A Diachronic Assessment of Health and Disease from the Adult Dentition of the Naton Beach Burial Complex in Tumon Bay, Guam

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Title:
A Diachronic Assessment of Health and Disease from the Adult Dentition of the Naton Beach Burial Complex in Tumon Bay, Guam
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1 online resource (279 p.)
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english
Creator:
Parr, Nicolette M
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University of Florida
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Degree:
Doctorate ( Ph.D.)
Degree Grantor:
University of Florida
Degree Disciplines:
Anthropology
Committee Chair:
Warren, Michael W
Committee Members:
Krigbaum, John S
Daegling, David
Steadman, David W
Schmidt, Christopher

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Subjects / Keywords:
anthropology -- carious-lesions -- dental-reduction -- guam -- linear-enamel-hypoplasia
Anthropology -- Dissertations, Academic -- UF
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Anthropology thesis, Ph.D.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
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Abstract:
The current study is a diachronic investigation of the prehistoric Chamorro in Guam to assess health and disease patterns over time.  The transition from the Pre-Latte to Latte periods displays a shift from horticultural to early agricultural practices;accompanying changes include increased population size and technologically advanced food processing and preparation techniques.  Likewise, these changes occur concomitantly with large-scale environmental and climatic fluctuations.  It is predicted that the cultural and environmental shifts will be accompanied by biological ones, due to increased stress levels associated with malnutrition, limited access to resources, and increased prevalence of disease. Analyses of odontometrics ,linear enamel hypoplasias, and carious lesions were performed and analyzed in concert with skeletal data collected by other researchers to construct a health profile of the prehistoric populations in Guam. Expected results include dental reduction over time coupled with an increase in linear enamel hypoplasias and carious lesions. The dentition display an 8% decrease from the Pre-Latte to Latte periods.  Increased reliance on starchy crops would have lead to selection for smaller dentition to minimize carious lesions. Additionally, sophistication in food processing techniques decreases the force necessary to break down tough food, leading to reduced functional demands of the masticatory apparatus.  Thus, this finding is best explained by a combination of the Selective Compromise Effect and Masticatory-Functional Hypothesis. Significant differences in linear enamel hypoplasia expression are also noted with an increase over time.  While not significant, the data suggests that there may have been differential access to resources as a result of gender roles associated with food procurement, where the females in the Latte period were much more highly susceptible to physiological stress than the males. Carious lesions are also significantly different over time; however, these findings do not follow the predicted pattern.  Caries frequency decrease over time likely due to the cultural practice of betel-nut chewing, which has cariostatic properties, in the Latte period. This study expands on the current knowledge of prehistoric health in Guam by demonstrating an overall decrease in health over time as a result of climatic instability and subsequent dietary transitions.
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In the series University of Florida Digital Collections.
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Includes vita.
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by Nicolette M Parr.
Thesis:
Thesis (Ph.D.)--University of Florida, 2012.
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Adviser: Warren, Michael W.
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RESTRICTED TO UF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE UNTIL 2014-08-31

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1 A DIACHRONIC ASSESSMENT OF HEALTH AND DISEASE FROM THE ADULT DENTITION OF THE NATON BEACH BURIAL COMPLEX IN TUMON BAY, GUAM By NICOLETTE M. PARR A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2012

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2 2012 Nicolette Maria Luney Parr

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3 To my mother and father for always providing me with endless support and encouragement

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4 ACKNOWLEDGMENTS First and foremost I would like to thank my committee members, Mike Warren, John Krigbaum, Dave Daegling, and David Steadman for yo ur ti reless advice and patience and Chris Schmidt for introducing and instilling in me a love for teeth. Drs. Warren and Krigbaum have been my mentors and friends since my undergraduate years and never hesited to give me guidance and encouragement along the wa y. They took the gamble to accept me back as a Ph.D. student for that, I will be forever grateful. This study would have never been conducted had it not been for the opportunities ites, and Garcia and Associates. I want to express my deepest gratitude to Pat and Nicole for introducing me to Guam and the Pacific region and for the countless dinners, drinks, birthday celebrations, holidays, and hours of laughter we shared together Your kindness and generosity will never be forgotten. Am am also lucky to have made wonderful friends in Guam: Jamey Cyrus, Marie, Justin and Patrick to whom I am thankful for so many great adventures and for reminding me to explore the island, eat goo d food, and spend time underwater when the going got tough. I am particularly thankful to Dave DeFant for access to the Naton Beach skeletal collection, as well as Cherie Walth, Sandy Yee, Lynn Leon Guerro, and Michelle Christy, from SWCA, for providing me with endless amounts of information regarding the Naton Beach site access to their library, and workspace in which I spent ma n y months I am also grateful to a number of people who have worked in Guam for many years and have provided me with valuable information, feedback, and thoughtful conversation regarding my research: Gary Heathcote, Rona Ikehara Quebral, Judith Amesbury, Rosanna

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5 Barcinas, Boyd Dixon, Jonn Peterson, Anne Stodder, Lawrence Cunningham, Lon Bulgrin, Michael Pietrusewsky, Br uce Anderson, Vince Sava, Michele Douglas, and Joanne Eakin. I would like to thank my fellow grad students from the Pound Lab: Laurel, Katie, Traci, Caroline, Sarah, Al lysha, Carrie, and Kristina. You have been with me through many highs and lows, have endured my caffeine induced hysteria and listened to my usually pointless stories. I am so grateful for all you have done for me from reading a myriad of grant proposals to giving me statistical advice, which does not even begin to enumerate it all A special thanks goes out to Carlos Zambrano for throughoughly reviewing this entire document and for putting up with me for six more years and then some To Lulu, Kim, Viviana, Amber, and Leila: much of who I am today I owe to havi ng had you in my life; thank you for always being there for me, no matter the distance between us. I am deeply indebted to my many family members who supported me even when they thought I would always be a perpetual student, particula rly Sonia, Casper, Piedad, and Jennifer. Last but not least, I would like to give my most heartfelt thanks to my mother and father who continuously provided me with unconditional love and support. You have always encouraged me to never stop learning and f or teaching me how to travel, eat good food, and to love lif e I could not have done this without you This dissertation was supported in part by the William R. Maples Memorial Scholarship, O. Ruth McQuown Supplemental Award, Wentworth Foundation William M. Goza Fellowship, Delores Auzenne Minority Dissertation Award, Ellis R. Kerley Forensic Sciences Foundation Scholarship, and the CA Pound Human Identification Laboratory.

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6 TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................ ................................ ................................ .. 4 LIST OF TABLES ................................ ................................ ................................ ............ 9 LIST OF FIGURES ................................ ................................ ................................ ........ 14 LIST OF ABBREVIATIONS ................................ ................................ ........................... 15 ABSTRACT ................................ ................................ ................................ ................... 17 CHAPTER 1 INTRODUCTION ................................ ................................ ................................ .... 19 Theoretical Framework ................................ ................................ ........................... 21 Biocul tural Approach to Bioarchaeology ................................ ........................... 22 Stress Models ................................ ................................ ................................ ... 23 Purpose and Research Objectives ................................ ................................ ......... 27 Objectives and Hypotheses ................................ ................................ .................... 28 Chapter Organization ................................ ................................ .............................. 30 2 NATURAL AND CULTURAL ENVIRONMENT ................................ ....................... 33 Study Location ................................ ................................ ................................ ........ 33 Natural Enviro nment ................................ ................................ ............................... 33 Biogeographical Divides ................................ ................................ ................... 34 Paleogeography ................................ ................................ ............................... 35 Paleoenvironment ................................ ................................ ............................ 37 Paleofauna ................................ ................................ ................................ ....... 39 Settlement History ................................ ................................ ................................ .. 40 Colonization of the Pacific Region ................................ ................................ .... 40 Archaeology ................................ ................................ ............................... 40 Linguistics ................................ ................................ ................................ .. 41 Biology ................................ ................................ ................................ ....... 42 Colonization of the Mariana Islands ................................ ................................ 44 Linguistics ................................ ................................ ................................ .. 45 Archaeology ................................ ................................ ............................... 46 Genetics ................................ ................................ ................................ ..... 48 Bioarchaeology ................................ ................................ .......................... 49 Settlement Summary ................................ ................................ ........................ 51 Marianas Chronological Sequence ................................ ................................ ......... 51 Pre Latte Period ................................ ................................ ............................... 53 Diet ................................ ................................ ................................ ............ 54 The Early Unai phase (3500 3000 BP) ................................ ...................... 56

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7 The Middle Unai period (3000 2500 BP) ................................ .................... 57 The Late Unai period (2500 2400 BP) ................................ ....................... 58 The Transitional (Huyong) period (400 1000 CE) ................................ ...... 59 Latte Period (1000 1668 CE) ................................ ................................ ............ 59 Latte architecture ................................ ................................ ....................... 60 Cooking and food processing ................................ ................................ ..... 61 Diet ................................ ................................ ................................ ............ 62 Post Contact Era ................................ ................................ .............................. 65 The Spanish Colonial Period (1521 1898 CE) ................................ ........... 66 The First American Period (1898 1941 CE) ................................ ............... 68 The Japanese World War II Period (1941 1944 CE) ................................ .. 68 The American World War II Period (1944 1948 CE) ................................ .. 69 The Second American Period (1945 Present) ................................ ........... 69 3 MATERIALS ................................ ................................ ................................ ........... 77 Archaeological Sample ................................ ................................ ........................... 77 Taphonomic Bias ................................ ................................ ................................ .... 78 Sample Popul ation ................................ ................................ ................................ .. 80 Pre Latte Demographics ................................ ................................ ................... 80 Latte Demographics ................................ ................................ ......................... 80 4 DENTAL REDUCTION ................................ ................................ ........................... 85 Background ................................ ................................ ................................ ............. 85 Current Study ................................ ................................ ................................ .......... 86 Expected Results ................................ ................................ ................................ .... 88 Methods ................................ ................................ ................................ .................. 90 Results ................................ ................................ ................................ .................... 92 Pre Latte and Latte Differences ................................ ................................ ........ 94 Male and Female Differences ................................ ................................ ........... 95 Interaction between Time and Sex ................................ ................................ ... 95 Discussion ................................ ................................ ................................ .............. 96 Tooth Summaries and Rate of Change ................................ ............................ 96 Hypothesis Testing ................................ ................................ ........................... 98 Time Period Differences ................................ ................................ ............ 98 Comp arison of dental, craniofacial, and postcranial changes across time ................................ ................................ ................................ ......... 99 Carious lesions ................................ ................................ ........................ 101 Mechanism for Dental Reduction ................................ ................................ .......... 102 Probable Mutation Effect ................................ ................................ ................ 102 In creasing Population Density Effect ................................ .............................. 103 Selective Compromise Effect ................................ ................................ ......... 104 Masticatory Functional Hypothesis ................................ ................................ 105 Conclusions ................................ ................................ ................................ .......... 105 5 DEVELOPMENTAL INSTABILITY: ENAMEL HYPOPLASIAS ............................. 110

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8 Enamel Hypoplasias ................................ ................................ ............................. 110 Health and Disease in the Shift to Agriculture ................................ ....................... 112 Current Study ................................ ................................ ................................ ........ 114 Materials and Methods ................................ ................................ .......................... 117 Results ................................ ................................ ................................ .................. 119 Age Differences ................................ ................................ .............................. 120 Sex Differences ................................ ................................ .............................. 120 Time Period Differences ................................ ................................ ................. 121 Discussion ................................ ................................ ................................ ............ 122 Age Differences ................................ ................................ .............................. 122 Sex Differences ................................ ................................ .............................. 123 Time Period Differences ................................ ................................ ................. 124 Broader Implications ................................ ................................ ............................. 127 Conclusions ................................ ................................ ................................ .......... 129 6 CARIOUS LESIONS ................................ ................................ ............................. 139 Formation of Carious Lesions ................................ ................................ ............... 140 Carious Lesions and Agricultural Intensification ................................ ................... 140 Culture History of Guam ................................ ................................ ....................... 142 Mate rials and Methods ................................ ................................ .......................... 145 Results ................................ ................................ ................................ .................. 146 Tooth Position ................................ ................................ ................................ 146 Age Differences ................................ ................................ .............................. 147 Sex Differences ................................ ................................ .............................. 147 Time Period Differences ................................ ................................ ................. 148 Discussion ................................ ................................ ................................ ............ 148 Tooth Position ................................ ................................ ................................ 148 Age Differences ................................ ................................ .............................. 150 Sex Differences ................................ ................................ .............................. 150 Time Period Differences ................................ ................................ ................. 151 The effects of diet ................................ ................................ .................... 152 The effects of betel nut chewing ................................ .............................. 156 Conclusions ................................ ................................ ................................ .......... 158 7 SUMMARY ................................ ................................ ................................ ........... 165 APPENDIX A DENTAL METRICS ................................ ................................ ............................... 170 B ANALYSES OF VARIANCE TESTS FOR DENTAL MEASUREMENTS .............. 206 LIST OF REFERENCES ................................ ................................ ............................. 249 BIOGRAPHICAL SKETCH ................................ ................................ .......................... 279

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9 LIST OF TABLES Table page 2 1 Archaeological and historical chronological sequences of the Marianas Islands ................................ ................................ ................................ ................ 71 2 2 Activities and artifact assemblage composition of latte sets ............................... 71 3 1 Radiocarbon dates from Naton Beach site, Tumon, Guam. ............................... 82 3 2 Pre Latte vs. Latte sample distribution ................................ ............................... 83 3 3 Okura dental sex distributions ................................ ................................ ............ 83 3 4 Okura dental age distributions ................................ ................................ ............ 83 3 5 Pre Latte dental sample by age and sex ................................ ............................ 83 3 6 Latte dental sample by age and sex ................................ ................................ ... 8 4 4 1 Tooth summary data ................................ ................................ ......................... 107 4 2 Tooth summaries of prehistoric Chamorro populations from Guam ................. 107 4 3 Tooth summaries of Pacific and circum Pacific samples ................................ .. 108 4 4 Mean cranial and mandibular measurements associated with masticatory apparatus ................................ ................................ ................................ ......... 109 5 1 Percentage of teeth with one or more linear enamel hypoplasia ...................... 132 5 2 Individual occurrence of Pre Latte and Latte linear enamel hypoplasias by age grouping ................................ ................................ ................................ ..... 132 5 3 Individual occurrence of linear enamel hypoplasias ................................ ......... 132 5 4 Tooth count of linear enamel hypoplasias ................................ ........................ 133 5 5 LEH frequencies in comparative populations ................................ .................... 134 5 6 Age differences of leh using a Pearson Chi Square Test ................................ 135 5 7 Sex differences of LEH using a Pearson Chi Square Test ............................... 135 5 8 Pre Latte and Latte differences in LEH using a Pearson Chi Square Test ....... 135 6 1 Individual occurrence carious lesion frequencies ................................ ............. 160

