe Site Sample (yrs BP) Occlusion Crowding Crowding

Pickwick Basin, AL % 80% 20% 0 Hunter-gatherers 7,000-3,000 n = 30 # 24 6 0 Pickwick Basin, AL % 53% 37% 10% Agriculturalists 1,200-1,500 AD n = 30 # 16 11 3 Jomon, Japan % 80% 20% 2% Hunter-gatherers 3,000-2,000 n = 45 # 36 9 1 Kofun, Japan % 54.5% 45.5% 9% Proto-Agriculturalists 1,800-1,600 n = 11 # 6 5 1 Medieval, Japan % 48% 52% 18% Agriculturalists 680-580 n = 50 # 24 26 9 Yedo, Japan % 44% 56% 25% Agriculturalists 400-100 n = 16 # 7 9 4 Japanese data from Hanihara et al. 1981: 66, North American data from Newman and Snow (1942): 397.

Onset of Dental Crowding in the Florida Archaic Dental crowding was investigated in subadult cohorts to observe the earliest onset of dental crowding within the Florida Archaic populations. The Windover collection provided the largest (n = 26) and most diverse subadult representation of the Florida Archaic samples, ranging from infancy to adolescence. The youngest individual who exhibited measurable dental crowding was a child aged 7-8 years from the Windover sample. The next youngest subadults exhibiting crowding were two children aged 8-9 years, also from the Windover sample. Dental crowding appears to manifest only with the eruption of the anterior permanent dentition and does not primarily affect the deciduous

141 dental arcade prior to the eruption of permanent dentition in this study; this contrasts with cases reporting anterior crowding in primary teeth (Alamoundi 1999, Prabhakar et al. 2008). There was no evidence of anterior crowding in the primary dentition nor was there any evidence of early loss of deciduous teeth; all mixed dentition appeared to be erupting in the normal dental developmental sequence based on Ubelaker’s (1989) study of dental development in Native Americans. However, possible cases of retention of deciduous teeth were found in the Windover population, which led to the impaction or malpositioning of the underlying permanent teeth in those individuals. It does not appear that habitual behaviors during childhood (e.g., thumb sucking) or early loss of deciduous teeth had any effect on the malocclusions recorded in these populations.

Dental Crowding Frequencies Between Age Cohorts Dental crowding was analyzed between age cohorts. This analysis revealed the demographic trends of crowding severity within the Windover population. Dental crowding increased greatly from the child group (4 percent) to the juveniles (13 percent). The majority of the child cohort consists of individuals who are younger than the earliest case of crowding (under seven years of age); therefore, the lower rates of dental crowding in children are a direct result of this age discrepancy. In the adult cohorts, there was a trend of decreasing crowding with age. The juvenile and young adult groups exhibited the greatest rates of total dental crowding and severe crowding within the Windover population. When dental crowding was analyzed within each cohort, a staggering 83 percent of all juveniles exhibit dental crowding, comprised mostly of severe crowding (58 percent), and 75 percent of young adults have dental crowding (45 percent of that being severe). In contrast, older adults show a 33 percent rate of dental crowding with the majority of that expressed as mild crowding (17 percent). Children have a 30 percent crowding rate with an equal distribution of moderate and severe crowding (15 percent). Middle-aged adults have the lowest rate of dental crowding (28 percent), with the majority of mild crowding cases (16 percent). At Windover, total crowding frequencies decline abruptly between the young adult (75 percent) and middle-aged adult groups (28 percent), followed by a slight increase in the older adult group (33 percent). In particular, severe crowding frequencies

142 reduce dramatically from the young adult group (45 percent) to the middle-aged adult group (8 percent). The reduction of dental crowding with age is most probably due to a combination of complications of dental crowding assessment with increased attrition and antemortem tooth loss, as well as improved tooth alignment from increased interproximal space as the result of advanced dental attrition. Tooth wear is greatly affected by age since older individuals have been exposed to the processes of attrition for a longer period of time (Larsen 1995). The reduction of tooth surface area from attrition complicates dental crowding assessments and may lead to an underrepresentation of observed dental crowding cases. Thus, dental crowding rates in older individuals at Windover may be higher than what is recorded in this study. This reduction in observed dental crowding in individuals with greater attrition levels may also support Begg’s (1954) notion of attritional occlusion (harmonious occlusion as the result of heavy tooth wear). The theory of attritional occlusion claims that individuals undergo continuous and dynamic changes to their dentition as a direct result of attritional tooth size reduction and movement throughout life. The greater the loss in tooth mass the more space is created between teeth, which is filled by compensatory tooth migrations, including mesial drift, continuous eruption and lingual tipping. However, Begg’s theory presents attritional occlusion as a preventative process rather than corrective. In the Windover sample, heavy attrition, coupled with biomechanical tooth migration processes, appears to have contributed to the correction of dental crowding by creating space and relieving anterior tooth malalignment. Therefore, this corrective attritional process in older Windover individuals could not be called attritional occlusion, although, it is based on similar processes. These factors seem to have a much greater effect on crowding frequency between young and middle-aged adults and are negligible between the middle and older adult groups. Attrition rates between these groups are consistent, increasing by approximately 50 points from one cohort to the next. Thus corrective attrition has the greatest effect on dental crowding in middle-aged adults but only has a minor effect on crowding in older individuals, despite increased attrition. Further studies of tooth migration and diminished anterior crowding in these samples need to be conducted to assess this process.

143 Dental Crowding Frequencies Between Sex Cohorts There were few differences in dental crowding frequency between the sexes in the Windover population. Both sexes were well represented in the Windover sample. Overall dental crowding was evenly distributed among the sexes for all severity ranks. Differences in crowding frequency between the sexes ranged from 1 – 4 percent. No significant differences were found between the female and male samples. Based on the current results, if cultural practices are the primary factor causing these malocclusions, these practices do not reflect a sexual division of labor. However, analyses of arch depth and arch width between sex and occlusal cohorts within the Windover population contradict these results, suggesting an association between dental crowding and cultural practices that are gender specific despite an even distribution of crowding between males and females (this is further described in the arch shape section below).

Windover Cluster Patterns for Dental Crowding and Dental Wear Patterns Dental crowding was plotted in conjunction with burial coordinates from Windover to investigate cluster patterns and distribution of dental crowding within the charnel pond. Total dental crowding has a broad distribution throughout the cemetery and showed no evidence of clustering in any of the subsections. However there is a cluster of severe crowding in the eastern portion of the northeast subsection of Windover pond (Figure 6.1). The hypothetical divide of this subsection was based on the division recorded by Stojanowski and Schillaci (2006) in their intra-cemetery biodistance analysis of Windover. Based on nonmetric frequency distributions, they suggest the presence of a minimum of two genetically distinct subpopulations within Windover. Their conclusions were based particularly on the east and west divisions of the northeastern subsection. Stojanowski and Schillaci (2006) also noted the higher degree of dental crowding severity in the east division of the pond. The severe dental crowding clustering pattern in this subsection supports the perimeters of this hypothetical divide. When tests of significance for dental crowding severity ranks were conducted between the east and west divisions, the severe crowding frequency was highly significant between the two sections (p = 0.0002) but tests of significance for mild and moderate crowding exhibited non- significant differences between the east and west divide.

144

Figure 6.1: Scatter plot of severity ranks with corresponding burial coordinates within the northeastern subsection of Windover pond. Circles indicate no dental crowding, squares represent mild crowding, triangles represent moderate crowding and diamonds represent severe crowding. The dividing line shows the hypothesized pond division between two groups based on dental crowding clustering. Note the high frequency of severe crowding in the eastern portion of the pond.