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10 6 2 Tooth count of carious lesion frequencies ................................ ........................ 160 6 3 Individual occurrence carious lesions by age ................................ ................... 160 6 4 Frequency of carious lesions by tooth class ................................ ..................... 161 6 5 Frequency of carious lesions by tooth position ................................ ................. 161 6 6 Pearson Chi Square Test on carious lesion expression ................................ ... 162 A 1 Descriptive statistics of dental measurements ................................ .................. 170 A 2 Descriptive statistics of dental measurements by time period .......................... 172 A 3 Descriptive statistics of dental measurements by time period and sex. ............ 176 A 4 Descriptive statistics of cross sectional area by time period ............................. 189 A 5 Descriptiv e statistics of cross sectional area by time period and sex ............... 190 A 6 Group comparisons of dental measurements ................................ ................... 193 A 7 Kolmogorov Smirnov a Test for normality ................................ .......................... 200 A 8 Levene's Test of Homogeneity of Variance based on the mean ....................... 204 B 1 Two Way Fa ctorial ANOVA for LMax I1 MD ................................ ..................... 207 B 2 Two Way Factorial ANOVA for LMax I1 BL ................................ ...................... 207 B 3 Two Way Factorial ANOVA for LMax I2 MD ................................ ..................... 208 B 4 Two Way Factorial ANOVA for LMax I2 BL ................................ ...................... 209 B 5 Two Way Factorial ANOVA for LMax C MD ................................ ..................... 210 B 6 Two Way Factorial ANOVA for LMax C BL ................................ ...................... 211 B 7 Two Way Factorial ANOVA for LMax P3 MD ................................ ................... 212 B 8 Two Way Factorial ANOVA for LMax P3 BL ................................ ..................... 213 B 9 Two Way Factorial ANOVA for LMax P4 MD ................................ ................... 214 B 10 Two Way Factorial ANOVA for LMax P4 BL ................................ ..................... 215 B 11 Two Way Factorial ANOVA for LMax M1 MD ................................ ................... 216 B 12 Two Way Factorial ANOVA for LMax M1 BL ................................ .................... 217

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11 B 13 Two Way Factorial ANOVA for LMax M2 MD ................................ ................... 217 B 14 Two Way Factorial ANOVA for LMax M2 BL ................................ .................... 218 B 15 Two Way Factorial ANOVA for LMax M3 MD ................................ ................... 218 B 16 Two Way Factorial ANOVA for LMax M3 BL ................................ .................... 219 B 17 Two Way Factorial ANOVA for RMax I1 MD ................................ .................... 219 B 18 Two Way Factorial ANOVA for RMax I1 BL ................................ ..................... 220 B 19 Two Way Factorial ANOVA for R Max I2 MD ................................ .................... 220 B 20 Two Way Factorial ANOVA for RMax I2 BL ................................ ..................... 221 B 21 Two Way Factorial ANOVA for RMax C MD ................................ ..................... 222 B 22 Two Way Factorial ANOVA for RMax C BL ................................ ...................... 223 B 23 Two Way Factorial ANOVA for RMax P3 MD ................................ ................... 223 B 24 Two Way Factorial ANOVA for RMax P3 BL ................................ .................... 224 B 25 Two Way Factorial ANOVA for RMax P4 MD ................................ ................... 224 B 26 Two Way Factorial ANOVA for RMax P4 BL ................................ .................... 225 B 27 Two Way Factorial ANOVA for RMax M1 MD ................................ .................. 225 B 28 Two Way Factorial ANOVA for RMax M1 BL ................................ ................... 226 B 29 Two Way Factorial ANOVA for RMax M2 MD ................................ .................. 226 B 30 Two Way Factorial ANOVA for R Max M2 BL ................................ ................... 227 B 31 Two Way Factorial ANOVA for RMax M3 MD ................................ .................. 227 B 32 Two Way Factorial ANOVA for RMax M3 BL ................................ ................... 228 B 33 Two Way Factorial ANOVA for LMand I1 MD ................................ ................... 228 B 34 Two Way Factorial ANOVA for LMand I1 BL ................................ .................... 229 B 35 Two Way Factorial ANOVA for LMand I2 MD ................................ ................... 229 B 36 Two Way Factorial ANOVA for LMand I2 BL ................................ .................... 230 B 37 Two Way Factorial ANOVA for LMand C MD ................................ ................... 230

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12 B 38 Two Way Factorial ANOVA for LMand C BL ................................ .................... 231 B 39 Two Way Factorial ANOVA for LMand P3 MD ................................ ................. 232 B 40 Two Way Factorial ANOVA for LMand P3 BL ................................ .................. 232 B 42 Two Way Factorial ANOVA f or LMand P4 BL ................................ .................. 234 B 43 Two Way Factorial ANOVA for LMand M1 MD ................................ ................. 235 B 44 Two Way Factorial ANOVA for LMand M1 BL ................................ .................. 235 B 45 Two Way Factorial ANOVA for LMand M2 MD ................................ ................. 236 B 46 Two Way Factorial ANOVA for LMand M2 BL. ................................ ................. 236 B 47 Two Way Factorial ANOVA for LMand M3 MD ................................ ................. 237 B 48 Two Way Factorial ANOVA for LMand M3 BL ................................ .................. 238 B 49 Two Way Factorial ANOVA for RMand I1 MD ................................ .................. 238 B 50 Two Way Factorial ANOVA for RMand I1 BL ................................ ................... 239 B 51 Two Way Factorial ANOVA for RMand I2 MD ................................ .................. 239 B 52 Two Way Factorial ANOVA for RMand I2 BL ................................ ................... 240 B 53 Two Way Factorial ANOVA f or RMand C MD ................................ .................. 240 B 54 Two Way Factorial ANOVA for RMand C BL ................................ .................... 241 B 55 Two Way Factorial ANOVA for RMand P3 MD ................................ ................. 242 B 56 Two Way Factorial ANOVA for RMand P3 BL ................................ .................. 242 B 57 Two Way Factorial ANOVA for RMand P4 MD ................................ ................. 243 B 58 Two Way Factorial ANOVA for RMand P4 BL ................................ .................. 244 B 59 Two Way Factorial ANOVA for RMand M1 MD ................................ ................ 245 B 60 Two Way Factorial ANOVA for RMand M1 BL ................................ ................. 246 B 61 Two Way Factorial ANOVA for RMand M2 MD ................................ ................ 246 B 62 Two Way Factorial ANOVA for RMand M2 BL ................................ ................. 247 B 63 Two Way Factorial ANOVA for RMand M3 MD ................................ ................ 248

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13 B 64 Two Way Factorial ANOVA f or RMand M3 BL ................................ ................. 248

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14 LIST OF FIGURES Figure page 1 1 Model for interpreting stress in skeletal populations ................................ ........... 31 1 2 The island of Guam and Naton Beach Site location ................................ ........... 32 2 1 The Mariana Island Chain ................................ ................................ .................. 72 2 2 Paleoshoreline notches along the western coast of Guam. ................................ 73 2 3 ite. .................. 73 2 4 Examples of Pre Latte Period pottery ................................ ................................ 74 2 5 Island of Guam depicting Latte Set density and distribution, follo wing s survey ................................ ................................ ..... 75 2 6 Examples of Latte Period pottery ................................ ................................ ........ 76 5 1 Sing le linear enamel hyoplasia in the mandibular lateral incisor, canine, and third premolar. ................................ ................................ ................................ .. 136 5 2 Multiple linear enamel hypoplasias in a single tooth. ................................ ........ 136 5 3 Labial abrasion in a Pre Latte Period individual. ................................ ............... 137 5 4 Betel nut staining in a Latte Period individual. ................................ .................. 137 5 5 Dental incising in a Latte Period Individual. ................................ ...................... 138 5 6 Frequency of linear enamel hypoplasias by tooth type ................................ .... 138 6 1 Pre Latte carious lesions in the anterior dentition. ................................ ............ 164 6 2 Pre Latte dental crowding. ................................ ................................ ................ 164 6 3 Betel nut with piper leaf and slacked lime. ................................ ....................... 164

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15 LIST OF ABBREVIATION S ANOVA Analysis of Variance AVG Average BL Buccolingual diameter of the tooth BP Before Present CE Common Era CX Cross sectional area of the tooth g/m 3 Grams per cubic meter IPDE Increasing Population Density Effect km Kilometers LEH Linear enamel hypoplasia MD Mesiodistal diameter of the tooth MF Masticatory Functional Hypothesis mm Millimeters PME Probable Mutation Effect SCE Selective Compromise Effect TS Tooth summary LMax I1 Left maxillary central incisor LMax I2 Left maxillary lateral incisor LMax C Left maxillary canine LMax P3 Left maxillary third premolar LMax P4 Left maxillary fourth premolar LMax M1 Left maxillary first molar LMax M2 Left maxillar y second molar LMax M3 Left maxillary third molar

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16 RMax I1 Right maxillary central incisor RMax I2 Right maxillary lateral incisor RMax C Right maxillary canine RMax P3 Right maxillary third premolar RMax P4 Right maxillary fourth premolar RMax M1 Ri ght maxillary first molar RMax M2 Right maxillary second molar RMax M3 Right maxillary third molar LMand I1 Left mandibular central incisor LMand I2 Left mandibular lateral incisor LMand C Left mandibular canine LMand P3 Left mandibular third premolar LMand P4 Left mandibular fourth premolar LMand M1 Left mandibular first molar LMand M2 Left mandibular second molar LMand M3 Left mandibular third molar RMand I1 Right mandibular central incisor RMand I2 Right mandibular lateral incisor RMand C Right mandibular canine RMand P3 Right mandibular third premolar RMand P4 Right mandibular fourth premolar RMand M1 Right mandibular first molar RMand M2 Right mandibular second molar RMand M3 Right mandibular third molar

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17 Abstract of Dissertation Present ed to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy A DIACHRONIC ASSESSMENT OF HEALTH AND DISEASE FROM THE ADULT DENTITION OF THE NATON BEACH BURIAL COMPLEX IN TUMON BAY, GUAM By Nicolette M.Parr August 2012 Chair: Michael Warren Major: Anthropology The current study is an investigation of the prehistoric Chamorro in Guam to assess health and disease patterns over time. The transition from the Pre Latte to Latte periods displays a shift from horticultural to early agricultural practices; accompanying changes include increased population size and technologically advanced food processing and preparation techniques. T hese c hanges occur concomitantly with large scale environmental and climatic fluctuations. It is predicted that the cultural and environmental shifts will be accompanied by biological ones, due to increased stress levels associated with malnutrition, limited ac cess to resources, and increased prevalence of disease. Analyses of odontometrics, linear enamel hypoplasias, and carious lesions were performed and analyzed in concert with skeletal data collected by other researchers to construct a health profile of the prehistoric populations in Guam. Expected results include dental reduction over time coupled with an increase in linear enamel hypoplasias and carious lesions. The dentition display an 8% decrease in size from the Pre Latte to Latte periods. Increased r eliance on starchy crops would have le d to selection for smaller dentition to

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18 minimize carious lesions. Additionally, sophistication in food processing techniques decreases the force necessary to break down tough food, leading to reduced functional demand s of the masticatory apparatus. Thus, this finding is best explained by a combination of the Selective Compromise Effect and Masticatory Functional Hypothesis. Significant differences in linear enamel hypoplasia expression are noted with an increase over time. While not significant, the data suggests that there may have been differential access to resources as a result of gender roles associated with food procurement, where the females in the Latte period were much more highly susceptible to physiologica l stress than the males. Carious lesions are significantly different over time; however, these findings do not follow the predicted pattern. Caries frequency in the Latte period decrease over time likely due to the cultural practice of betel nut chewing, w hich has cariostatic properties This study expands on the current knowledge of prehistoric health in Guam by demonstrating an overall decrease in health over time as a result of climatic instability and subsequent dietary transitions.

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19 CHAPTER 1 INTRODU CTION In these Proes 1 several hundred Leagues, the Sun serving them for a compass by day and longer at a loss to know how th e Islands lying those Seas came to be Captain James Cook, The Journals of Captain James Cook on His Voyages of Discovery (in Beaglehole, 1955) Since the dawn of European exploration into the Pacific, Captain James Cook, and others, pondered how the earliest peoples came to inhabit the most remote of the Pacific islands. This journey across thousands of kilometers of ocean to reach small landmasses represents t he last migration into unchart ed territory (Spate, 1979) and is a 67). T he relative isolation of the Pacific Islands makes their populations ideal subjects for evolutionary studies (Howells, 1973). Small populations, as are often found on islands are more sensitive to random genetic changes in comparison to large populations (Turner, 1987). Thus, populations on small, isolated island s are more likely to undergo more noticeable phenotypic modification due to environmental changes and subsequent selective p ressures (Houghton, 1991 a ). Additionally, the diversity of investigating on a broad comparative scale the effect of environment on phenotypic : 154). 1 Cook is referring to proas maritime sailing vessels used in the Pacific.

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20 The Pacific is the focus of much bioarchaeological research; however, Guam and its people, the Chamorro, are underrepresented in the scientific literature in comparison with other island groups. As Howells noted in 1973 many books have been w ritten about where the Polynesians c ame from but nobody cares a stra w where the Guamanians came from. And yet it is probable that they can tell at least as much about the peopling of the Pacific as can the Polynesians (p. 248). Almost 40 years later, th e paucity of skeletal research in Guam still remains, regardless of evidence that places the earliest colonization of R emote Oceania in the Marianas Islands ( Carson 2008; Clarke et al., 2010; Carson, 2010 ) M ost research in Guam is often contracted out to specialists by cultural resource management firms (Howells, 1973; 1989; Pietrusewsky, 1990; Houghton, 1996) and many studies combine collections from different time periods while overlooking important differences that may be gleaned from a more thoroug h diachronic study of its populations. Exposure of the remains to a tropical environment over a long period of time leads to poor preservation and high fragmentation of the skeletal remains, thus explaining the dearth of skeletal research in Micronesia (H anson and Butler, 1997). Burial reports, often difficult to procure, contain the majority of the details, discussions, conclusions, and raw data regarding these remains ( e.g. Graves and Moore, 1985; Hanson, 1991; Heathcote, 1991; Douglas and Ikehara, 199 2; Heathcote, 1994; Pietrusewsky and Ikehara Quebral, 1994; Ikehara Quebral, 1998; Pietrusewsky, 1988; Ikehara Quebral, 1999; Heathcote, 2006). Few published articles on bioarchaeological research in the Marianas are available, however and most are restri cted to a single published volume (vol. 104, 1997)

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21 in The American Journal of Physical Anthropology (Ambrose et al., 1997; Arriaza, 1997; Douglas et al., 1997; Hanihara, 1997; Hanson and Butler, 1997; Hanson and Pietrusewsky, 1997; Ikehara Quebral and Doug las, 1997; Ishida and Dodo, 1997; Pietrusewsky et al., 1997; Stodder, 1997); while others are scattered throughout additional peer reviewed journals ( e.g. Leigh, 1930; Rothschild and Heathcote, 1993; Rothschild and Heathcote, 1995; Rothschild and Rothschil d, 1995) or in small Pacific based publications ( e.g. Underwood, 1973, 1976; Houghton, 1991 b ; Heathcote et al., 1996; Suzuki, 1986; Pietrusewsky, 1990; Turner, 1990; DeFant, 2008). Furthermore, the majority of these studies focus primarily on craniometric data for population history and health and disease of the Chamorro population. Few studies concentrate on the dentition (Leigh, 1930; Brace et al., 1981; Brace et al., 1990; Hanihara,1990; Turner, 1990; Turner, 1992, Heathcote, 1994), even though the tee th are often the most highly preserved portion of human skeletal remains, in pre historic and archaeological contexts, making them an excellent repository of biological information (Brace et al., 1987; Kieser, 1990; Hillson, 1996). Previous dental studies address biological distance between geographical groups of the Pacific region (Brace et al., 1981; Brace et al., 1990; Hanihara and Ishida, 2005). However, these studies combine disparate time periods in their population analyses, thus obscuring importan t differences that may be inferred through a more fine grained analysis of temporal variation. Theoretical Framework This research adopts a biocultural approach, which combines biological, cultural, and archaeological data to analyze adaptations associate d with subsistence patterns and health status in prehistoric populations (Armegalos and Van Gerven, 2003)

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22 Central to the biocultural analysis of human health and disease is Goodman and ehistoric skele tal assemblages, which will be discussed in further detail below. Biocultural Approach to Bioarchaeology Biocultural studies focus on the population, rather than individual typological traits, as the unit of evolutionary change within the environment (Wellin, 1978). This method models to understand interrelatedness of biological, cultural, and environmental variables, as well as behavioral patterning occurring within a population (Blakely, 1977). Thus, evaluating information gathered from biological, archaeological, and cultural contexts could provide much needed insights to dynamic evolut ionary processes and adaptations in prehistory. In the 1970 s the term bioarchaeology was coined independently within the United Kingdom (U.K.) and the United States (U.S.), each with different definitions. Whereas in the U.K. the term was originally assoc iated with the study of faunal remains (Clark, 1972), bioarchaeology as defined in the U.S., was coined by Buikstra in an edited volume by Blakely (1977a) Biocultural Adaptation in Prehistoric America. Bioarchaeology as it is known in the U.S., and now i n other parts of the world, stems approach to population based, ecological research (Buikstra, 1997; Buikstra and Beck, 2006). Bioarchaeology can be defined as a context ual study of human skeletal remains from archaeological contexts, which integrates biology, culture, and enviro nment to better understand health, disease, and demography in prehistoric human populations

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23 (Buikstra; 1977; Larsen, 1997; Buikstra and Beck, 200 6). Unlike osteological investigations where skeletal data were largely typological and descriptive, the bioarchaeological approach emphasizes problem oriented research to reconstruct prehistoric lifeways through the integration of biological and archaeol ogical data (Buikstra, 1977). Bioarchaeologists often take a biocultural approach to understanding evolutionary dynamics allowing for a holistic analysis of prehistoric populations and their interactions with the surrounding environment through a synthesis of biology and culture (Blakely, nor biological adaptation, but through biocultural Recent studies emphasize a biocultural approach by comb ining archaeological and osteological research to answer significant questions about adaptation and the force s that drive biological change such as weaning and dietary shifts in Guatemala (Wright and Schwarcz, 1999); demographic collapse in Spanish Florida (Griffin and colleagues, 2001; Stojanowski, 2003; 2005) ; climatic variability in Ja pan ( Temple, 2007; Temple and Larsen, 2007; Oxenham and Matsumura, 2008; Temple, 2010 ) ; ecological and demographic pressure in the Nile Va lley (Starling and Stock, 2007) and mobility in Northern Africa (Stojanowski and Knudson, 2011) Stress Models Early approaches to the study of stress focused on how external and environmental parameters place strain on a given organism (Goodman et al., 1988). In the middle of the twent ieth century, Hans Selye (1936 1956, 1973) introduced a new perspective on the study of stress where the focus shifted from strain as a result of environmental stressors to looking at physiological change (i.e. health and

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24 where alterations of the environment alters the normal and steady state of an organism. Central to the Selyean concept o f stress is the general adaptation syndrome, a defense mechanism in which homeostatic mechanisms are activated to alleviate a long and co ntinued stress event (Selye, 195 6). Classic l aboratory studies showed that increased exposure to stressors (i.e. nois e, cold, and heat) lead to developmental disturbances in rodents (Siegel and Smookler 1973; Siegel and Doyle 1975a,b; Siegel et al. 1977; Doyle et a l. 1977; S ciulli et al. 1979). Recently rodent studies have demonstrated that stress stimuli impairs memory (Luine et al., 1994; Conrad et al., 1996); alters cardiovascular activity (Rudyk et al., 2001), and a ffects sexual maturity and weight ( Rodriguez et al. 2007) In terms of human skeletal and dental remains, nonspecific indicators of stress represent an adaptive response to a stressor, which occurred during the development of an individual (Roberts and Manchester, 2005). Paleopathology at the Origins of Agriculture the authors present a model for stress app licable to skeletal populations in which health is the fundamental variable in examining the adaptive processes of an individual or population (Larsen, 1997; Goodman and Martin, 2002). Subsequently, the model has been reworked by Goodman and colleagues to include feedback systems and indicators of stress in the skeleton and provides a systematic framework to analyze the effects of a physiological disruption (Goodman 1991; Goodman and Armelagos, 1989; Goodman et al. 1988; Goodman and Martin, 2002).

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25 The mod el presented in Figure 1 1 begins with the environment, which provides certain necessary resources (e.g. food, shelter, and water) as well as stressors (e.g. extreme temperatures, parasites, predators) that may affect the health of a population (Goodman and Armelagos, 1989; Goodman and Martin, 2002). Technological advances in culture, such as clothing and shelter, may often provide a buffer to stressors. In cases where the cultural system does not adequately buffer the environmental constraints, the str essors will reach the individual or population. impact will be noted in individuals who can combat any given stressor, however, some individuals may be unable to resist the stressor, due to genetic susceptibility, disease, malnutrition, age, sex, and/or resiliency (Goodman et al., 1988; Goodman and Martin, 2002). The severity of and duration of the stress response may be viewed as a function of the degree of cultural and environmental constraints and stressors, balanced against the adequacy of the cultural buffering system and individual resistance resources (Goodman and Martin, 2002: 18). If an individual fails to fight a stressor, a physiological disruption or biological stress response may occur, resulting in permanent and visible changes in the body (Goodman et al., 1988; Goodman and Armelagos, 1989; Larsen, 1 997; Goodman and Martin, 2002). S oft tissue responds more quickly to stress and disease than the skeleton The refore a stress event must be severe or endure for a prolonged period before the bone or teeth are affected (Goodman et al., 1988). Interpreting of skeletal markers of stress is difficult and while some diseases, such as tuberculosis, syphilis, and lepro sy, leave diagnostic lesions on the skeleton, many other pathogens elicit the same response fo r a number of given stressors

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26 (Goodman et al., 1998). For example, periostitis is one of the m ost common diseases indicative of trauma or infection ; however, periosteal bone formation occurs in a number of other infectious diseases ( Ortner, 2003 ). As such, identification of an infection in the skeleton may seem simple; however, diagnosis of its etiology proves quite difficult Additionally, some diseases, suc h as influenza and other viruses that may result in decreased health and in some cases death leave no evidence in the skeleton, which confound s the interpretation of health and disease from skeletal remains (Goodman et al., 1998) In what is now known as the Osteological Paradox, Wood and colleagues (1992) criticized the conclusions regarding population health in the transition to agriculture. The authors identified key conceptual problems that compli cate interpretations of health from skeletal remains. Most pertinent to the current study is the concept of hidden heterogeneity which refers to the amount of frailty of any given individual in essence, how prone an individual is to disease and death. Individuals who are more are most frail may succumb quickly to external stressors, leaving no markers of bony response of the disease pr ocess on the skeleton, while others who are exposed to moderate stressors, may survive through the stress event and thus elicit skeletal markers of stress. Some studies, however, have shown correlations between disadvantaged populations and disease. For e xample, studies in living human and non human primate populations have shown that individuals with a higher prevalence of linear enamel hypoplasias are not at a greater advantage than those without (Zhou and Corruccini, 1998; Guitelli Steinberg and Benderl ioglu 2006 ). Nonetheless, since its publications

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27 researchers have suggested several methods to correct for issues related to the Osteological Paradox. Goodman (1993) suggests that an analysis of multiple stressors, instead of an individual trait, would reduce the likelihood of misinterpreting health status. Ideal stress indicators are those associated for use with the health index as proposed by Steckel and Rose (2002): stature, hypoplasias, anemia, dental health, skeletal infections, degenerative join t disease, and trauma (Goodman and Martin 2002). Furthermore, an interdisciplinary approach to health status may prove helpful in an interpretation of skeletal lesions in archaeological populations. Larsen (1997: 337) states that in order to get a clear evaluation of population health, biological indicators of dis other lines of evidence, including subsistence and settlement, environmental context, cultural contex t, Thus, a n analysis of the cultu ral and environmental factors in association with biological indicators of stress will provide biocultural understanding of adaptation in archaeological skeletal samples. Purpose and Research Objectives The current study investigates evolutionary dynamics of the prehistoric Chamorro across time to see how they relate to biocultural and environmental changes in prehistoric society. Between the Pre Latte and Latte time periods in Guam, there are changes in population size and subsistence strategies ( Hunter A nderson and Butler, 1991 ). Likewise, they changed many of their food procurement and preparation strategies ( Amesbury, 1999; Moore, 2005 ; Amesbury, 2007 ) These transitions occur concomitantly with large scale environmental and climatic fluctuations such as sea level decline and increased storminess, aridity, and drought ( Hunter Anderson and Butler 1991); Nunn, 2007, Hunter Anderson, 2010). It is predicted that these cultural and

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28 environmental modifications will be accompanied by biological ones. For e xample, increase in stress levels associated with malnutrition, limited access to resources, and increased prevalence of disease, may be evidenced in the dentition as dental reduction, linear enamel h y poplasias, and carious lesions. In this study, I adopt a diachronic approach to assess change in the dentition as it correlates to environmental and cultural changes over time. A diachronic analysis of the dentition allows for an investiga tion into biological processes that can lead to biological change in hu man populations. An analysis such as this may uncover very small changes that occur between the two time periods that are often lost in broader studies that do not take temporal differences into account. Bellwood (1989: 4) emphasizes the need to evaluate processes that lead to cultural and linguistic diversification such as These te mporal changes in Guam represent the type of transitions that Bellwood feels are needed to analyze biological processes. Objectives and Hypotheses The current study focuses on the Naton Beach mortuary sample, which was excavated in response to cultural res ource management litigation. 2 This sample includes both the Pre Latte (n = 103) and Latte p eriods (n = 112) and is located on the west coast of Guam, in the Western Pacific, in Northern Tumon Bay at the Na ton Beach location (Figures 1 2 ). Four 14 C dates were obtained from conus shell bead necklaces associated with the earliest burials and range from 2 790 to 2330 BP (DeFant, 2008) 2 The Naton Beach site is often referred to colloquially as the Okura site in response to its location at the previous Okura Hotel, which was subsequently renovated to become the Guam Aurora Villas & Spa.

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29 This sample represents the largest Pre Latte skeletal assemblage to be excavated on Guam and one of the oldest and la rgest in the remote Western Pacific. Dates from the Chelechol ra Orrak cemetery in Palau (also in the Western Pacific) range from 3 000 BP to 200 CE ( Fitzpatrick, 2003; Fitzpatrick and Nelson, 2008 ) placing it either slightly earlier or contemporary to the Naton Beach site ; however, the sample size is limited to 26 individuals (Nelson and Fitzpatrick 2006 ; Fitzpatrick and Nelson, 2011). An overall review of the known settlement history of the Pacific and the M arianas Islands will be presented using four independent lines of evidence: linguistics, archaeology, genetics, and bioarchaeology. Evidence of cultural change between the Pre Latte and Latte time Periods will be evaluated in the archaeological record in an attempt to define triggers that may have led to change. These cultural changes will be analyzed in concert with the dental data to determine i f biological change has occurred following cultural shifts The hypotheses are as follows: Hn1: There is no s ignificant difference in the dental dimensions between the Pre Latte and Latte time periods. Hn2 : There is no significant difference in the frequency of linear enamel hypoplasias between the Pre Latte and Latte time periods. Hn3 : There is no significant difference in the frequency of carious lesions between the Pre Latte and Latte time periods. A focus on temporal differences in Guam will help clarify health and disease patterns in the prehistoric Chamorro, which until recently, w as only known for the late prehistoric peoples. Analysis of the dental data are combined with other indicators of disease from the postcranial skeleton, gleaned from published and unpublished reports, and evaluated within the broader frame of the changing ecosystems that coincides with the Pre Latte and Latte transition. Addit ional ly, subsistence adaptations are analyzed to