Age appears to be a confounding factor, however, in severe dental crowding frequency between the east and west portions. The ratio of elder adults (combination of middle-aged and older) is significantly greater in the west portion than the east. In particular, five juveniles and six young adults constitute 50 percent of the east division’s

145 sample size collectively. In the western division, 15 middle-aged adults and 16 older adults comprised 60 percent of the total sample in the western division. As discussed previously, severe dental crowding rates in the juvenile and young adult groups are more than twice as high as the severe crowding rates in the middle-aged and older adult groups. Therefore, age demographics between the east and west division may be influencing severe crowding rates. To test for cultural differences between these subpopulations, cluster pattern analyses of dental wear patterns were also conducted. Two dental wear patterns (interproximal grooved wear and lingual root wear) exhibited cluster patterns in the NE subsection at Windover, showing high frequencies in the western portion of the subsection. However neither of these wear patterns showed significantly different frequencies between the east and west portions. During this study, a previously undocumented lingual dental wear pattern close to the root of the posterior teeth (termed lingual root wear) was identified in the Windover sample. This wear pattern was not observed in the other comparative samples, and further demonstrates unique cultural practices at Windover. Similar to the dental crowding cluster patterns, cluster analyses of this lingual root wear in conjunction with burial coordinates in the Windover sample revealed clustering on the west side of the northeast subsection. The frequencies of this pattern are not significantly different between the eastern and western divisions, but further study of the wear pattern is warranted

Dental Crowding Between Windover and the Comparative Samples In hunter-gatherer societies, teeth typically fully erupt into proper alignment (including the third molars) and function in mastication (functional occlusion) (Kaifu et al. 2003). Heavy attrition and functional occlusion are the norm in the available literature on hunter-gatherer populations. Therefore the high rate of dental crowding and other malocclusions found in Windover are atypical. Figs. 6.2 – 6.3 are examples of maxillary good occlusion and severe dental crowding cases recorded in the Florida Archaic samples under study.

146

Figure 6.2: Example of good occlusilusion (functional occlusion) in the Gauthier (left) (specimcimen from Box 36A BL5 B-7846) and Republic GroGroves (right) (Burial 133) samples. Note the slight bilaterateral winding in the Republic Groves individual.

Figure 6.3: Example of severe dentaental crowding in the Windover (left) (Burial 86) and Gauthauthier (right) (specimen from Box 23B B-22) samsamples. Note the retention of the right second deciduousus molarm as well as premolar rotations and displacelacements in the Windover individual; also the conical mesiesiodens in the Gauthier individual. Although thehe rretention of the deciduous molar, premolar rotationss andan the mesiodens may have exacerbateded ccrowding, severe anterior crowding was already evidentent in these cases.

Dental crowding devevelopment in the Windover sample may have beeneen influenced from their practice of aquatictic foraging. It is possible the consumption of soffterte meats associated with riverine resouources may have an effect on arch size developmement, however, as discussed previously, thee aaquatic hunter-gatherer groups from the Jomonn periodp in Japan had a low malocclusionion rate. To test for aquatic subsistence factors on dental

147 crowding in this study, comparative analyses were conducted on other Florida Archaic populations practicing hunter-gatherer subsistence with a primary emphasis on riverine or marine resources. When compared to other prehistoric groups in the literature (discussed previously in this chapter) (see Table 6.1), Tick Island’s total dental crowding (32 percent) and mild/moderate crowding rates (26 percent) are higher than the recorded dental crowding frequencies in North American Pickwick Basin and Japanese Jomon period hunter- gatherer groups (20 percent), but lower than the Pickwick Basin agriculturalists (37 percent) or Japanese emergent agriculturalists (45.5 percent) and agriculturalists (52 – 56 percent). The 6 percent severe crowding frequency at Tick Island is also higher than the hunter-gatherer groups but lower than the emergent agriculturalists. The dental crowding rates at Bay West (11 percent), Republic Groves (zero cases of dental crowding), and Little Salt Springs (zero cases of dental crowding) are lower than the Japanese and Pickwick Basin hunter-gatherer groups. The total crowding rate recorded at Gauthier (20 percent) is comparable to the Pickwick Basin and the Jomon period hunter-gatherers but Gauthier’s severe crowding rates (10 percent) are higher than the hunter-gatherer groups and more similar to the Pickwick Basin agriculturalists (10 percent) and Kofun period emergent agriculturalists (9 percent). However, the severe crowding frequency at Gauthier is lower than the Japanese agricultural groups (18-25 percent). Although Gauthier shows high rates of severe dental crowding for a hunter-gatherer society, the total dental crowding frequency from this site is comparable to other hunter-gatherers. Windover’s severe crowding rates are more than ten-times greater than the other hunter- gatherer samples and are more comparable to the more recent Yedo agricultural population (400-100 BP). Numerous studies have confirmed Windover as a hunter-gatherer society (Doran 2002, Quinn et al. 2008, Tuross et al. 1994). As a consequence, the high crowding rates found at Windover cannot be explained by means of agricultural influence. The lower dental crowding rates in the Florida Archaic comparative samples also discounts possible influences from the practice of aquatic foraging. These results are similar to findings by von Cramon-Taubadel (2011), who recorded no significant differences in jaw morphology between hunter-gatherer diets utilizing terrestrial fauna versus aquatic

148 hunter-gatherers. Thus, other factors are involved in the development of dental crowding in the Windover sample. In extant slash and burn horticulturalists from Brazil, the Xavante and Yanomami Indians exhibit disparate dental crowding rates (Table 6.2). A study by Neel et al. (1964) of the Xavante Indians recorded a dental crowding frequency of 5 percent in this population. However, in the Yanomami population, Pereira et al. (1972, 1994) recorded a dental crowding rate of 53 percent. Neither of these populations exhibited observable cases of severe crowding. The differences in dental crowding frequencies between these populations may reflect differential environmental exposures. In particular, Pereira et al. (1978) noted an increase in dental crowding development due to the habitual use of tobacco wads placed between the mandibular incisors and lips. Therefore, it may be possible that dental crowding differences between Windover and the Florida Archaic samples are due to differential extramasticatory influences on the dentition as a result of different cultural practices.

Table 6.2 Dental Crowding Frequencies in Extant Horticulturalists from Brazil

Good Total Dental Severe Site Date Range Sample Occlusion Crowding Crowding

Yanomami, Brazil % 47% 53% unknown Slash and Burn Horticulture Modern n = 148 # 70 78 unknown Xavante, Brazil % 95% 5% 0 Slash and Burn Horticulture Modern n = 37 # 35 2 0 Yanomami data from Pereira et al., (1994): 9, Xavante data from Neel et al. (1964): 107. Percentages are rounded.

Dental crowding is known for its multifactorial etiology (Bernabé and Flores-Mir 2006 (a), Harris and Corruccini 2008, Mossey et al. 1999) and it is possible that these results indicate an underlying genetic origin for mild/moderate crowding within the Florida Archaic samples. Overall, the mild/moderate crowding rates throughout all seven Florida groups range from 0 - 26 percent. These rates are comparable to other hunter- gather rates from the literature. Severe dental crowding is not typical in hunter-gatherer societies, and the low dental crowding rates recorded in many of these populations are

149 predominately comprised of mild/moderate crowding (Mockers 2003, Niswander 1964). These data suggest the possibility that the rate of mild/moderate crowding seen in these groups (approximately 25 percent) represents a normal occlusal distribution. In other words there is a general non-pathological predisposition for approximately 25 percent of individuals to exhibit mild to moderate crowding despite heavy masticatory use (this argument is discussed further in the Conclusions Chapter). On the contrary, since severe dental crowding is atypical in prehistoric groups, the rates of severe dental crowding in the Windover and Gauthier samples may be the result of environmental influences. Based on the current study, it seems likely that the severe crowding frequencies recorded at Gauthier and Windover are the result of similar cultural practices between these groups (most likely cultural practices involving the dentition). Further supportive evidence is provided by nonmetric, cranial metric, and attrition analyses, which are discussed below.

Nonmetric Analyses It is difficult to isolate genetic and environmental factors in dental crowding analyses. Environmental influences resulting in orofacial change might be a direct factor in dental crowding development but it also may accentuate previously masked genetic effects (Corruccini 1991). To investigate the relationship between the dentition and crania, nonmetric analyses were conducted between Windover and the comparative samples. Nonmetric analyses revealed similarities between Windover and Tick Island. Small sample sizes coupled by the low population frequencies of many of these traits complicated nonmetric tests of significance. Given small sample sizes, an aggregate sample of Little Salt Springs, Warm Mineral Springs and Bay West was employed to compare these sites to Windover. All dental nonmetric traits showed statistically similar (non-significant) rates between all of the comparative samples and Windover with the exception of maxillary second molar enamel extensions and tympanic dehiscence. Gauthier exhibited the greatest number of significant differences with Windover; significant differences were recorded for enamel extensions of maxillary second molars (p ≤ 0.001) and for tympanic dehiscence (p = 0.0157). Only Tick Island’s enamel

150 extensions frequency was similar to Windover. The presence of Uto-Aztecan premolars (UAP) and talon cusps, though not significantly different from any of the comparative samples, are notable because of their rarity. Talon cusps and UAPs were only observed in the Windover and Tick Island samples. Windover is the oldest population in the world demonstrating UAPs (Johnson et al. 2011) and the oldest population in the New World to exhibit talon cusps (Stojanowski et al. 2010). The presence of these traits in both of these populations further demonstrates dental similarities between Windover and Tick Island, and perhaps a genetic affinity between these two groups. Windover, however, was the only population with examples of concrescence and gemination. The incisor geminations, particularly their manifestation in the permanent dentition, and the concrescence are exceptionally rare dental anomalies (Nunes et al. 2002, Romito 2004), demonstrating the unique genetic composition of the Windover population. Windover was also the only population to exhibit cases of inca bones, including a rare partial bipartite variant. The presence of these rare traits, however, may be a reflection of the large size of the Windover sample since a larger sample size provides a greater opportunity to observe rare traits. Overall, the dental and cranial nonmetric trait frequencies were similar between the comparative samples and Windover, suggesting that these groups are more genetically similar to one another than distinct. Of the comparative samples under study, Tick Island demonstrated more similarities to Windover. However, dental crowding at Tick Island primarily consisted of mild/moderate dental crowding with only a 6 percent severe crowding rate. Although the severe dental crowding frequency at Windover is much higher than Tick Island, the mild/moderate crowding rate at Windover (22 percent) is comparable to Tick Island (26 percent). The nonmetric similarities between Windover and Tick Island support the possibility that mild/moderate crowding rates between these two samples represent a normal variation for this malocclusion. In other words, mild/moderate crowding (at rates of 0-26 percent) falls along the spectrum of normal occlusal variation within these groups and is not a disorder. Conversely, severe dental crowding appears to have a different etiology. Gauthier exhibited the most significant differences compared to Windover and was the only comparative sample to have a high rate of severe crowding. It is possible that these populations, though the least similar

151 genetically, share similar cultural practices influencing severe crowding development. These inferences are supported by attrition and cranial metric analyses and lend further support to the cultural (rather than genetic) etiology for severe crowding at Windover. Unfortunately, small sample sizes complicated tests of significance between these populations. To further assess genetic similarities between the Florida Archaic populations, future inter-cemetery biodistance analyses should be conducted between the samples to assess relatedness using dental and cranial metric and non-metric traits.

Tooth Size Tooth size and arch-size factors in dental crowding development were analyzed between all groups. Due to small sample sizes the comparative populations were combined and the aggregate sample was compared to Windover. This aggregation was justified based on the non-significant results of a Kruskal-Wallace nonparametric test between the comparative samples. Overall, there were high levels of variation in the samples for many of the teeth (particularly in the Windover sample), as reflected in the high standard deviations. Based on MD means comparisons between occlusion cohorts, there appeared to be no correlation between dental crowding and tooth size (particularly in the maxilla). Within Windover, significant differences between MD tooth width and sex cohorts were recorded for the maxillary canine (p = 0.033) and mandibular lateral incisor (p = 0.002). The maxillary canine exhibited higher MD averages in the males and the mandibular lateral incisor exhibited higher means in the females. The differences in the male and female canines are to be expected, as these are typically the most dimorphic teeth in sexually dimorphic primates. The mandibular lateral incisor was exceptional in that it exhibited a larger mean in the female group than the males. Slightly higher tooth width averages were recorded in the dental crowding group compared to the good occlusal group within Windover and the aggregate comparative sample. Although these higher averages are not consistent for all tooth types, only mandibular incisors (p ≤ 0.005) in the Windover sample exhibited significant differences between dental crowding and good occlusion groups. The differences in the other anterior teeth, combined with the high levels of variation for many of the teeth, may indicate that

152 the size of specific teeth, rather than the dentition in general, is contributing to dental crowding at Windover. These observations are similar to the results found by Mills (1964) and Radnzic (1988). Overall, it does not appear that tooth size alone is contributing to dental crowding development in either Windover or the comparative samples.

Arch Shape Inadequate space in the dental arcade for proper tooth eruption is known as tooth size arch size discrepancy (TSASD) and is the most common cause of dental crowding (Alt and Türp 1998b). To investigate arch shape factors on dental crowding etiology, arch depth and arch width averages were compared within Windover and between Windover and the comparative samples. Arch depth and arch width results show a high degree of variability between the samples (particularly for the mandible). The high variability among the seven samples may reflect measurement complications from heavy attrition (particularly interproximal attrition), antemortem tooth loss and postmortem damage. Despite the high variability of arch shape between these samples, tests of significance only showed non-significant differences between Windover and the comparative samples. The good preservation and large sample size of the Windover collection allowed more confident arch depth and arch width assessments within this sample. Assessments of arch depth and arch width between sex and occlusal cohorts in Windover showed correlations between dental crowding and increased arch depth (maxillary and mandibular) in the females (p ≤ 0.002) but not in the males. Correlations between dental crowding and reduced mandibular arch width in the Windover males (p = 0.026) and females (p = 0.040) were also observed in this study. The sex differences between arch shape and dental crowding correlations are interesting because males and females exhibit similar dental crowding severity frequencies. Correlations between dental arch elongation and arch width reduction are similar to findings by Kelly and Harvey (1977). They described the trend of palatal narrowing and deepening in industrialized groups with increasing frequencies of dental crowding. However, these results also contradict other tooth-size arch-length studies that found

153 reduced arch depth correlated with dental crowding (Bishara et al. 1995, Warren and Bishara 2001, Radnzic 1988). At Windover, a correlation between the reduction in mandibular arch width and dental crowding was recorded in both males and females. These data suggest similar environmental influences acting on the mandible are contributing to dental crowding development in both males and females. However differences in the maxillary and mandibular arch depth between the sexes demonstrates sex specific correlations between arch depth and dental crowding. Despite the equal distribution of dental crowding between males and females, these differences suggest gender-specific environmental effects on arch shape between males and females in Windover. It seems that practices unique to each sex, or differences in primary resource access, led to differential masticatory stressors affecting both arch depth and dental occlusion. There also appears to be a greater correlation with mandibular arch shape and dental crowding than in the maxilla. These results are similar to data recorded by von Cramon-Taubadel (2011) who found that mandibular shape was more susceptible to environmental factors (particularly masticatory stress) than the maxilla (von Cramon- Taubadel 2011).