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30 determine the impact of agricultural intensification on the oral health of the C hamorro. If dental reduction, linear enamel hypoplas ias, and cariou s lesion differences are found, this s tudy will delineate causative factor s associated with these changes through further analysis of the archaeological record. In conclusion, an analysis of dental changes in the prehistoric Chamorro will e lucidate a shift in not only within island phenomena but also provide a framework for interpreting biocultural interactions of the Chamorro and the dynamic environment in which they lived. Chapter Organization This dissertation is organized into seven chap ters. The current chapter includes an introduction to the study, outlines the theoretical goals, and provides a brief overview of the site location. It also outlines the research problem and presents three key hypotheses to be tested using the recovered s keletal remains The second chapter details the natural and cultural hi story of Guam, beginning with the study location, paleogeography/environment/fauna, terminating with a review of the settlement history of the Pacific and the Marianas Islands In Cha pter 3, the sample materials are described with a focus on taphonomic biases and pop ulation demography. The fourth, fifth and six chapters report the background, data collection methods, including statistical procedures, results, and discussion of the re sear ch. In Chapter 4, odontometric analyses are performed to elucidate differences between the populations with a focus on mechanisms for dental reduction over time. Physiological stress, as evidenced by linear enamel hypoplasias, is analyzed in Chapter 5 and highlights trends associated with climatic variability between the populations The seventh and final chapter discusses the study as a whole and identifies future studies that can be conducted for a more holistic interpretation of the lifest yles of the prehistoric Chamorro

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31 Figure 1 1. Model for interpreting stress in skeletal populations. Redrawn after Goodman AH, Armelagos GJ. 1989. Infant and childhood morbidity and mortality risks in archaeological populations. World Arch 21 (page 226 Figure 1 )

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32 Figure 1 2. The island of Guam and Naton Beach Site location ( Map courtesy of Rad Smith/GANDA )

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33 CHAPTER 2 NATURAL AND CULTURAL ENVIRONMENT Study Location The Mariana Islands are an archipelago in Micronesia that forms a chain of 15 islands extending north south between 13 and 20 N latitude These islands are located approximately 2200 km southeast of Japan and approximately 6000 km west of Ha 1). The Marianas lie west of th e Marianas Trench Subduction Zone, where the Pacific and Philippine tectonic plates meet. The larger Pacific Plate is subsumed beneath the Philippine plate (Rainbird, 1994; Steadman, 2006). The five southern islands (Guam, Rota, Aguiguan, Tinian, and Sai pan) are the oldest and largest of the chain and are composed primarily of raised limestone, while the ten northern islands are volcanic in nature, eight of which are still active (Steadman, 2006). Guam is the southern most and largest of the islands for ming the Marianas chain and is approximately 50 km long and ranges between 6 and 19 km wide with an area of approximately 554 square km (Thompson, 1932; Karolle, 1993; Mylroie et al 2001; Gingerich, 2003). Geologically, Guam is divided by the Pago Adelu p fault line, which separates the northern low relief limestone plateau and southern volcanic cuesta with an uplifted limestone component on the eastern coast (Tabrosi et al., 2005). Natural Environment A basic understanding of the natural environment of G uam is necessary to elucidate the complexity of human cultural adaptations over time, particularly when ev aluating the archaeological record. This overview provides insight into environmental factors that may have led to biocultural changes in the Chamorr o.

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34 Biogeographical Divides In the 17 th classification system for the peoples and islands of the Pacific (Melanesia, Micronesia, and Polynesia) based on typological physical characte ristics of incredibly diverse and utility in terms of biological, cultural, and historical processes, save for Polynesia which has proven more homogeneous, both cultural ly and historically (Kirch, 2000; 2010); nonetheless, his classification system has become ingrained in Western thought and the terms continue to be utilized today (Thomas, 1989). As with most broad geographical groups throughout the world, the population s of the Pacific are not linguistically, culturally, and biologically homogeneous; instead variation across the Pacific displays clinal trends rather than sharp boundaries (Thomas 1989; Terrell 1986; Bellwood 1989). While not particularly useful from a b land masses in the Pacific and includes New Guinea, the Solomons, Vanuatu (previously known as New Hebrides), New Ca encompasses five main archipelagos: Palau (Belau), the Marianas, the Carolines, the Marshalls, and Kiribati (formerly the Gi only ~ 2,700 km 2 with Guam be ing the largest of the islands ( 582 km 2 ) (Kirch, 2010). Lastly, es the far eastern Pacific islands, including

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35 2010). Near Oceania to include New Guinea, the Bismarck Archipelago, and the Solomon Islands, and Remote Oceania which encompasses the entirety of the Pacific islands east of the Solomon Islands. This terminology is heavily rooted in archaeological findings that have demonstrated the antiquity of human settlement in Near Oceania extending into the Pleistocene, compared to Remote Oceania with earliest colonization ranging from 4,000 years ago, in the Western Pacific, to 1000 years ago in the Eastern Pacific (Green, 1991; Kirch, 2000; 2010). Thus, the suggested and continued classification scheme is in better standing with current linguistic, biological, and archaeological data. Paleogeography Guam has undergone dramatic changes in coastline due to changes in sea level and bioturbation from storm and wave acti vity (Bath, 1986; Kurashina and Clayshulte, 1983; Dickinson 2000; 2003; Carson, 2011). During the mid Holocene highstand (~6000 and 4 000 BP ) sea level elevations ranged between 1.6 m to 2.6 m above modern day sea levels in the tropical Pacific Islands (including Mariana and eastern Caroline Islands, Samoa, Fiji, Tonga, and Molokai) ( Dickinson 2001; 2003). Atolls, barrier reefs and most land masses in the Pacific Ocean were completely submerged except for the highest volcanic ridges ( Dickinson 2001; Amesbury and Hunter Anderson, 2008). After the mid Holocene sea level decrease began in 2200 BP in the northwest and southwest pacific; however in the eastern boundaries of the Pacific, sea

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36 levels did not begin to decline until ~ 0 CE and in some cases as late as 800 CE ( Dickinson 2003). In Guam, significant sea level changes can be measured directly from the wave cut notches of exposed limestone faces and emergent mid Holocene reef flats (see Figure 2 2 ), indicating that between 5400 and 3050 BP the sea level was over 1.8 m higher than in the present day (Easton et al., 1978; Dickinson 2000; 2003; Kayanne et al., 1993). This finding is in accordance with the mid Holocene highstand estimate (Dickenson 2003 ) Archaeological findings have placed arri val of the earliest colonizers ~ 3500 BP (see expanded settlement discussion below), which coincides with the mid Holocene highstand. Thus, settlers encountered high sea level paleoshorelines with fringing reefs, coastal flats, mangrove lined lagoons, stab le islets, and estuaries ( Dickinson 2003; Amesbury and Hunter Anderson, 2008). Approximately 300 years after settlement, in 3200 BP the post mid Holocene sea level decline changed the appearance of the coastline by expanding preferable habitation areas into wide sandy beaches along the coast and also allowed for more widespread dispersal of inhabitants throughout the island (Amesbury et al., 1996; Dickinson 2003). Archaeological studies in Tumon Bay have found evidence of coastal progradation associate d with sea level decline (Graves and Moore, 1985; Bath, 1986; Olmo, 1997; Magnuson et al., 2000). Additionally, this sea level decline necessitated cultural adaptations of the population to new environmental conditions (Carson, 2011). Understanding sea level and ecosystem changes in Guam helps clarify spatial differences in the locations of Pre Latte and Latte sites as well as variation of shellfish exploitation between the periods. Prograding coastlines and sea level drawdown explains why earlier Pre L atte sites are

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37 usually found more inland in comparison with the later Latte sites: the sandy beach areas, closest to the present shoreline, had not yet emerged in the Pre Latte times (Bath, 1986; Graves and Moore, 1985; Amesbury, 1999; Carson, 2011). Addi tionally, the shift from bivalve to gastropod consumption may be an indirect result and subsequent cultural adaptation due to altered ecosystems and the disruption of mangrove swamps, which are the preferred habitat of bivalves (Amesbury, 1999). Thus, dat a on sea level changes, coupled with archaeological data from prehistoric habitation sites, has shed light on the relationship between the dynamic alterations of paleocoastlines and associated cultural changes (Carson, 2011). Paleoenvironment The paleoenvironmental record has proven difficult to interpret however, several researchers provide valuable data to better understand prehistoric environmental conditions (Athens and Ward, 1993; 1995; 1999; 2004; Ward, 1994; 1995; Nunn, 1999; Nunn, 2007 ; Nunn et al., 2007). The mid Holocene highstand corresponds with the Holocene Climatic Optimum (HCO) in the Pacific region, which occurred between 6000 and 3000 BP, where higher sea levels, temperatures, and an abundance of organisms allowed for increase d diversity in habitat (Nunn, 1999). Since the end of the HCO attributed to climate change (Nunn, 2007: 2). Cool temperatures remained stable until ~ AD 750, during the Little Climatic Optimum (LCO also known as the Medieval Warm Period), when temperatures began to rise slowly, rainfall decreased, and sea levels once again rose, until approximately AD 1300 (Nunn, 2007; Nunn et al., 2007). The transitional phase f where rapid cooling temperatures, decline in sea levels, and increased storminess

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38 resulted in greater climatic variability, during this Little Ice Age (Bridgman, 1983; Nunn and Britton, 2001; Nunn, 2007; Nunn et al., 2007). The AD 1300 Event may well be the (Nunn, 2007: 1) and is associated with societal disruption, subsistence change, and move ment to inland habitatio n areas (Nunn, 2000; Nunn and Britton, 2001; Nunn, 2007) This pattern is observed throughout the Pacific Basin (Nunn, 2000; Nunn and Britton, 2001; Nunn, 2007). Further, the dramatic ecosystem fluctuations between the LCO and the Little Ice Age, associa ted with the AD 1300 Event, correspond with the shift from the Pre Latte to Latte time period and concomitant cultural modifications. Wetland sedimentary cores conducted at various locations throughout Guam have generated a continuous record of paleoclimat e and vegetation changes of the island landscape that predates human settlement (Ward, 1994; 1995; Athens and Ward, 1993; 1995; 1999; 2004). Pollen analysis from the IARII Laguas Core (Athens and Ward, 1999; 2004), which is the most detailed and complete paleoenvironmental record from Guam, indicates that this island was largely forested during the early Holocene. Between 4405 and 2956 cal. BP, forest and swamp/mangrove taxa begin to decline in conjunction with the arrival of the first human settlers. Li kewise, Lycopodium and Gleichenia ferns begin to appear, circa 3,900 cal. BP, indicating possible gardening and resource collecting. By 2900 cal. BP, ferns, grasses, and charcoal are in abundance, suggesting a shift to a savannah like habitat of open area s with grass cover, possibly augmented by both intentional and unintentional fires. By 2300 cal. BP, very little of the native forest persisted on the island; instead the majority of the island had been converted to the savanna landscape typical of modern day Guam.

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39 Paleofauna Research by Pregill and Steadman (2009) has provided valuable information on the prehistoric fossil record in Guam and documents the extensive faunal loss due to human colonization and the resulting habitat destruction, human predatio n, and introduction of exotic predatory animals. Terrestrial vertebrates were collected from two caves, Ritidian Cave 1 and Gotham Cave, both located on the northernmost area of Guam. Ten native reptiles were identified as well as two prehistorically int roduced species: the mangrove monitor lizard ( Varanus indicus ), introduced a round 2900 BP (Liston et al., 1996; Wiles et al., 1989), and blind snakes ( Ramphotyphlops sp.). Of the 10 native reptiles identified, the gekkonid lizard (Gekkonidae new sp.) is e xtinct. Seventeen species of bird were identified, of which five are currently extinct (Duck: Anas oustaletii ; Rail: Porzana undescribed sp.; Parrot: cf. Vini undescribed sp.; White eye: Zosteropidae new sp.), two are extirpated, and eight lost in histor ic times. Pregill and Steadman (2009) also found evidence for the introduction of the rat, Rattus rattus circa 800 to 1000 CE approximately 2000 years after human colonization and corresponding to the shift between the Pre Latte and Latte time periods. However, no chicken, dog, or pig rema ins were found in the prehisto ric skeletal assemblage in Guam, although they have been found in nearly all other Pacific Islands (Wickler, 2004), Far more lizard species have survived into modern times in comparison to birds. Extinction of native lizard species occurred after European contact due to habitat destruction, competition for resources, and predation by rats, the brown tree snake ( Boiga irregularis ), and other animals. There are currently only five ( of twent y four documented ) extant species of birds (herons two species, swifts, starlings, and crows)

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40 left on Guam. This extreme decline in birds is attributed to the introduction of the brown tree snake around World War II (Pregill and Steadman, 2009). Settle ment History In order to fully understand colonization in Guam and the Mariana islands, an overview of peopling of the Pacific region will first be presented, followed by a more detailed overview of settlement history of Guam. Colonization of t he Pacific Region The peopling of the Pacific is an area that has received a great deal of attention since the European discovery of the Pacific Islands. Over the past several decades, there has been much dispute as to the actual origins of the Pacific Islanders, i ncluding one hypothesis of settlement from South America (Heyerdahl, 1952), which has not been substantiated. Currently researchers are much more united in their ideas on the regions and dates of colonization and typically believe that the wide amount of population variation seen in the Pacific region is due primarily to regional processes of diversification (Bellwood, 1989). Several independent lines of data have recently come together for a more unified theory of the colonization of the Pacific. While t here are still some debatable issues, approaches from linguistic, archaeological, and biological perspectives have shed light on the peopling of the Pacific. A temporal framework for the colonization of the Pacific Islands is followed by a review of the a rchaeological, linguistic, and biological evidence. Archaeology The Pacific is characterized by two major colonization events (Thomas, 1999). The first settlement of the Pacific occurred in Near Oceania around 40,000 to 30,000 BP (Kirch, 1997). The secon d settlement event began around 3,500 BP with the rapid

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41 spread of the Lapita Cultural Complex and the Austronesian languages throughout Remote Oceania (Green, 1979; Kirch, 1997; Pawley, 1999; 2002). More specifically, colonization dates, based on archaeol ogical data for the geographical areas of the Pacific are as follows: Mariana Islands circa 3,500 BP; Eastern Melanesia (Santa Cruz region through Vanuatu and New Caledonia) circa 3,300 to 3,200 BP; eastern Micronesia circa 2,000 BP; Polynesia circa 2,000 BP; and the last colonized areas are New Zealand circa 1,000 BP, and Chatham Islands circa 500 BP (Sutton, 1980; Davidson, 1984; Bonhomme and Craib, 1987; Kirch and Hunt, 1988; Green 1991b; Craib, 1993; Butler 1994; Anderson, 1991; 1996). Linguistics The t hree major language groupings in the Pacific are Australian, Papuan, and Austronesian (Bellwood, 1989). Populations from the region of interest for the current study are all members of the Austronesian language family with approximately 1,200 modern lang uages, thus this family will be discussed in more detail (Kirch, 2010) Pawley and Green (1984) advocate a dialect chain model in contrast to many of the hierarchical family tree models often discussed in reference to Pacific language groupings (e.g. Terre ll, 1986; Terrell et al 1997). The Proto Austronesian homeland is believed to have originated in Taiwan approximately 6000 5000 BP before spreading across the Pacific region (Bellwood, 1991; Bellwood, 1997; Pawley, 1999). As outlined in Bellwood (2000 : 7 ) the spread of the Austronesian language is as follows: a subgroup of the Proto Austronesian subgroup developed into the Proto Malayo Polynesian family with colonization of the Philippines circa 4500 BP; rapid movement through island South East Asia and western Micronesia permitted the spread of the Malayo Polynesian subgroups between 4000 and 3000 BP; finally, the Proto Oceanic group developed in