Cranial Shape The Disuse Theory and masticatory function hypothesis have been used to explain and describe craniofacial changes, resulting from mastication stressors that lead to dental crowding. The shift from hard to soft textured foods is believed to be the primary force in the recent human evolutionary trend toward craniofacial gracialization consisting of more posteriorly placed faces and a general reduction in size and robusticity of faces, jaws and teeth (Carlson and Van Gerven 1977, Corruccini 1999, Larsen 1995, Rose and Roblee 2009). Carlson and Van Gerven (1977) developed the masticatory function hypothesis based on skeletal analyses of the Nubian transition from hunter-gatherer to agricultural subsistence practices. They argued that craniofacial gracilization was not a result of genetic factors but a direct result of a reduction in chewing stress during development. Corruccini (1999) built upon the disuse theory and the masticatory function hypothesis by

154 emphasizing that a reduction in chewing stress leads to under-development of maxillary and mandibular bones, creating insufficient space for proper tooth positioning. To investigate cranial shape factors in dental crowding etiology in the Florida Archaic, comparisons were conducted for cranial measurements between Windover and the comparative samples. Cranial measurements could not be conducted on the Tick Island skeletal collection because of heavy taphonomic damage. Due to small sample sizes and heavy postmortem damage, Little Salt Springs and Warm Mineral Springs were also excluded from these analyses. Therefore only Gauthier, Republic Groves, and Bay West could be compared to Windover. Overall, Republic Groves was the most statistically similar to Windover, followed by Gauthier (3 significant differences) and Bay West (5 significant differences) (see Tables 5.17 -5.19). Out of the 32 cranial measurements assessed, only ten cranial measures could be analyzed for Republic Groves, 11 for Bay West, and 27 for Gauthier. It is possible that the low number of measures analyzed between Republic Groves and Windover has obfuscated any cranial differences between these two groups. On the other hand, despite the low number of cranial measurements assessed between Windover and Bay West, a large number of significant differences were revealed. In regards to Gauthier, almost all 32 cranial measures could be compared to Windover and only three measurements were significantly different between the groups. The significant differences between Gauthier and Windover did not involve craniofacial metrics, with the exception of chin height. The other four comparative samples all demonstrated significant differences of numerous craniofacial metrics with the Windover sample. Although Gauthier demonstrates significant differences with Windover in nonmetric trait frequencies, this population appears to be one of the most similar to Windover in cranial shape. Due to the plasticity of cranial shape (Carlson and Van Gerven 1977, Corruccini 1999, Larsen 1995, Rose and Roblee 2009), these results may indicate that Windover and Gauthier, despite being genetically dissimilar, utilize similar cultural practices resulting in comparable cranial shape. When sex was controlled for in the Windover sample, sexual dimorphism was found to have a significant effect on cranial metrics in this population. Cranial metrics were found to be highly sexually dimorphic within the Windover population. Out of the

155 32 cranial metrics analyzed, 22 were significantly different between the sexes (see Table 4.26). All of the metrics (significant and non-significant) exhibit higher means in the male sample than the female. When cranial metrics were analyzed by sex between the good occlusion and dental crowding groups, only one metric was statistically significant between occlusal cohorts in the female group (mandibular angle), and three metrics were significant in the males (cranial base length, basio-prosthion length, and the occipital chord). The large degree of sexual dimorphism within these populations may have an effect on cranial metric comparisons between populations if sex is not accounted for. Unfortunately, small sample size prevents cranial metric analyses by sex in the comparative samples.

Attrition Analyses The heavy attrition levels recorded at Windover are similar to those observed in other hunter-gatherer populations. This makes the Windover malocclusion rate appear even more unusual because it contrasts with two long-held concepts of dental evolution: Disuse Theory (Price 1936) and the masticatory function hypothesis (the relationship between craniofacial shape and size and the functional mechanics of mastication) (Carlson and Van Gerven 1977). To investigate exogenous effects on the dentition in these populations, attrition analyses were conducted. Heavy chewing stress usually results in heavy attrition; thus, dental wear severity is a good indicator of the degree of chewing stress and dietary coarseness. The heavy masticatory requirements of hunter- gatherer diets lead to external abrasive stresses of the teeth resulting in increased dental attrition. Thus, one would expect to find dental traits and pathologies typical of prehistoric North American foraging societies in the populations under study. Attrition assessments were conducted to observe masticatory stress between each sample. Windover and Gauthier exhibited the greatest attrition means within each age cohort of the samples. Tests of significance between Windover and the comparative samples showed Gauthier to be the most similar to Windover in the young adult and middle-aged adult groups, while the aggregate sample is the most statistically different from Windover in the middle-aged and older adult groups. The similar (higher) rates of attrition between Windover and Gauthier suggest similar masticatory stressors between

156 these populations. This finding further supports an environmental etiology for the similar severe dental crowding frequencies between these samples. Attrition comparisons were also conducted within Windover between age cohorts as well as sex and occlusal cohorts to further investigate a possible correlation between attrition rates and dental crowding development. Dental attrition increased between each age cohort at a steady rate of approximately 50 points in both the maxilla and mandible. Overall, dental attrition averages were higher in the good occlusion groups than in the dental crowding groups, with the exception of the middle-aged and older adults. When attrition scores were analyzed by sex, tests of significance between occlusal cohorts in the male and female samples only showed non-significant differences between good occlusion and dental crowding groups (maxillary and mandibular). These results suggest that males and females show no dental wear distinction based on attrition scores alone. Detailed dental wear pattern analyses are needed to assess possible differences in extramasticatory use between the sexes.

Summary Previous studies do not support the notion that Windover practiced agriculture or horticulture (Quinn et al. 2008, Tuross et al. 1994); all research indicates that this population was a hunter-gatherer society that primarily subsisted on riverine resources. However, Windover exhibits agricultural-like dental crowding frequencies. The comparative samples in this study were chosen for their spatial and temporal similarities to Windover, as well as similar foraging practices (utilizing riverine or marine resources). Gauthier is the only Florida Archaic population that is similar to Windover for severe dental crowding (though at a much lower frequency). Gauthier also has the most similar cranial shape and attrition rates to Windover, indicating these two groups share similar environmental influences (most likely cultural resulting in extramasticatory tooth use) affecting cranial shape and dental wear. However, Gauthier also exhibits the most nonmetric significant differences with Windover suggesting that these populations are genetically distinct. Therefore, the similarity of severe dental crowding frequencies between these groups appears to be the product of similar environmental factors rather than genetic variants.

157 Conversely, Tick Island exhibited the most nonmetric similarities with Windover (inferring a genetic affinity between these populations) as well as statistically similar general dental crowding rates. Tick Island, however, exhibits lower attrition levels and severe crowding rates than Windover, suggesting that although the two groups are genetically similar, they are influenced by different environmental factors affecting dental wear and severe crowding development. Tick Island’s crowding rates (with the exception of one severe case) are predominately mild or moderate crowding scores. The mild/moderate dental crowding frequencies from Windover and the general dental crowding rates at Tick Island are more comparable, though slightly higher, to other prehistoric foragers. Therefore, it appears that there is a predisposition (i.e., a normal occlusal variation) in prehistoric and historic peoples to exhibit mild/moderate dental crowding rates as high as 26 percent. However, alongside this frequency of normal occlusal variation, environmental factors appear to have influenced dental crowding development as well (particularly severe dental crowding). It is possible that dental crowding originating from primarily environmental factors manifests along a continuum and therefore may also be contributing to a portion of the mild/moderate crowding frequencies. The greater correlations in Windover, between arch shape and dental crowding rather than tooth size, also support the likelihood of an environmental etiology for dental crowding since arch shape is more environmentally influenced and tooth size is more genetically determined. The significant differences in dental crowding rates between Windover and the comparative samples suggest that aquatic foraging subsistence does not affect dental crowding development. If environmental factors are the primary cause of dental crowding, then the high degree of dental crowding in the subadult sample implies that this practice begins early in life. Ethnographic and prehistoric studies (including studies of Windover) have shown that many adult behaviors associated with hunting and gathering begin during early childhood (Marlowe 2010, Thomas 2011). Tool-use involving the dentition is a possible environmental factor affecting dental crowding in these populations. Using teeth as tools (e.g., in the production of baskets, fibers, or cordage) would utilize different muscles than mastication and may alter craniofacial formation differently than heavy masticatory use. Ethnographic and

158 osteological studies of prehistoric and extant hunter-gatherer societies demonstrate frequent teeth-assisted tool use (Price 1936, Scott and Turner 1988) and habitual practices involving the teeth (Pereira and Evans 1975) in these societies. However, for tool-use to be a factor in dental crowding etiology, the practice(s) would have to lead to different craniofacial robustiticty/gracilization than is discussed in Carlson and Van Gerven’s (1977) masticatory function hypothesis, resulting in the odd combination of heavy attrition and dental crowding. If using teeth as tools is influencing severe dental crowding development in Windover (and Gauthier), it would have to include practices unique to the culture, since using teeth as all-purpose tools is common in hunter-gatherer societies and these groups usually exhibit low frequencies of dental crowding. Also, to affect craniofacial shape, these practices would have to occur frequently during development. The high rates of subadult dental crowding support this notion, implying that dental crowding is a product of tooth size (particularly permanent tooth size) arch size discrepancy rather than biological processes that occur with age, such as mesial drift. It is interesting that dental crowding appears to be mitigated by heavy attrition and compensatory tooth migrations. The Windover differences found in arch depth and arch width further support a TSASD etiology for severe crowding. Environmental factors appear to be affecting differences in female and male arch width and female arch depth. Both of these morphological changes correlate with dental crowding. However, the question also arises as to why only certain individuals develop severe dental crowding while others retain good occlusion? In this study, the greatest attrition level variability within Windover is recorded in the young adult age cohort (the age group with the greatest dental crowding frequency). This variability may reflect the first observable assessments of individual levels of labor and/or artistic divisions within these populations; even if these divisions are occurring while the individual is a subadult, it takes time for extramasticatory activities (resulting in tooth wear) to reflect in the dentition. It is unfortunate that detailed comparisons of craniofacial shape needed to test this hypothesis could not be undertaken with the cranial metrics methodology utilized in this study. Cranial shape assessments were further constrained by small sample sizes in the comparative samples. Future cranial morphometric studies should be conducted,

159 including the use of a digitizer or three-dimensional scanner to more accurately assess cranial form and robusticity differences between populations. In particular, future arch depth and arch width analyses should be conducted using the jaw corpus and alveolar regions instead of dentition in these prehistoric samples to reduce complications from heavy attrition and antemortem tooth loss. Digitization and 3-dimensional models also would be more effective to demonstrate morphological differences between arch shape and dental crowding severities. To test for non-masticatory differences between the samples, detailed analyses of dental wear patterns (e.g., flat wear, cupped wear, sloped wear, etc.) need to be conducted to address the influences of tool-use on dental crowding development. Few studies have concentrated their efforts on the effects of habitual behaviors and tool-use reflected in the dentition (Scott and Turner 1988). This is a promising line of dental anthropological research that may provide insights into these intriguing Florida Archaic populations.

160

CHAPTER 7 CONCLUSION

Anterior dental crowding has a multifactorial etiology, consisting primarily of environmental influences and polygenic factors. High rates of dental crowding are most prominent in societies practicing an agricultural subsistence and are rare in foraging societies (Begg 1954, Corruccini 1999, Larsen 1995, Price 1936, Rose Roblee 2009). This trend has been associated with masticatory stress and its effects on the craniofacial and dental arch shape, as described in Carlson and Van Gerven’s (1977) masticatory function hypothesis and Price’s (1936) Disuse Theory. This study has investigated the high rates of dental crowding in the Early Archaic Windover sample from Florida. Comparative tests were conducted on other Early and Middle Archaic samples from Florida including: Little Salt Springs, Gauthier, Harris Creek at Tick Island, Warm Mineral Springs, and Bay West.

Methodology and Future Research The methodology developed in this study should prove useful to future research projects involving the collection and analyses of similar data sets. Small sample sizes and taphonomic damage complicated many of the analyses in this study. To further assess dental crowding etiology in the Florida Archaic, future investigations employing geometric morphometric analyses for cranial shape, in particular arch size, should be assessed using digitization methodologies as well as three-dimensional scans. To avoid complications from dental attrition, landmarks should be assessed from the corpus of the maxillary and mandibular arcades rather than the dentition itself. In addition, dental wear patterns should also be recorded from these populations to investigate non-masticatory influences on dental crowding development. This study also highlights the necessity of reporting frequencies of specific dental conditions, such as anterior dental crowding, rather than lumping a host of different dental malpositions under the term malocclusion. This is particularly important because different dental conditions have different etiologies (Harris 2008). Reporting specific conditions allows for direct comparisons between

161 studies, which may further uncover factors involved their development. For instance, in this study there is evidence for different developmental origins within dental crowding, resulting in differential dental crowding severity. It is imperative to evaluate standardized assessments for different dental conditions and to avoid using the term malocclusion as a blanket variable in both orthodontic and dental anthropological studies.

Research Results In this study, analyses of dental crowding frequencies, cranial and jaw shape, tooth-size, and dental alterations (masticatory and non-masticatory) were conducted in the Windover and comparative samples. The primary goal of this study was to assess dental crowding within the Florida Archaic peoples and to ascertain the primary factors influencing its etiology.

Primary Hypothesis: The incidence of dental crowding in the Windover population is anomalous compared to other hunter-gatherer populations including Florida Archaic peoples. This study supports the primary hypothesis: the Windover Early Archaic sample presents an unusual case of high dental crowding frequencies in a hunter-gatherer society. The rate of malocclusions in this sample is anomalous when compared to other hunter- gatherer populations - both prehistoric and modern (Brace 1988, Larsen 1995, Sciulli 1997). Overall, Windover exhibits rates nearly three-times higher than other prehistoric groups, rates that are more comparable to agricultural societies than foragers. When dental crowding is analyzed according to severity within Windover, severe crowding frequencies are similar to rates recorded in agricultural societies. However, mild/moderate crowding rates within Windover are comparable to other hunter-gatherer societies recorded in the literature and in the comparative samples. These data demonstrates that the Windover population is unique among hunter-gatherers in severe crowding frequencies but not in mild/moderate dental crowding rates. Amongst age cohorts in the Windover sample dental crowding rates are the highest in the juvenile and young adult groups and exhibit a trend of decreasing dental crowding frequency with age in the young adult though older adult groups. This is the

162 result of two factors: 1) complications of dental crowding assessment due to heavy dental wear with age, and 2) the corrective effects of heavy dental attrition, resulting from increased interproximal space coupled with migratory processes (e.g., mesial drift and lingual tipping). Dental crowding rates were found to be similar between sex cohorts within the Windover sample, suggesting that prescribed cultural gender roles did not have a differential effect on dental crowding rates. Compared to other Florida Archaic samples, Windover only exhibited similar rates of total dental crowding to the Tick Island sample, and similar severe dental crowding rates to the Gauthier sample. However, mild/moderate crowding rates were similar between all groups.