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42 the Bismark Archipelago, followed by the spread of the Lapita cultural complex which occurred alongside the spread of the Oceanic languages throughout western Polynesia between 33000 and 2800 BP. Most languages of island Melanesia, Micronesia, and Polynesia (except for those of Western New Guinea, Palau, and Guam) stem from Oceanic subgroup of the Austronesian fam ily (Bellwood 1989; Pawley, 1972). The spread of Austronesian languages throughout the Pacific region has been shown to integrate well with settlement patterns as demonstrated archaeologically (Green, 1999; Spriggs, 1999; Bellwood, 2000) Biology Biolo gically, colonization studies can be subdivided into anthropometric genetic, and biodistance studies (Shapiro and Buck, 1936; Howells, 1970; Brace and Hinton, 1981; Serjeantson, 1985; 1989; Brace and Hunt, 1990; Brace et al., 1990; 1991; Pietrusewsky, 199 0a; 1990b; Houghton, 1991 b ; 1996; Hanihara, 1992; Turner, 1990a; 1990b; Scott and Turner, 1997; Hurles et al., 2002; Lum et al., 2002; Stephan and Chapman, 2003). For the purpose of this study, evidence will be paid primarily to the genetic, skeletal, and dental evidence of colonization of the Pacific. Genetic: Genetic analyses grouped the human leukocyte antigen (HLA) into two clusters: island Melanesian and Australian, and western Melanesian (Serjeantson, 1985; 1989). These studies demonstrated that p roto Polynesians likely traveled along the northern coast of New Guinea before arrival into Polynesia. Y chromosome studies have suggested Island Southeast Asia as the ancestral group to both Near and Remote Oceania (Hurles et al., 2002; Lum et al., 2002) Pietrusewsky (2006) outlines three conclusions from the genetic studies regarding peopling of the Pacific. First, common origins for Remote Oceania is likely from a region extending from Island Southeast Asia,

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43 the Bismarck Archipelago, and the northern New Guinea coast. Second, admixture between indigenous groups and Austronesian migrants likely occurred during eastward expansion across the Pacific. Lastly, differential settlement or gene flow patterns are postulated between males and females due to t he diversity between mtDNA and Y chromosome evidence. Cranial: The majority of multivariate cranial analyses have shown a close association between Polynesians and Micronesians and have suggested a homeland origin of Island Southeast Asia. Additionally, t he Polynesians and Micronesians are differentiated from the closely related Australians and Melanesians. Likewise, the Melanesian and Australian groups display more heterogeneity and variation than what is found within the more homogeneous Polynesian grou ps (Pietrusewsky, 1990a; 1990b; Hanihara, 1992; Stephan and Chapman, 2003). These findings are consistent with two colonization events of the Pacific (Pietrusewsky, 2006). Brace and coworkers (Brace et al., 1990; 1991) provided evidence of Japan as the h omeland for Pacific Island groups; however, no other studies have corroborated this finding. Dental: Odontometric studies have shown that Australians and Melanesians have some of the largest teeth in the world, followed by the smaller Micronesian and Polyn esian teeth (Brace and Hinton, 1981; Brace et al., 1990; 1991). Non metric dental variation displays two contrasting patterns between the Sinodont dentition of Polynesians and Micronesians and the Sundadont pattern found in Southeast Asia. While Australi ans and Melanesians do not fit either pattern, they are closest to the Southeast Asia Sundadont pattern (Turner, 1990a; 1990b; Scott and Turner, 1997). Likewise Hanihara (1992) found that Polynesians and western Micronesians had a

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44 closer affinity with Sou theast Asians than with Melanesians. These dental studies also support a two part settlement strategy for colonization of the Pacific. Archaeological, linguistic, and biological studies have already shown that there appears to be a two prong settlement pa ttern in the Pacific. However colonization of the Marianas is still unclear (Irwin 1992). The following section will outline the known evidence of settlement patterns in this region. Colonization of the Mariana Islands Lowered sea levels following the m id Holocene in the Pacific Ocean resulted in the appearance of attractive coastal environments, which subsequently allowed for rapid dispersal and init ial human colonization in the western parts of Remote Oceania (Dickinson, 2001; Carson, 2011). Most rese archers agree that the islands were colonized by horticulturalists, with sophisticated sea faring technology, from Island Southeast Asia (Bellwood, 1975; 1979; Hanson and Butler, 1997; Kirch, 2000). Intentional and planned migration is supported by analys is of pottery, language, and DNA (Hanson and Butler, 1997; Lum and Cann, 1998; 2000; Callaghan and Fitzpatrick, through Palau and Yap, however, this theory has not been supp orted by the linguistic and archaeological data (Anderson ; 2005; Clark, 2005). Most recently, Hung and colleagues (2010) suggest the Northern Philippines as the most likely point of origin for the earliest colonizers based on shared cultural similarities Archaeological and paleoenvironmental studies date initial settlement to approximately 3,500 BP (Spoehr, 1957; Craib, 1993; Butler, 1994; Carson, 2008, 2010; Clark et al., 2010); however one study, based on sedimentary cores, suggests an earlier date of 4,300 BP (Athens and Ward, 1999; Athens and Ward, 2004). If these theories are correct, Guam was

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45 colonized after an open sea voyage of 2,600 km, which is three times the previously reported longest ocean crossing in prehistory (900 km, between Vanuatu an d Fiji) by the Lapita people (Green, 1979; Keegan and Diamond, 1987). Additionally, this voyage would have occurred 500 years prior to the Lapita expansion (Craib, 1999; Spriggs, 1999), making Guam the earliest colonized island in Remote Oceania. A four prong approach, focusing on linguistics, archaeology, genetics, and skeletal data, is utilized to review colonization patterns of the Marianas. Linguistics A substantial amount of debate has occurred over the linguistic position of the indigenous Chamorro language (Blust, 2000; Reid 2002). Three viewpoints summarize the theories regarding the Chamorro language. The first suggests that Chamorro is closest to Philippine languages (Safford, 1909; Topping et al., 1975); the second theory supports tha t Chamorro is most closely related to Indonesian languages (Zobel, 2002), and the last viewpoint, and currently accepted interpretation, suggests that Chamorro is not closely related to any subgroupings of the larger Austronesian language family (Dyen, 196 5; Blust, 2000; 2009; Reid, 2002). The current assertion about the position of the Chamorro language places it in the Western Malayo Polynesian (WMP) group, which is spoken in the Marianas, Palau, Philipines, Malaysia, parts of Indonesia, coastal southern Vietnam, and Madagascar. The WMP group, along with the extra Formosan Austronesian languages, is part of the reconstructed Proto Malayo Polynesian language, which began to differentiate in northern Southeast Asia (Ross et al., 1998; Blust, 2000; 2009; Paw ley, 2002). The Chamorro language has a distinctly separate geographic source from the Lapita Proto Oceanic group, and historical linguistics suggest direct colonization from Island

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46 Southeast Asia (Kirch, 2010), and more specifically, from the central or northern Philippines (Blust, 2000; Reid, 2000). Archaeology The earliest occupation sites in the Marianas are found along the shorelines associated with dynamic and biologically diverse marine ecosystems (Hanson and Butler, 1997). Radiocarbon analysis on materials recovered from sites in the Marianas, including Achugao on Saipan (Butler 1994; 1995), Unai Chulu on Tinian (Craib, 1993; Craib 1998; Haun et al., 1999), and Unai Bapot on Saipan (Bonhomme and Craib, 1987; Carson, 2008; Carson and Welch, 2005; C lark et al., 2010) provide the earliest dates of settlement for the Mariana Islands and correspond to the Early Unai Phase. The Achugao site on Saipan provides the earliest radiocarbon dates for the Marianas from charcoal samples, 3470 120 BP and 3120 50 BP, and is associated with a large assemblage of Marianas Red slipped pottery, some with curvilinear or straight lines and lime filled decorations, termed Achugao Incised, that serves as the archetype for ceramics associated with early period settlem ent sites (Butler, 1994; 1995). Thirteen radiocarbon dates from charcoal samples at the Unai Chulu site on Tinian place occupation between 3400 and 2900 BP. Like the Achugao site, the Unai Chulu site is also associated with Marianas Red incised sherds, si milar to the Achugao Incised type (Craib, 1993; Craib 1998; Haun et al., 1999). The Unai Bapot site on Saipan is associated with a diverse assemblage of early decorative redware and blackware, with slipped surfaces, and has been established as one of the earliest sites in the Marianas, with 31 radiocarbon dates, from charcoal, shell, and wood, placing settlement between 3400 to 3200 cal. BP (Bonhomme and

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47 pushes o ccupation even further back to 3600 to 3 42 0 BP which pre dates Lapita expansion into Remote Oceania. Most of the earliest sites in Guam have questionable dates. Matapang Beach Park, in Tumon Bay, has been dated as early as 5260 100 BP, however, this sample is fr om a scattered charcoal deposit with no clear provenience. Dates with more secure stratigraphic contexts from the same site, were obtained from fire pit features and range from 3880 90 BP to 3170 70 BP (Bath, 1986). The Huchunao site is associated wi th Achugao Incised pottery has b een dated more securely to 3 690 to 2830 BP (Dilli et al., 1998). Most recently, the Ritidian site in northern Guam, dates from the beginning and end of the Early Unai Period. The early settlement dates to 3547 to 3323 BP a nd is associated with very thin, redware pottery. The slightly younger site, with thicker and coarser pottery, dates to 3 056 to 2842 BP (Carson, 2010). This site illustrates chronological change in pottery sequence over time with solid radiocarbon dates. slipped, circle and punctate Philippines (Hung et al., 2010: 913). The Neolithic and Iron age site of Nagsabaran, which is located within the Cagaya n Valley, dates to 4000 and 3300 BP ; thus encompassing the earliest settlement dates from the Mariana Islands. While decoration of pottery sherds is rare in Nagsabaran, the motifs are similar to those found in the Early U nai Phase; additionally the incised designs are often filled with lime (Hung et al., 2010). Thus, the archaeological evidence supports the linguistic findings of settlement from the Philippines.

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48 Genetics Lum and Cann (1998) analyzed five mtDNA region V length polymorphisms from over 800 individuals. This polymorphism is frequent in Island Southeast Asia, Micronesian, and Polynesia, suggesting an Island Southeast Asia origin for the latter two groups. Further, the data were compared with linguistic and geographic interactions spheres and revealed that there was extensive amount of gene flow in much of Micronesia, except for in the Marianas and in Palau (the two non Oceanic speaking populations), suggesting that these regions may have been isolated from o ther Micronesian groups. A subsequent study (Lum and Cann, 2000) sought to elucidate the relationship between Micronesians and Polynesians, who share a number of cultural (e.g. kava drinking only in Central and not Western Micronesia) and biological (cra niofacial measurements) traits (Pietrusewsky, 1990 a,b; Lum and Cann, 2000). The study found that 89% of Micronesians and Polynesians had shared mtDNA control region sequences. Additionally, Micronesians and Polynesians have been clu s tered into the same lineage group (Lineage group I.1), along with Indonesians. A nodal sequence (L22) of another lineage group (Lineage group I.2), is also shared by Micronesians and Polynesians, as well as populations from the Philippines and Borneo. However, regional diff erences in shared lineages have been found between Western and Central Eastern Micronesia. A chart plotting multidimensional chord distances between populations groups the central Micronesian islands (Nauru, Pohnpei, Kiribati, Kosrae, and Marshalls) more closely with Polynesian islands. The western Micronesian islands (Marianas, Palau) are further removed, with the exception of Yap. Looking specifically at the at the Marianas sequence it is obvious that it remains isolated and lies in between

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49 Near Oceani a and Remote Oceania groups, in concordance with its geographical position. Further, sequences of the Western Micronesia groups suggest that they were each settled independently, but directly from Island Southeast Asia. Post settlement gene flow between the Marianas and central eastern Micronesia is posited due to the low frequency and diversi ty of its group I sequence. The s e data are further evidence Yap. Bioarchaeol ogy Howells (1970; 1973; 1977; 1979) was influential in collecting and publishing anthropometric and cranial data on large numbers of living Pacific populations. Multivariate analysis of anthropometric data grouped Micronesians closely with Melanesians su ggesting gene flow and intermarriage between those populations (1970). Craniometric analyses, however, closely link Polynesians and Micronesians as distinct from Australians and Melanesians who share similar features in skull form (1977). esearch (1990a; 1990b; 1994; 1995; 1996; 2000; 2005; 2008) Micronesia populations in comparison to the rest of Remote Oceania, Near Oceania, Australia, Island Southeast Asi a, mainland Southeast Asia, east Asi a, and North Asia. M ultivariate craniometric analyses (27 measurements in 63 crania l series) on 2,805 male crania suggest that the crania from Guam (n = 46) are most closely related to the Polynesian series (Tonga Samoa Hawaii, Rapa Nui, Gambier Islands, Marquesas Islands, Society Islands, Tuamotu Archipelago, and Chatham Islands) (Pietrusewsky, 2005) A dendogram of Mahalanobis D 2 shows that the Polynesian and Guam grouping

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50 connects with a larger cluster formed by East Asian, North Asian, and Southeast Asian crania which are closest to cranial series from Island Southeast Asia. Thus, the Polynesian and Guam populations have likely originated from Island Southeast Asia. Ishida and Dodo (1997) analyze d 22 non metric cranial traits from the Marianas populations and compare d them with twelve other groups across the Pacific, mainland Southeast Asia, East Asia, mainland Asia, and North with Southeast Asian and East Asian populations. findings on Ch amorro settlement patterns is that their sample s are derived from the Hornbostel Thompson Collection 1 of skeletal material from the Mariana Islands originally housed in the Bishop Museum in Hawai i. These series of skeletons were collected by Hans Hornbos tel and J C Thompson in the 1920 s from various locations across Guam, Saipan, and Tinian (Ishida and Dodo, 1997). Most of these skeletons date to the pre contact Latte period and were found in association with Latte sets; however, the remains are poorly provenienced and the result of selective recovery for the most well preserved elements (Hanson and Butler, 1977). Thus, these skeletal remains are not representative of a typical mortuary population. While their research represents the largest undertakin gs in understanding the Chamorro settlement patterns and relationships to other island groups, a similar study on a Pre Latte sample would likely provide complementary information regarding early Chamorro populations. 1 The Hornbostle collection has since been repatriated to Guam and is currently being curated in the