Genetic Sub-hypothesis: The high dental crowding rates recorded at Windover are the result of predominant genetic factors with high degrees of phenotypic expression despite heavy attrition and adequate arch size. In this study, this sub-hypothesis is partially accepted. In the samples analyzed, as well as comparative samples in the literature, it was found that approximately 26% of mild/moderate crowding frequencies represented normal occlusion variants that are non- pathological. Normal variation is a condition of organic existence and is non-pathological or incidental; it is a reflection of the variation (within definite limits) inherent in organisms under basic conditions (Hrdlička 1934: 253). While normal variation is non- pathological and is not governed by any single gene, it is the product of the overall normal genetic variability within that organism. To test for genetic factors in dental crowding etiology, nonmetric analyses were conducted to assess relatedness between Windover and the comparative samples. Measurements of tooth size and arch size were also employed to investigate tooth size arch size discrepancy (TSASD) within this population, a condition found to be the product of environmental influences (Beecher and Corruccini 1981, Brown and Maeda 2004, Carlson and Van Gerven 1977, Cassidy et al. 2010, Conceição and Cardoso 2010, Rose and Roblee 2009, Schoenau and Fricke 2008, von Cramon-Taubadel 2011). Overall mesiodistal tooth width means within Windover did not exhibit significant correlations between dental crowding and tooth size but correlations were observed between longer

163 arch depth means in females and shorter arch width means in males. These findings suggest that dental crowding in Windover has a greater association with tooth size arch size discrepancy than tooth size alone. Mild/moderate crowding rates are similar between Windover and other prehistoric populations (including the comparative Florida Archaic samples) suggesting that a low frequency of mild/moderate crowding (ranging from 0-26 percent) is a normal occlusal variation found in many prehistoric groups. This is supported by the nonmetric and dental crowding similarities between Windover and Tick Island. Tick Island was the most similar to Windover in nonmetric frequencies and is also the only sample similar to Windover for total dental crowding frequency. Tick Island’s crowding rates (with the exception of one severe case) are predominantly mild or moderate crowding scores. Tick Island exhibits lower attrition levels and severe crowding rates than Windover, suggesting that although the two groups are genetically similar, they are influenced by different environmental factors affecting dental wear and severe crowding development. The mild/moderate crowding similarities between Windover and Tick Island, therefore, most likely reflect a normal occlusal variant while severe dental crowding presents a differential etiology.

Environmental Sub-hypothesis: The high dental crowding rates recorded at Windover are the product of unique cultural practices: resulting in craniofacial changes leading to inadequate arch size to accommodate the size of the teeth. This study supports the environmental sub-hypothesis with regard to severe dental crowding in the Florida Archaic populations. This was found to be particularly true for the Windover and Gauthier samples. The plasticity of cranial and orofacial shape are described in the Disuse Theory and masticatory function hypothesis which maintain that changes to the cranium can occur as a result of mastication stress (or the relaxation of that stress) and can lead to dental crowding (Carlson and Van Gerven 1977, Corruccini 1999, Larsen 1995, Price 1936, Rose and Roblee 2009). Unlike tooth shape, craniofacial development can be greatly influenced by environmental factors. Increased stimulation to the jaws (via masticatory or non-masticatory stressors) can alter the dental arch shape and lead to

164 inadequate jaw space for the size of the teeth (Corruccini and Potter 1980). My research supports an environmental etiology for this malocclusion, as evidenced by the correlations in Windover between arch shape and dental crowding, rather than tooth size. It is possible that only severe crowding is effected by TSDAS in this sample; unfortunately, small sample sizes made it impossible to assess correlations between arch shape and dental crowding severity ranks to investigate differences between mild/moderate and severe crowding. My suggestion that severe dental crowding is predominately environmentally influenced is also supported by the similarities between Windover and Gauthier. Gauthier is the only comparative sample to exhibit statistically similar severe crowding rates to Windover. Gauthier is also the most similar to Windover for cranial shape and attrition rates, but was the most dissimilar dataset for nonmetric frequencies. These results suggests that Gauthier and Windover are genetically distinct but share similar masticatory stresses, further supporting an environmental etiology for the similar severe dental crowding frequencies between these samples. Another example of environmental influence on dental crowding development are the differences in the maxillary and mandibular arch depth between males and females within Windover. These differences showed sex specific correlations between arch depth and dental crowding. Though males and females exhibited similar dental crowding frequencies, it appears that differences in primary resource access or gender-specific cultural practices led to different stressors that affected arch depth and dental crowding development. There was also a correlation between a reduction in arch width and increased dental crowding in both sexes. Interestingly in this study, Windover (one of the few known cultures in the world to have practiced underwater burial) appeared to share the greatest genetic affinity not with groups practicing similar mortuary pond burial techniques (Little Salt Springs, Republic Groves, Bay West) but rather with a group who utilized shell middens to inter their dead (Tick Island). The genetic affinity between Tick Island and Windover is particularly evident in the presence of rare traits such as the Uto-Aztecan premolar and talon cusp variant within these samples. It is also interesting that data from this study suggest that similar environmental/cultural factors that influenced cranial shape, dental wear, and possibly severe dental crowding development in Windover were found in the

165 Gauthier sample; Gauthier represents a temporarily more recent skeletal assemblage buried along elevated sandy ridges. Therefore it appears that practices, such as burial techniques, cannot alone infer genetic and cultural similarities between groups from the Florida Archaic.

Summary Dental crowding development within Florida Archaic populations appears to have different primary influences based on crowding severity. Mild/moderate crowding in the Windover and comparative samples is similar to rates recorded in other prehistoric foraging societies. Therefore, this malocclusion might be a normal occlusal variant within these samples. However, severe dental crowding is unique amongst hunter-gatherer societies and indicates a predominantly environmental etiology. Despite statistically similar severe crowding rates between Windover and Gauthier, the crowding frequency at Windover is more comparable to agricultural societies while Gauthier’s is not. Yet Windover predates the onset of agriculture in Florida by at least 5,500 years. It is possible that the environmental influence on severe dental crowding development is the result of cultural and sociological peculiarities of an isolated society. Despite the similarities between Windover, Tick Island and Gauthier, Windover exhibits a much greater frequency of dental crowding (particularly severe dental crowding) than these samples and shows more distinctions from the comparative samples than similarities (including rare genetic anomalies and unique dental wear patterns). This suggests that although Windover has a genetic affinity with Tick Island, it still exhibits cultural and genetic distinctions from these Florida Archaic groups. In particular, non-masticatory cultural practices (e.g., using teeth as tools) might alter cranial-facial formation differently than is discussed in Carlson and Van Gerven’s (1977) masticatory function hypothesis; these practices might have caused the odd combination of heavy attrition and severe dental crowding. It should be emphasized that dental crowding does not increase in frequency simply because of a change in subsistence. Dental crowding development is a product of a complex of masticatory processes and non-masticatory practices adopted by a group and is not necessarily governed solely by the overarching subsistence practice itself. There are other factors to consider rather than simply looking at the food acquisition of a

166 society. Windover is exemplary of a non-agricultural group whose high dental crowding frequencies are attributable to cultural practices. A comprehensive view of what activities affect dental crowding is necessary to isolate influential factors governing its development.

167

APPENDIX I Dental Nonmetric Frequencies

Republic Tick Bay Trait Sample Windover LSS Gauthier WMS Groves Island West % 55% 0 40% 75% 80% 100% 100% Bilateral winging UI1 Sample # 36 0 5 4 10 1 2 (+ = ASU 1) Present # 20 0 2 3 8 1 2 % 97% 0 60% 67% 100% 100% 100% Shoveling UI1 Sample # 31 0 5 3 19 1 2 (+ = ASU 2-6) Present # 30 0 3 2 19 1 2 % 34% 0 14% 0 31% 100% 0 Double Shoveling UI1 Sample # 38 0 7 4 13 1 2 (+ = ASU 2-6) Present # 13 0 1 0 4 1 0 % 7% 0 0 0 13% 0 0 Peg-reduced UI2 Sample # 45 0 8 4 15 1 2 (+ = ASU P or A) Present # 3 0 0 0 2 0 0 % 24% 0 0 50% 40% 0 0 Tuberculum dentale UI1 Sample # 33 0 4 2 15 1 2 (+ = ASU 2-6) Present # 8 0 0 1 6 0 0 % 34% 0 0 0 60% 100% 50% Tuberculum dentale UI2 Sample # 41 0 5 2 15 1 2 (+ = ASU 2-6) Present # 14 0 0 0 9 1 1 % 8% 0 0 67% 33% 0 67% Tuberculum dentale UC Sample # 13 0 7 3 18 1 3 (+ = ASU 2-6) Present # 1 0 0 2 6 0 2 % 3% 0 17% 0 8% 0 0 Tuberculum dentale LC Sample # 35 0 6 3 12 1 0 (+ = ASU 2-6) Present # 1 0 1 0 1 0 0 % 26% 0 33% 0 20% 0 0 Distal accessory ridge UC Sample # 23 0 3 1 10 1 2 (+ = ASU 2-5) Present # 6 0 1 0 2 0 0

168 Republic Tick Bay Trait Sample Windover LSS Gauthier WMS Groves Island West % 29% 0 25% 0 33% 100% 0 Distal accessory ridge LC Sample # 17 0 4 0 6 1 0 (+ = ASU 2-5) Present # 5 0 1 0 2 1 0 % 8% 0 0 0 14% 0 0 UtoAztecan Premolar UP3 Sample # 25 1 1 1 14 1 0 (+ = ASU P or A) Present # 2 0 0 0 2 0 0 % 0 0 0 0 15% 0 0 C3 UM1 Sample # 27 1 1 1 13 0 0 (+ = ASU 2-5) Present # 0 0 0 0 2 0 0 % 29% 0 0 0 10% 0 0 C3 UM2 Sample # 24 0 2 1 10 0 1 (+ = ASU 2-5) Present # 7 0 0 0 1 0 0 % 15% 0 0 0 23% 0 0 C5 UM1 Sample # 27 1 1 1 13 0 0 (+ = ASU 2-5) Present # 4 0 0 0 3 0 0 % 8% 0 50% 0 17% 0 0 C5 UM2 Sample # 2 0 1 0 2 0 0 (+ = ASU 2-5) Present # 24 0 2 1 12 0 1 % 6% 0 0 0 10% 0 0 Carabelli’s cusp UM1 Sample # 66 3 16 5 20 1 6 (+ = ASU 2-7) Present # 4 0 0 0 2 0 0 % 2% 0 0 0 0 0 0 Carabelli’s cusp UM2 Sample # 66 2 17 0 16 1 6 (+ = ASU 2-7) Present # 1 0 0 0 0 0 0 % 0 0 0 0 0 0 0 Carabelli’s cusp UdM2 Sample # 13 0 1 0 1 0 0 (+ = ASU 2-7) Present # 0 0 0 0 0 0 0 % 34% 0 12% 20% 42% 0 0 Enamel extensions UM1 Sample # 77 2 15 5 12 0 0 (+ = ASU 1-3) Present # 26 0 2 1 5 0 0

169 Republic Tick Bay Trait Sample Windover LSS Gauthier WMS Groves Island West % 70% 0 13% 20% 45% 0 0 Enamel extensions UM2 Sample # 73 1 16 5 11 0 0 (+ = ASU 1-3) Present # 51 0 2 1 5 0 0 % 52% 0 0 0 0 0 0 Enamel extensions LM1 Sample # 46 0 3 0 7 0 0 (+ = ASU 1-3) Present # 24 0 0 0 0 0 0 % 68% 0 0 0 0 0 0 Enamel extensions LM2 Sample # 47 0 2 0 9 0 0 (+ = ASU 1-3) Present # 32 0 0 0 0 0 0 % 6% 0 0 0 5% 0 0 Parastyle UM1 Sample # 77 3 16 5 19 1 5 (+ = ASU 1-5) Present # 5 0 0 0 1 0 0 % 9% 0 0 0 0 0 20% Parastyle UM2 Sample # 68 2 17 5 15 1 5 (+ = ASU 1-5) Present # 6 0 0 0 0 0 1 % 6% 0 0 0 0 0 0 Deflecting wrinkle UM1 Sample # 16 0 0 1 0 0 0 (+ = ASU 2-3) Present # 1 0 0 0 0 0 0 % 9% 0 0 0 0 0 0 Deflecting wrinkle LM1 Sample # 11 0 2 0 0 0 0 (+ = ASU 2-3) Present # 1 0 0 0 0 0 0 % 20% 0 0 0 0 0 0 Deflecting wrinkle Ldm2 Sample # 5 0 1 0 0 0 0 (+ = ASU 2-3) Present # 1 0 0 0 0 0 0 % 0 0 0 0 0 0 0 Cusp 7 LM1 Sample # 20 0 3 0 7 0 0 (+ = ASU 2-4) Present # 0 0 0 0 0 0 0 % 0 0 0 0 0 0 0 Cusp 7 LM2 Sample # 19 0 2 0 9 0 0 (+ = ASU 2-4) Present # 0 0 0 0 0 0 0

170 Republic Tick Bay Trait Sample Windover LSS Gauthier WMS Groves Island West % 4% 67% 7% 0 7% 0 0 Protostylid LM1 Sample # 47 3 14 3 13 0 0 (+ = ASU 2-6) Present # 2 2 1 0 1 0 0 % 11% 50% 8% 0 0 0 0 Protostylid LM2 Sample # 44 2 13 4 12 0 0 (+ = ASU 2-6) Present # 5 1 1 0 0 0 0 % 27% 0 0 0 40% 0 0 Anterior fovea LM1 Sample # 15 0 0 0 5 1 0 (+ = ASU 2-4) Present # 4 0 0 0 2 0 0 % 61% 0 0 0 56% 0 0 Anterior fovea LM2 Sample # 18 0 0 0 9 1 0 (+ = ASU 2-4) Present # 11 0 0 0 5 0 0 % 60% 0 0 0 0 0 0 Anterior fovea Ldm2 Sample # 5 0 0 0 0 0 0 (+ = ASU 2-4) Present # 3 0 0 0 0 0 0

171 APPENDIX II Cranial Nonmetric Frequencies

Republic Tick Bay Trait Sample Windover LSS Gauthier WMS Groves Island West % 0 0 0 0 0 25% 0 Metopic Suture Sample # 95 6 15 4 1 4 6 (+ = B&U 2) Present # 0 0 0 0 0 1 0 % 90% 100% 100% 100% 0 100% 100% Supraorbital Notch Sample # 93 6 14 4 0 4 7 (+ = B&U 1-4) Present # 84 6 14 4 0 4 7 % 28% 50% 60% 100% 100% 0 43% Supraorbital Foramen Sample # 92 6 15 1 1 4 7 (+ = B&U 1-2) Present # 44 3 9 2 2 0 3 % 50% 50% 20% 0 0 33% 14% Infraorbital Suture Sample # 52 6 5 0 2 3 7 (+ = B&U 1-2) Present # 26 3 2 0 0 1 1

Multiple Infraorbital % 17% 33% 0 0 0 50% 75% Foramina Sample # 46 6 4 0 2 2 4 (+ = B&U 1-3) Present # 8 2 0 0 0 1 3 % 89% 67% 57% 75% 100% 100% 80% Zygomatico-facial Foramina Sample # 73 6 7 4 2 2 5 (+ = B&U 1-6) Present # 65 4 4 3 2 2 4 % 73% 0 50% 75% 100% 75% 0 Parietal Foramen Sample # 84 1 14 4 1 5 6 (+ = B&U 1-2) Present # 61 0 7 3 1 3 0 % 6% 0 7% 25% 0 0 0 Epiteric Bone Sample # 78 1 15 4 0 3 5 (+ = B&U 1) Present # 5 0 1 1 0 0 0 % 12% 0 0 0 0 0 0 Coronal Ossicle Sample # 86 1 14 2 0 4 5 (+ = B&U 1) Present # 10 0 0 0 0 0 0 % 0 0 0 0 0 0 0 Bregmatic Bone Sample # 90 2 14 4 0 4 5 (+ = B&U 1) Present # 0 0 0 0 0 0 0 % 6% 0 0 0 0 0 0 Sagittal Ossicle Sample # 80 2 15 2 0 4 5 (+ = B&U 1) Present # 5 0 0 0 0 0 0