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51 Unfortunately, as discussed above, Pr e Latte samples are typically very small and extremely fragmentary, thus precluding this type of analysis. Settlement Summary The linguistic, archaeological, genetic, and skeletal findings are consistent in their findings of settlement of the Marianas from Island Southeast Asia. Most recently, archaeology and genetic advances have been able to pinpoint a more precise location for area of origin the Philippines. As Ra inbird (2003: 85) notes a voyage from the uld constitute the longest sea crossing undertaken by terms of population history and initial expansion into Remote Oceania, but also represents development of new technological advances in precise navigation and sea faring skills. Marianas Chronological Sequence Spoehr (1954, 1957) was the first to place colonization of the Mariana Island chain at 3500 BP, based on radiocarbon dates of 3,479 200 years BP from oyster shells at t he Chalan Piao site on the island of Saipan, located approximately 220 km north of Guam (Figure 2 recent age dating to 1,700 years later (Cloud et al 1956). Nonetheless, the majority of t he earliest radiocarbon dates across the Marianas cluster around 3500 to 3000 BP, of Micronesia and Remote Oceania (Reinman 1977; Kurashina and Clayshulte 1983; Butler 19 95; Carson 2008; Carson 2010; Clark et al. 2010; Hung et al. 2011). Athens and Ward (2004) push this chronology as far back as 4,300 BP, based on the

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52 appearance of charcoal particles in the Laguas paleocore, however, this date has not been supported by ar chaeological findings. The history of the Marianas can be divided into the pre contact and post contact eras, each of which may be further subdivided based on archaeological and historical periods. While various chronological sequences have been suggested based primarily on different temper types found in ceramics, the standard terminology is adapted from that of by Spoehr (1957) and Moore (1983). Table 2 1 provides an overview of the standardized chronological sequences used in the Mariana Islands. Sp oehr (1957) proposed the chronology of the Marianas dividing the prehistory into tw o periods: the Pre Latte (3500 BP 1000 CE) and Latte Periods ( 1000 1521 CE ). Named after megalithic architecture, the latte are composed of two parts: coral limestone p illars, the haligi which are topped by a hemispherical stone cap, the tasa (Figure 2 3 ) This monumental architecture is only associated with the Latte Period Grouped latte stones are known as latte sets and were arranged in parallel rows designed to support wooden structures and are thought to be the foundation for prehistoric houses and meeting halls (Thompson, 1932; Thompson, 1940; Morgan 1988) stinguished between ceramic types with the Pre Latte Marianas Redware and the Latte Period Marianas Plainware. is based on analyses of ceramic sequences from Tarague Beach in No rthwestern Guam. Her classification was later refined into the current standard which has replaced 1) The following

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53 overview of the chronological sequence is based on a combination of Spoehr (1957) Pre Latte Period The peopling of Micronesia was accomplished by people with sophisticated ocean faring technology who already had horticultural capabilities (Bellwood 1979) (see more detailed discussion below). The Pre Latte p eriod (3500 BP 1000 CE ) in the Marianas begins with initial human colonization around 3,500 years ago and concludes with the onset of latte construction in the first millennium CE Very few Pre Latte sites have been found in the Marianas as older sites are more prone to erosional and depositional disturbances due to frequent storms. Additionally, wave activity removes the original soil horizon and depos its materials further inland, causing major alterations to archaeological deposits. Thus, many Pre Latte sites have not been preserved intact and have often been reworked, mixed, moved, or eradicated by storms (Kurashina and Clayshulte, 1983; Hunter Ander son and Butler, 1991). The Pre Latte period is characterized by small populations with low population densities and semi permanent habitation sites situated near the coastal margins (Hunter Anderson and Butler, 1991). However, Moore and colleagues (1988) suggest that the Pre Latte were not sedentary and moved seasonally in accordance with resource procurement strategies. There has been little research on terrestrial animal exploitation of the Pre Latte period in Guam and most information comes from the n orthern Marianas Islands. At Unai Chulu on the island of Tinian, located approximately 160 km north of Guam, bones from the fruit bat and rails were found (Haun et al., 1999). However, additional analysis to interpret food processing, cooking, or breakag e patterns were not conducted

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54 (Amesbury and Hunter Anderson, 2003). On the islands of Tinian and Aguiguan, Steadman (1999) found burned animal bone indicative of human cookery dating to the Pre Latte period. In the Railhunter Rockshelter on Tinian, datin g to approximately 2400 to 2200 BP, Steadman (1999) found fish, lizard, snake, and several bird species restricted to a layer that is associated with human activity. Likewise, the Pisonia Rockshelter on Aguiguan, also contained burned remains of fish, liz ard, Rallidae sp. (rails), and Gallicolumba xanthonura (white throated ground dove) (Steadman, 1999). Diet The majority of the Pre Latte diet came from bivalves, shellfish, and reef fishing (Amesbury et al., 1991). Archaeological studies on Saipan have shown that the Pre Latte period has a much higher percentage of fishing gear and related fishing production debris per temporal unit in comparison to the Latte Period (44% and 14%, respectively, of total shell artifacts per unit) (Butler 1995). Studies ha ve shown that bivalves, specifically Andara antiquate (Blood clam), Gafrarium tumidum (Tumid venus clam), and Gafrarium pectinatum (Comb venus clam), were found in abundance in Pre Latte deposits in Ypao Beach on Guam, as well as on Chalan Piao in Saipan ( Leidemann, 1980; Amesbury et al., 1996). Likewise, Graves and Moore (1985) found that bivalves, specifically Arcidae (Ark clam) comprised between 61% and 71% of the bivalve assemblage in the Pre Latte units from Tumon Bay. In the Tarague cultural sequence Kurashina and Clayshulte (1993) found Tridacna cf gigas (Giant clam) in the oldest stratigraphic layer, with radiocarbon dates of 3435 70 BP, placing it in the Pre Latte period. This finding is rare, as T. gigas is not known to have existed in the Ma rianas during the Holocene. Thus, presence of T. gigas

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55 may represent goods brought by the early island colonizers or may be the result of trade or communication networks with other Pacific populations. Zooarchaeological assemblages have also shown that a significant proportion of consumed foods came from fish in the Pre Latte period. Archaeological excavations conducted in Pagat (Craib, 1986) on the east coast of Guam yielded higher concentrations of pelagic fish bone in the Pre Latte Layer (density of 37 8.37 grams per cubic meter) in comparison to the Latte layer (187.20 g/m 3 ). Leach and Davidson (2006a) report on approximately 20 pelagic fish species excavated from the Mangilao Golf Course site on the east coast of Guam, and determined that there are differences in their frequency over time. While the number of species exploited between Pre Latte and Latte periods increase, the percent of some species represented in the archaeological samples decrease throughout prehistory. For example, the most comm on fish in the archaeological assemblage, parrotfish (Scaridae), decreases in frequency from 60%, in the Early Pre Latte, to 27% in the Intermediate/Transitional Pre Latte, and then increase s to 43% in the Latte period. Emperorfish (Lethrinidae) and wrass es/tuskfish (Coridae/Labridae) decrease in frequency over time, while other pelagics, mahimahi (Coryphaenidae) and swordfish/marlin (Istiophoridae/Xiphiidae) increase in the Latte period. McGovern and Wilson (1996) similarly found elevated 15 N values i n a small Pre Latte sample from Saipan, suggesting increased exploitation of pelagic species, rather than dependence upon coastal and marine habitats. Evidence from Ylig Bay, on the east coast of Guam, showed no significant differences from frequencies of fish remains

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56 between the Transitional Pre Latte and Latte/Historic phases (Leach and Davidson, 2006b). The impact of coastal deposition due to eroding hillsides suggests that the landscape was extensively altered and likely utilized for horticultural pr actices around 2,400 BP (Athens and Ward, 2004). This evidence of landscape modification suggests that horticultural practices and foreign cultigens were likely brought with the early colonizing populations (Hunter Anderson, Butler, 2001). The appearance of Lycopodium and Gleichenia ferns early on in paleoenvironmental sediment cores suggests small scale gardening and resource collecting by initial settlers (Athens and Ward, 2004). Breadfruit ( Artocarpus ) and taro pollen ( Colocasia esculenta ) are also not ed early in the coring record, and around 1100 BP. Further evidence of taro comes from identified taro starch grains from Pre Latte pottery sherds, where Cordyline (Cabbage tree) fish and, shellfish, were also found, suggesting their importance and use a mong the foods cooked in the Pre Latte (Loy, 2001a,b; 2002; Loy and Crowther, 2002). The Pre Latte phase is further subdivided into four phases based on differences in pottery sequences from Tarague Beach on the west coast of Guam: the Early, Middle, and Late Unai, and the Transitional/Huyong (Moore, 2002). These phases will be discussed in terms of pottery characteristics and cultural materials associated with each phase. Known sites dated to these periods will also be addressed. The Early Unai p hase ( 3500 3000 BP ) The first 500 years following initial human settlement constitute the Early Unai Phase, which is characterized by non decorated Marianas Redware (Moore, 2002). These vessels have thin walls, restricted openings, everted rims, and slipped ext eriors

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57 (Moore, 2002). Other pottery types, such as Achugao, with curvilinear or straight lines and dentate infilling, and San Roque, characterized by curvilinear lines with stamped circles, have also been found in small frequencies in Saipan (Butler, 1995 ; Moore, 2002) (Figure 2 4). Coastal locations with access to marine resources were likely habitation areas for the earliest settlers (Hunter Anderson and Butler, 1991). Cultural materials associated with this phase include a high proportion of bivalve re mains, stone and shell tools, fishing equipment, and shell ornaments (Bath, 1986; Graves and Moore, 1995). Few sites of this earliest phase are known for Guam, but exceptions include Huchunao in Mangilao; Ypao and Matapang in Tumon Bay; and Ritidian in th e north (Leidemann, 1980; Bath, 1986; Dilli et al., 1998; Carson 2010). The Middle Unai p eriod (3 000 2500 BP ) The Middle Unai Phase spans the next 500 years following the Early Unai Phase. Archaeological assemblages are very similar to the Early Unai Phas e, with the exception of pottery. Marianas Redware usage is continued in this phase with everted or flared unthickened rims. Many of the vessels have calcareous or volcanic sand tempering. As seen in the Early Unai, many of the surfaces are plain or sli pped, however, new surface treatments also emerged, characterized by polishing or burnishing, in the Middle Unai (Moore, 2002). By the Middle Unai Period, the Achugao and San Roque designs were abandoned and replaced by Ipao Stamped, after Craib (1990), wi th bold lime filled designs with combinations of straight lines, circles, half circles and chevrons (Moore 2002: 8). Over 30 different types of band designs have been reported from across the Marianas islands.

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58 Ray (1981) reports another pottery type cha racteristic to the Middle Unai, know as Tarague Striated, which has scrape marks and impressions parallel to the rim seen on both the inside and outside of the vessel. Other observed surface treatments during this time period include fingernail impression s, dots, and ridges (Ray, 1981; De Roo and Goodfellow, 1998; Moore, 2002). Known sites dating to the Middle Unai increase in abundance in comparison to the Early period. These sites are primarily located along the coast and in caves and rockshelters along the shoreline. Some sites associated with the Middle Unai featuring Ipao Stamped pottery are located in Tumon Bay: Ypao (Leidemann, 1980; Olmo and Goodman, 1994), Kallingal Property (Moore et al., 2001), and at the site of the current study, Naton Beach (Defant, 2008). The Late Unai period ( 2 500 2400 BP ) The Late Unai phase begins around 2,500 years ago and lasts 100 years. This period is characterized by a decrease in complexity in design of vessel forms (Moore and Hunter Anderson, 1999). Ipao Stamped designs are found on a small percentage of bowls, however, the impressions are limited to the rim of the vessel (DeRoo and Goodfellow, 1998; Ray, 1981; Moore, 2002). Thick flat bottomed pans become common with wide openings and shorter heights (Moore, 1 983). Matt impressions are also noted, primarily on the base exterior, however they are sometimes observed on the interior as well (DeRoo and Goodfellow, 1998). The matt impressions are similar to a weaving style found on contemporary sleeping mats from the Caroline Islands made from the pandanus tree ( Pandanus tectorius ) (Safford, 1905; Hunter Anderson et al., 1998). This finding suggests that the pan was placed on the mat while drying to prevent collapse prior to firing (Moore, 2002).

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59 Quartz inclusio ns originating from Saipan have been found on a small number of sherds from this time period suggesting that ceramics were brought to Guam from Saipan (Dixon et al., 1999; Moore, 2000; Hunter Anderson et al., 1998). These foreign sherds are found primaril y on the west coast of Guam, including in Tumon Bay, indicating that inter island contact occurred but was restricted to the western portion of the island. This theory is plausible since the high cliff lines along the east coast would make access to the sh oreline much more difficult by canoe (Moore, 2002). The Transitional (Huyong) period (400 1000 CE ) In the period following AD 400, there is a decline in the use of flat bottomed pans and a change in vessel form with thin walls, round bases, and slightly in curvate rims. Various tempers, including calcareous sand, mixed sands, and volcanic sands, are common with plain or polished/burnished surface treatments. Decorated rims are accompanied by wall perforations, likely for practical and not decorative purpos es, are also seen in this time period (Moore, 2002). Change between vessel form, surface treatment, and tempers were gradual and did not occur concurrently across sites. There are numerous sites dating to this time period, which are found along rivers a nd the coast and include both open air sites and rockshelters (Tomonari Tuggle and Tuggle, 2003). Most notable is Tarague Beach, just north of Tumon Bay. Coastal and river sites of this period have an abundance of cultural materials in comparison to the more inland sites (Moore, 2002). Latte Period (1000 1668 CE ) An increase of charcoal within paleoenvironmental sediment cores around 1,800 BP is suggestive of an intensification of land use and corresponds to the shift from the Pre Latte to Latte periods (Athens and Ward, 2004). Unlike Pre Latte sites, the Latte

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60 deposits did not experience extensive disturbances from the long term effects of storm events, wave action, and sea level changes. Thus, most of the current knowledge of rom the better preserved Latte period sites (Hunter Anderson and Butler, 1991). While ideal settlem ent areas remained in sandy coastal locations, populations also began to move inland to more marginal environments, such as the interior uplands (Figure 2 5 ). Latte architecture The Latte Period begins around AD 1000 and is characterized by the construction of latte sets and Marianas Plainware pottery (Spoehr, 1957; Graves, 1986) (Figure 2 6) The construction of the latte sets began at approximately AD 1000 and by AD 1325 had spread to the northern Mariana Islands, including the northernmost populated island, Pagan (Graves, 1986). Latte use and construction continued throughout the Latte period and into the first part of the Spanish Colonial period (Tomonar i Tuggle et al., 2005). The exact end date for latte construction is unknown, however, it is believed that the disbanding of indigenous settlements resulted in cessation of latte production (Moore, 2002). Several early hypotheses speculate on the functi on and significance of latte sets houses, canoe sheds, and residential structures for the elite (Thompson, 1940; Thompson, 1945; Spoer, 1957; Reinman, 1977). A survey of cul tural material in and surrounding latte sets in Guam, Saipan, Tinian, and Rota revealed varied artifact categories and assemblages that indicating that a wide range of domestic activities were associated with the latte structures and they were not, as prev iously thought, specialized to one function. Artifacts associated with cooking, food preparation, tool

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61 manufacture and maintenance, fishing, and warfare were found homogenously in all the latte sets analyzed (Table 2 2). Additionally, human burials are u niformly found within or around Latte structures (Graves 1986). Most latte sets are arranged parallel to the coastline, however, some are perpendicular to the coast (Thompson, 1940). Multiple latte sets are distributed in a linear pattern, if located alon g a steep cliff, or clustered along long stretches of sandy beach or limestone terraces. In areas where several latte sets are clustered together, the largest latte set is centralized with the smaller sets along the periphery (Graves, 1986). Archaeologic al evidence points to a stratified society with chiefs who organized the labor for construction of the latte sets. It has been hypothesized that latte were (Hunter Anderson, 1989). Further, the practice of burying family members within the latte set further legitimizes the claim through a direct link to the ancestors (Hanson an d Gordon, 1980). At the time of contact, settlements consisted of groups of latte houses nucleated into villages (Tomonari Tuggle et al., 2005). Cooking and food processing Marianas Plainware is thick walled with no slip and little decoration. Pots were typically hemispherical or globular with restricted mouths, thick, incurving rims and rounded bottoms (Moore, 2002). Some vessels displayed grooves perpendicular to the rim, suggestive of being secured with ropes, possibly for suspension, handle for grip ping/pouring, or to secure a lid (Reinman, 1977; Davis et al., 1992; Wickler, 1993; Moore and Hunter Anderson, 2001; Moore, 2002). Many of these potsherds, primarily those with plain and wiped or brushed surfaces, from the Latte Period have charred

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62 residu e on their interior surfaces that is likely associated with cooking (Loy, 2002; Moore, 2002). The wide array of food processing artifacts in the Latte Period are indicative of a more sedentary population with an increased reliance on agriculture ( Hunter Anderson and Butler, 1991 ; Moore, 2005). Stone mortars (Chamorro: lusong) and pounders are found ubiquitously throughout Guam and date almost exclusively to the Latte Period. husk plant products, such as rice, and blades usually made from local shells, are associated with digging into the soil and for scraping, peeling, and cutting tubers, roots, and plant stalks (Moore, 2 005). Additionally, archaeological investigations in Saipan showed a 35% increase of adze production from the Pre Latte to Latte periods (Butler, 1988). Diet Starch grain analysis: Analysis of starch grain residues on 35 Latte period potsherds identifie s taro, Cordyline (type of palm common name Cabbage tree) rice, sugarcane, and possibly shellfish and fowl indicative of cooking these foods Taro was identified on more than half (n = 18) of these potsherds (Loy, 2001a,b; Loy and Crowther, 2002; Crow ther et al. 2003), suggesting preference for and possibly cultivation of the introduced tuber. Radiocarbon dates associated with the archaeological sites from which the potsherds originated, indicate that taro was cooked in clay pots as early as 2000 year s BP (Hunter Anderson et al., 2001; Moore, 2005). Likewise, analysis of pollen in sediment cores indicates that taro was present just prior to and during the early Latte Period (Cummings and Puseman, 1998; Athens and Ward, 1999).

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63 Pollen and phytolyth anal ysis: Pollen data from paleoenvironmental sediment cores have identified a wide range of cultivable plant foods including betel palm, breadfruit, coconut, pandanus, Cordyline Ipomoea (morning glory), and aroids ( Alocasia, Colocasia [taro] and Cyrtosperma ) (Athens and Ward, 1995; Athens and Ward, 1999; Cummings and Puseman, 1998; Dixon et al., 1999; Ward, 2000; Cummings, 2002; Athens and Ward, 2004). Likewise, phytolith analyses from soil samples and sediment cores have also found evidence of breadfruit a nd betel palm, in Anderson, 1994; Pearsall and Collins 2000; Collins and Pearsall, 2001a; Collins and Pearsall 2001b). Lastly, analysis of charred wood samples from southwestern Guam ide ntified coconut, breadfruit, and pandanus (Murakami, 2000). Stable isotope analysis: Studies focusing on stable isotopes for prehistoric diet reconstruction in the Marianas are limited in number and generally have very small sample sizes. However, resear ch has been conducted in Guam, Rota, and Saipan to estimate the proportion of marine versus terrestrial foods in the prehistoric diet of populations inhabiting the Mariana Islands (Hanson, 1989; Quinn, 1990; McGovern Wilson and Quinn, 1996; Ambrose et al., 1997; Pate et al., 2001). In an analysis of four individuals from the Duty Free site on Saipan, Hanson (1989) found consistently low 13 C values indicating a significant dependence on terrestrial foods. The 15 N values are more variable and suggestive of a combination of different marine resources in the diet with a higher emphasis on lagoon resources. Nonetheless, the isotope signatures are reflective of a diet comprised much more of terrestrial foods than marine foods, which is surprising given the p roximity to the ocean.

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64 revealed one individual with a high proportion of open water foods and no reef foods in his diet. Likewise, another male with no access to reef or lagoon areas had a high proportion of the diet comprised of open water marine foods. McGovern Wilson and Quinn (1996) found that marine resources contributed 30% (based on mean 15 N value) and 40% (based on mean 13 C value) of the diet in Saipan, while these val ues are still low, they are higher than expected given the faunal record. Exploitation of pelagic fish is also revealed by the isotope analysis. Additionally, McGovern Wilson and Quinn found no age or sex differences in access to differential resources. Ambrose and colleague between diet reconstruction from collagen versus carbonate carbon isotopes. Bone collagen suggests a 17% marine component to the diet, while the carbonate data indicates a much hi gher estimate of 60%. Nonetheless, the very high apatite carbonate 13 C values suggest a large component of the Saipan diet came from C 4 or marine plants with very low protein component, such as sugar cane and seaweeds. Small amounts of marine foods, lik ely reef and lagoon fish and/or shellfish, contributed to the diet (Ambrose et al., 1997). Data from Rota are similar, yet more diverse than what was noted in Saipan. Pate and colleagues (2001) found much variability between individuals and determined that 22% (ranging from 10% to 41%) of the diet in Rota was derived from marine foods. Some individuals have elevated 15 N values suggesting exploitation of pelagic fish, while others have negative 13 C and 15 N values indicative of a diet dominated by terrestrial foods, suggesting differential access to resources between individuals.

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65 Likewise, Ambrose and colleagues (1997) found that marine resources contributed between 21% and 40% to the diet in Rota. High 15 N values suggest a significant contribution of pelagic fish to the diet in some individuals, while other individuals have low 13 C values suggestive of a predominately terrestrial diet. Stable isotopes in Guam reveal a similar signature as those reported for Rota, however, the isotopic analysis is limited to only five individuals. Marine foods contribute between 27% to 35% to the diet. Collagen 15 N and 13 C values are similar to what is seen in Rota, however, the 15 N values in Guam are slightly lower than those from Rota. This suggests that the diet was comprised primarily of terrestrial C 3 resources, such as rice, root crops, and vegetables, and some marine protein, but little reliance on seaweeds or C 4 p lants (Ambrose et al., 1997). Shellfish consum ption: A transition from bivalve to gastropod consumption (1980) at Ypao Beach, in Tumon Bay, found that Strombus gibberulus gibbosus ( gibbose conch ) far outnumbered any other family. Graves and Moore (1985) also noted the overabundance of gastropods in comparison with bivalves in Tumon Bay, with Strombidae (conchs) comprising 90% of the gastropod assemblage. This shift is attributed to changes in sea level and concomit ant changes in local ecosystems where preferable mangrove environments dissipate once in sea levels decrease (Amesbury, 1999; Graves and Moore, 1985). Post Contact Era The arrival of Ferdinand Magellan and his crew in 1521 on the southern coast of Guam mar ks the beginning of the post contact era as well as the first contact between

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66 Europeans and any indigenous Micronesian populations (Alkire, 1977; Lvesque, 1995). The post contact area is commonly divided into the Spanish Colonial, First American, Japanes e World War II, American World War II, and Second American periods. The Spanish Colonial Period (1521 1898 CE ) At the time of Spanish contact, the indigenous peoples of the Mariana Islands were described as a single population with shared language, cultur e, and customs (Driver, 1983). In 1565 Miguel Lopez de Legazpi landed in Guam and the Marianas from Spain. This led to the start of the Manila galleon route where ships sailed from the Philippines to Acapulco and upon the return voyage, stopped in Rota o r Guam where they bartered iron for fish, fruit, coconut, and rice, with the locals, before the final return back to the Philippines (Schurz, 1939; Driver, 1983). Contact between Europeans and the Chamorro remained limited until 1668 when the Spanish miss ionaries settled in Guam to convert the Chamorro to Christianity. By this time, the missionaries characterized Guam as a thriving culture, with 180 villages, the largest of which had 150 houses (Lvesque, 1995). Juan Pobre, a Franciscan brother who live d on Rota for seven months in 1602, provides the most detailed description of the Marianas in the post contact period. While descriptions of agricultural practices were not reported, he noted that the Chamorro used wooden sticks to dig into the soil. Tar o, yams, rice and a type of sweet potato were grown inland and these agricultural products were traded with villagers living along the coast for fish (Driver, 1989). Local population size for the island at the start of the 17th century is estimated at 20,0 00 but by the end of that century diminished to approximately 1,600 individuals due

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67 to widespread epidemic disease and rebellion after European contact (Russell and Fleming, 1990; Tomonari Tuggle et al., 2005). The Spanish instituted the Reduccin which c onsolidated the entire Chamorro population of the Mariana Islands into seven mission centers, six of which were located in Guam and one in Rota which were under rigid control by the Spanish missionaries and military officers (Rogers, 1995; Tomonari Tuggle et al. 2005). Latte sets were eventually replaced by lanchos small, elevated thached houses, which served as subsistence farms (Carano and Sanchez, 1964; Rogers, 1995). Rice became much more frequently utilized and introduced maize gained dietary importa nce over the previous staples of taro and breadfruit (Rogers, 1995). Other introductions included mango, pineapple, papaya, citrus, hot peppers, and cassava (Carano and Sanchez, 1964; Rogers, 1995). The Spanish also imported farm animals, such as carabao (subspecies of the water buffalo indigenous to Southeast Asia), cattle, horses, deer, pigs, and goats (Farrell, 1991; Hunter Anderson et al., 2001). The presence of alternative animal protein contributed to an overall decline in fishing in favor of farmi on the 17 th century map from the Jesuit Charl es Le Gobien (1700) yet, according to Kurishina et al. (1987; 1988), little activity is known to have occurred in the area durin g this period due to the limited number of Spanish artifacts found during archaeological testing (Kurashina 1987). However, Tumon Bay was used by the Spanish for fishing and likely contributed to subsistence practices of both the Chamorro and Spaniards (B urtchard 1991).

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68 The First American Period (1898 1941 CE ) During the Spanish American War, the United States took control over Guam in 1898, after Captain Henry Glass attacked the Spanish Forts in Apra Harbor and demanded that the Spanish surrender Guam to the US. However, American rule wa s not firmly set until the 1900 s when a centralized, yet unfortified, military presence was established at Orote Peninsula (Rogers, 1995). In 1922, all coastal defenses were removed from Guam with the signing of the Naval Limitations Treaty, leaving Guam unprotected and vulnerable. According to Safford (1903), the local residents resided in six towns and were primarily farmers who traveled to the lanchos for subsistence farming, while fishing practices were greatly reduced. Population estimates for the six villages were: Agafia with 6,400 inhabitants; Sumai, 9oo; Ynarajan, 550; Agate, 400; Merizo, 300; and Umata, 200. No population records for the Tumon Bay area are available for this time period, however, hamlets along the coast composed of few houses are described. The Japanese World War II Period (1941 1944 CE ) On the morning of December 8, 1941 (UTC/GMT +10 hours), the same day of the December 7 th of bombing by Japanese air forces culminated with a three pronged ground attack at Tumon Bay, Agaa Bay, and Merizo, and the forced surrender of Guam to the Japanese by the American gov ernor (Rogers, 1995). The majority of the local population fled to the countryside in an effort to escape the Japanese regime who attempted to enculturate the Chamorro population with forced language change in the school system. During this period, the C hamorro reverted back to traditional practices

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69 of fishing and agriculture, particularly in more remote areas (Russel et al., 1993; Rogers, 1995). Theater, the Chamorro were placed into forced labor camps (Sanchez, 1979). The defense strategy included fortifications along the entire western coast of Guam, particularly Agat and Asan, but in Tumon as well (Craib and Yoklavich, 1996). Pillbox fortifications and gun emplacements were scattered along Tumon Bay and some can still be seen there today. The American World War II Period (1944 1948 CE ) On July 8, 1944 the US began air attacks on Guam and landed on Agat and Asan on July 21 of the same year. After the Orote Peninsula wa s succeeded to the Americans on July 27 th the Japanese abandoned the southern portion of the island and fled to the north for the last hold out. In August, the Americans had officially re captured Guam, although fighting continued until the last of the J apanese forces surrendered in September 1945 (Gailey, 1988; Denfeld, 1997). The Second American Period (1945 Present) A massive military build up by the US armed forces began prior to the end of WWII to assist in continued bombardment of Japan. In 1948, c ontrol of Guam was transferred from the US Navy to the Department of the Interior and the Organic Act of 1950 gave the Chamorro US citizenship. In 1970, the locals elected their first governor and today Guam is a US territory with a continued US military presence (Welch et al., 2005). Extensive damage to Guam occurred due to both massive bombing and post war modification of the island landscape through bulldozer clearing and leveling by the US Navy Seabees and extensive construction. The area of Tumon Bay is currently a

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70 popular tourist area and has undergone significant alterations due to post war construction of resorts and parks for recreational use (Welch et al., 2005).