172

Republic Tick Bay Trait Sample Windover LSS Gauthier WMS Groves Island West % 14% 0 15% 0 0 0 0 Apical Bone Sample # 86 2 13 4 0 4 5 (+ = B&U 1) Present # 12 0 2 0 0 0 0 % 49% 100% 18% 0 0 0 20% Lambdoid Ossicle Sample # 80 2 11 3 0 3 5 (+ = B&U 1) Present # 39 2 2 0 0 0 1 % 8% 0 18% 0 0 0 0 Asterionic Bone Sample # 87 2 11 2 0 4 4 (+ = B&U 1) Present # 7 0 2 0 0 0 0 % 18% 0 20% 0 0 0 0 Occipitomastoid Ossicle Sample # 83 2 10 3 0 4 4 (+ = B&U 1) Present # 15 0 2 0 0 0 0 % 7% 50% 8% 0 0 0 0 Parietal Notch Bone Sample # 87 2 12 3 0 4 4 (+ = B&U 1) Present # 6 1 1 0 0 0 0 % 2% 0 0 0 0 0 0 Inca Bone Sample # 91 2 15 4 1 4 6 (+ = B&U 1-4) Present # 2 0 0 0 0 0 0 % 83% 100% 0 0 0 0 83% Condylar Canal Sample # 52 1 11 0 1 1 6 (+ = B&U 1) Present # 43 1 0 0 0 0 5 % 17% 0 20% 0 0 0 0 Divided Hypoglossal Canal Sample # 54 2 10 1 1 1 4 (+ = B&U 1-4) Present # 9 0 2 0 0 0 0

Right Flexure of Superior % 24% 0 40% 0 0 67% 83% Saggital Suture Sample # 82 1 5 2 0 3 6 (+ = B&U 1) Present # 20 0 2 0 0 2 5

Left Flexure of Superior % 9% 100% 0 0 0 33% 17% Saggital Suture Sample # 82 1 5 2 0 3 6 (+ = B&U 2) Present # 7 1 0 0 0 1 1

Bifurcate Flexure of Superior % 67% 0 60% 100% 0 0 0 Saggital Suture Sample # 82 1 5 2 0 3 6 (+ = B&U 3) Present # 55 0 3 2 0 0 0

173 Republic Tick Bay Trait Sample Windover LSS Gauthier WMS Groves Island West % 21% 0 29% 0 0 0 14% Foramen Ovale Incomplete Sample # 61 3 7 2 0 1 7 (+ = B&U 1-2) Present # 13 0 2 0 0 0 1

Foramen Spinosum % 32% 33% 43% 33% 0 0 14% Incomplete Sample # 65 3 7 3 0 1 7 (+ = B&U 1-2) Present # 21 1 3 1 0 0 1 % 43% 0 57% 100% 0 0 0 Pterygo-spinous Bridge Sample # 46 1 7 1 0 1 5 (+ = B&U 1-3) Present # 20 0 4 1 0 0 0 % 11% 0 40% 0 0 0 0 Pterygo-alar Bridge Sample # 44 1 5 0 0 1 4 (+ = B&U 1-3) Present # 5 0 2 0 0 0 0 % 68% 100% 27% 75% 100% 100% 75% Tympanic Dihiscence Sample # 76 2 11 4 1 3 4 (+ = B&U 1-2) Present # 52 2 3 3 1 3 3 % 0 0 0 0 0 0 0 Auditory Exostosis Sample # 93 2 13 4 0 4 4 (+ = B&U 1-3) Present # 0 0 5 0 0 0 0 % 51% 50% 100% 75% 0 33% 0 Temporal Mastoid Foramen Sample # 91 2 2 4 0 3 4 (+ = B&U 1) Present # 46 1 2 3 0 1 0 % 15% 0 0 25% 0 0 0 Sutural Mastoid Foramen Sample # 91 2 2 4 0 3 4 (+ = B&U 2) Present # 14 0 0 1 0 0 0 % 2% 0 0 0 0 0 0 Occipital Mastoid Foramen Sample # 91 2 2 4 0 3 4 (+ = B&U 3) Present # 2 0 0 0 0 0 0

Both Sutural and Temporal % 11% 50% 0 0 0 33% 75% Mastoid Foramen Sample # 91 2 2 4 0 3 4 (+ = B&U 4) Present # 10 1 0 0 0 1 3

Both Occipital and Temporal % 2% 0 0 0 0 33% 0 Mastoid Foramen Sample # 91 2 2 4 0 3 4 (+ = B&U 5) Present # 2 0 0 0 0 1 0

174 Republic Tick Bay Trait Sample Windover LSS Gauthier WMS Groves Island West % 8% 25% 13% 22% 0 50% 0 Mental Foramen Sample # 88 4 15 9 10 2 1 (+ = B&U 2-3) Present # 7 1 2 2 0 1 0 % 0 0 0 0 0 0 0 Mandibular Torus Sample # 89 4 18 9 9 3 3 (+ = B&U 1-3) Present # 0 0 0 0 0 0 0

Mylohyoid Bridge Near % 4% 0 8% 0 0 0 0 Mandibular Foramen Sample # 88 4 13 7 4 3 1 (+ = B&U 1) Present # 4 0 1 0 0 0 0

Mylohyoid Bridge Center of % 23% 25% 31% 14% 25% 0 0 Groove Sample # 88 4 13 7 4 3 1 (+ = B&U 2) Present # 20 1 4 1 1 0 0

Double Mylohyoid Bridge % 3% 0 0 29% 0 0 0 with hiatus Sample # 88 4 13 7 4 3 1 (+ = B&U 3) Present # 3 0 0 2 0 0 0

Double Mylohyoid Bridge % 3% 0 0 29% 0 0 0 without hiatus Sample # 88 4 13 7 4 3 1 (+ = B&U 4) Present # 3 0 0 2 0 0 0

175

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BIOGRAPHICAL SKETCH

Kathryn O’Donnell Miyar began her college education as a classically trained pianist pursuing a degree in piano performance at the Cincinnati Conservatory and later at the University of Miami. During the third year of her undergraduate education she began a second major in anthropology. While at the University of Miami she participated in forensic excavations with the Miami-Dade police department, assisted with archaeological excavation and historic preservation in Key West at the Audubon House, and conducted cultural anthropological fieldwork analyzing population immigration and integration on the island of Santa Cruz in the Galapagos, Ecuador. In 2004 she received her B.A., in music and anthropology, from the University of Miami in Coral Gables, FL. During her graduate studies Miyar assisted on a Late Pleistocene Neanderthal Excavation in Pinilla del Valle, Spain, under the direction of Dr. Arsuaga and collected cranial measurements from crania casts at the American Museum of Natural History in New York City to test the taxonomic integrity of Homo heidelbergensis. Her master’s thesis focused on the epidemiology of spina bifida in Ireland. She conducted analyses of skeletal remains at the Archaeological Developmental Services Ltd. Kells laboratory and at the National Museum of Ireland for her master’s thesis research. Miyar received her M.A. in 2007 from Florida Atlantic University in Boca Raton, FL. During her doctoral studies at Florida State University, Miyar conducted stable isotopic analyses at the National High Magnetic Field Laboratory in Tallahassee, FL regarding dietary changes from prehistoric Florida populations through modern times. She also conducted archival research at the National Archives of Ireland and collected residue from Famine Pots for trace chemical analyses as preliminary research investigating the presence of mycotoxins during the Irish potato famine. In her fourth year she assisted Mr. Johnson and Dr. Stojanowski in the investigation of the presence of the Uto-Aztecan nonmetric variant of maxillary 3rd premolars in Florida Archaic populations. While at Florida State University she was the instructor for the Human Osteology course (ANT 4525). Miyar continues to work as an osteologist at the Southeast Archeological Center in the NAGPRA division.

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