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71 Table 2 1. Archaeological and historical chronological s equences of the M arianas Islands Dates Spoehr (1957) Craib (1990) Moore and Hunter Anderson (1999) Hunter Anderson and Moore (2001) 0 BP AD 1521 Historic (Protohistoric) Latte AD 1521 AD 1521 500 BP 1000 BP Latte Latte Mochong Latte Latte 1500 BP Pre Latte Transitional Ypao Transitional Huyong 2000 BP Intermediate Pre Latte Late Unai 2500 BP Intermediate Pre Latte 3000 BP Tarague Early Pre Latte Middle Unai 3500 BP Early Unai Table 2 2. Activities and artifact assemblage composition of latte sets a Cooking Tool Manufacture/Maintenance Fishing Warfare Prepared coral floors Shell and stone debitage Stone and shell net sinkers Slingstones Ovens/cooking areas Unfinished fish hooks Stone and shell line sinkers Bone spear points Fire cracked rock and charcoal Tridacna shell blanks Bone and shell fishhooks Pottery Chert or basalt cores Fish gorges Storage and cooking vessels Hammerstones Mortars and pestles Adzes Stone drills/perforators Shell/coral files Bone awls and needles a. Compiled from Graves (1986) survey

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72 Figure 2 1. The Mariana Island Chain ( Source: http://commons.wikimedia.org/ wiki/File:Casta_Marianas.jpg Accessed on 22 July 2012)

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73 Figure 2 2. Paleoshoreline notches along the western coast of Guam (Photo by author) Figure 2 3 (Photo by author)

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74 Figure 2 4 Examples of Pre Latte Period pottery. Photo taken at Cultural Park Museum (Photo by author)

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75 Figure 2 5 Island of Guam depicting Latte Set density and distribution, follo wing s survey ( Map courtesy of Rad Smith/GANDA )

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76 Figure 2 6 Examples of Latte Period pottery. Photo taken at Cultural Park Museum (Photo by author)

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77 CHAPTER 3 MATERIALS Archaeological Sample From 2006 to 2008, 376 prehistoric Chamorro burials were excavated by Paul H. Rosendahl, PhD, Inc., a cultural resource management firm, which was later incorporated into SWCA Environmental Consultants, during an archaeological mitigation project for the renovation of the Guam Aurora Villas & Spa in the Naton Beach Site on the northern end of Tumon Bay, Guam. Of these, approximately 177 are associated with the Pre Latte Period and 190 belong to the Latte Period. The location, associated artifacts, and radiocarbon dates (DeFant, 2008). Dating to roughly 2,500 BP, the Pre Latte group repres ents some of the earliest settlers in Guam and is the largest Pre Latte mortuary sample discovered to date (Table 3 1). There are no radiocarbon dates for the Latte sample; however, archaeological materials associated with the Latte remains place the rema ins in the Latte period between AD 1000 to 1521. A comprehensive osteological analysis was conduct ed by Ms. Cherie Walth, of SWCA All demographic information, as well as information on cranial and postcranial metrics and non dental pathological condition s, cited within this dissertation has been provided by Ms. Walth and SWCA. A complete archaeological and osteological report on the Naton Beach Burial Complex including preliminary results from the current study is being prepared for submission as an un published manuscript to the Guam Historic Preservation Office.

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78 Taphonomic Bias Bioarchaeological analysis in the Marianas is constrained by a number of taphonomic factors. As is the case in many island environments, poor bone preservation is the result of fluctuating temperatures, differential exposure to wet and dry environments, and soil factors such as limestone based soil matrices (Hanson and Butler, 1997). Coastal skeletal assemblages are further perturbed by the effects of changing sea levels (Dicki nson, 2003) the mechanical effects of the waves leading to soil and burial disturbance (Graves and Moore, 1985; Butler, 1995) and commingling of neighboring burials. A more prolonged exposure of Pre Latte skeletal remains to the natural environment, cha nging ecosystem, and human interaction may explain the dearth of mortuary assemblages from the earliest inhabitants of Guam. Additionally, the Pre Latte settlements and burials are located much deeper in the ground and thus are not as frequently encounter ed (Butler, 1995; Hunter Anderson and Butler, 1991). Thus, an overwhelming majority of skeletal studies in Guam focus on the Latte Period as Pre Latte remains were rarely recovered or too fragmented to provide any information on the biological variation o Disturbances due to human activity also affect bone preservation, as a rchaeological studies have noted that in many cases, skulls and long bones are often missing from primary interments (Thompson, 1932; Spoehr, 1957; Reinman, 1977). Ethnohistoric accounts note possessed power, which could be called upon by descendants for supernatural purposes (Thompson, 1940; Driver, 1983). Thus, skeletal remains were b uried beneath the Latte sets, as a means for the familial social unit to demonstrate their loyalty and

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79 identify with a particular lineage as well as to show strength through close proximity to their ancestors (Thompson, 1945; Graves, 1986). Further, Latt e burials were often placed in shallow pits, close to the surface, allowing for easy access and selective removal of skeletal elements: skulls were most often removed from male burials, while long bones were often taken from female burials (Spoehr, 1957; Yawata, 1961; Reinman, 1977; Hanson and Gordon, 1989). A regard are the skulls of the ancestors, especially those of their paren ts and which were often used in ritualistic activities. Long bones, on the other hand, were more utilitarian in purpose and were used to make tools and weapons, such as spearpoints, harpoons, and awls (Hanson and Gordon, 1989). Prehistoric habitation sites along Tumon Bay have also been impacted by modern modifications to the landscape. During the 1930 s the Tumon coastline was altered during the occupation of the Japanese who built commercial establishments and extensi ve fortifications around the bay (Defant 2002). Post war modification of the entire island landscape occurred through bulldozer clearing and leveling performed by U.S. Navy Seabees after the American take over (Fulmer et al., 1999). Additionally, in rece nt times, Guam has become an increasingly popular destination for tourists over the past two decades. The increase in tourism brought with it rapidly expanding economic growth (Hanson and Butler, 1997). Thus, new developments such as hotels, strip malls, and parks are being built along the bay leading to the accidental disinterment of skeletal assemblages. Effects of building and construction, using large machinery can result in fragmentation and commingling of skeletal elements.

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80 Sample Population A to tal of 215 individuals dentitions were analyzed from the Naton Beach mortuary sample, 103 from the Pre Latte Period and 112 from the Latte Period (Table 3 2). Females, n = 77, were slightly more represented than males, n = 62; however, 76 were of un determ ined sex (Table 3 3). Age cutoffs for the current study are as follows: child < 10 years of age; juvenile = 10 to 18 years of age; young adult = 18 to 25; middle aged adult = 30 to 45; older adult > 40 years of age. The majority of the assemblage is comp osed of adult individuals (n = 179). O nly juveniles with pe rmanent dentition (n = 36) were analyzed are included in this analysis. The majority of the sample is composed of young adults (n = 107), followed by middle aged adults (n = 55), and old adults ( further delineation of age grouping (Table 3 4). Pre Latte Demographics The Pre Latte sample is represented by 103 individuals, 38 females and 33 males. Sex could not be deter mined for 32 individuals. All but 11 individuals are adults (n = 93). More than half of the Pre Latte assemblage is composed of young adult individuals (n = 64; 62.1%), followed by middle aged adults (n = 23; 22.3%). No old adults were analyzed. For 5 individuals, only a category of adult age was possible (Table 3 5). Latte Demographics The Latte sample is represented by 112 individuals. Males (n = 29) are underrepresented in the sample in comparison with females (n = 39). Sex could not be det ermined for 44 individuals. The majority of the Latte Period individuals are adults (n = 87), with only 25 subadults represented in the sample. Young adults (n = 43; 38.4%) make up the majority of the population, followed by middle aged adults (n = 32; 2 8.6%),

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81 and lastly old due to lack of other skeletal indicators to further delineate age (Table 3 6)

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82 Table 3 1. Radiocarbon dates from Naton Beach site, Tumon, Guam. a Sample No. Provenience Material Measured radiocarbon age Conventional radiocarbon age age range Beta 238482 Feature 2 Soil 170040 BP 168040 BP AD 250 430 Beta 238483 Burial 173 b Conus shell beads 249040 BP 294040 BP 770 400 BC Beta 238484 Burial 156 b Conus shell beads 233040 BP 279040 BP 590 330 BC Beta 238485 Burial 273 b Conus shell beads 249040 BP 286040 BP 720 360 BC Beta 238486 Burial 286 b Conus shell beads 264040 BP 297040 BP 790 490 BC a Table recreated from Defant (2008:150). b All burials are from the Pre Latte assemblage.

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83 Table 3 2. Pre Latte vs. Latte sample distribution Time Period N Pre Latte 103 Latte 112 Total 215 Table 3 3. Okura dental sex distributions Frequency Percent M 62 28.8 F 77 35.8 INDT 76 35.3 Total 215 100 Table 3 4. Okura dental age distributions Frequency Percent CLD 18 8.4 JUV 18 8.4 YA 107 49.8 MA 55 25.6 OA 4 1.9 ADT 13 6 Total 215 100 Table 3 5. Pre Latte dental sample by age and sex Pre Latte Male Pre Latte Female Pre Latte Indet. Total n % n % n % n % CLD 0 0.0 0 0.0 6 18.8 6 5.8 JUV 0 0.0 2 5.3 3 9.4 5 4.9 YA 19 57.6 30 78.9 15 46.9 64 62.1 MA 13 39.4 6 15.8 4 12.5 23 22.3 OA 0 0.0 0 0.0 0 0.0 0 0.0 Adult 1 3.0 0 0.0 4 12.5 5 4.9 TOTAL 33 100.0 38 100.0 32 100.0 103 100.0 % is shown as percent of female/male/sex indeterminate quantity except for totals which are based on percent of total sample.

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84 Table 3 6. Latte dental sample by age and sex Latte Male Latte Female Latte Indet. Total n % n % n % n % CLD 0 0.0 0 0.0 12 27.3 12 10.7 JUV 1 3.4 3 7.7 9 20.5 13 11.6 YA 14 48.3 16 41.0 13 29.5 43 38.4 MA 11 37.9 16 41.0 5 11.4 32 28.6 OA 1 3.4 3 7.7 0 0.0 4 3.6 Adult 2 6.9 1 2.6 5 11.4 8 7.1 TOTAL 29 100.0 39 100.0 44 100.0 112 100.0 % is shown as percent of female/male/sex indeterminate quantity except for totals which are based on percent of total sample.

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85 CHAPTER 4 DENTAL REDUCTION Over time, and throughout the world, there has been an overall decrease in tooth size in the human dentition (Brace et al. 1987). The majority of the studies looking at dental reduction focus on dental trends over relatively long periods of time, from pr ehistoric Pleistocene populations to contemporary populations (Dahlberg, 1960; Greene, 1972; Carlson and van Gerven, 1977; van Gerven et al., 1977; Smith et al., 1984; Calcagno, 1986; Smith et al., 1986; Brace et al., 1987; Calcagno, 19 1989). O thers have focused on microevolutionary trends of dental reduction across shorter periods of time. For example, Christensen (1998) performed an odontometric analysis in a prehistoric Oaxaca V alley population spanning from 2600 BP to 1521 CE He found a dramatic 4.4% reduction in size between the earliest and the latest temporally dispersed groups. Pinhasi and colleagues (2008) found uniform reduction in the buccolingual dimensions of the dentitions of Southern Levant populations ranging from 12,000 to 7,000 BP Brace and colleagues (1987) suggest that dental reduction began in humans during the Late Pleistocene at a rate of 1% over 2,000 years. However, the rates of reduction have increased to 1% every thousand years, from the Post Pleistocene to the current day (Brace et al., 1987) Background The exact mechanism for decrease in tooth size is unknown, however, four main theories aim to explain this phenomenon. The Probable Mutation Effect (PME), as proposed by Brace and colleagues (Brace 1963; Brac e and Mahler 1971; Brace and Hinton 1981; Brace 1987), suggests that mutations are the primary forces acting on dental reduction. Whereby, a relaxation of selective forces due to a change in food

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86 processing techniques, may allow for an accumulation of mut ations, which lead to decrease d tooth size. Calcagno (1986, 1989) introduced the Selective Compromise Effect (SCE), suggesting that decreased surface area and tooth complexity results in such, selection for smaller teeth is the result of an overall decrease of dental dimensions leading to dental crowding and increased potential for cariogenic disease. The Increasing Population Densi ty Effect (IPDE) suggests that selection for smaller body size, due to increased population density, sedentism, and reduced nutritional requirements, results in a reduction of tooth size (Macchiarelli and Bondionli 1986). Lastly, Carlson and Van Gerven (1977) adopt a biomechanical approach in their Masticatory Functional Hypothesis (MFH) and suggest that shifts in subsistence patterns, such as consuming softer and more processed foods, led to selection for smaller teeth They propose that a shortening of the craniofacial complex, as a result of decreased functi onal demands of mastication, led to a compensatory reduction in the size of the dentition. Current Study The current study investigates evolutionary dynamics of the prehistoric Chamorro population to see how they relate to biocultural and environmental c hanges in prehis toric society. The Pre Latte (3500 BP 1 000 CE) and Latte (1 000 1521 CE ) periods in Guam convey distinctions, not only in population size, but also in diet and subsistence strategies. The Pre Latte are characterized by small population s with semi permanent settlements, that subsisted on bivalve shellfish, reef fishing, minimal agriculture, and limited terrestrial resources consisting of birds, crabs, and bats (Amesbury et al., 1991; Hunter Anderson and Butler, 1991; Pietrusewsky et al., 1997).

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87 Thin pottery, associated with cooking, has also been found in association with Pre Latte sites (Amesbury et al., 1991). A significant increase in population numbers and densities occurred during the Latte period ~ 1000 AD coupled with an intensi fication of agriculture ( Hunter Anderson and Butler, 2001 ) While archaeological records indicate that there was a shift from bivalves towards pelagic fish and gastropods (Graves and Moore, 1985), exploitation of marine resources declined in the Latte Per iod (Butler, 1988; Ambrose et al., 1997; Pietrusewsky et al., 1997). Material artifacts associated w ith food procurement and cooking include mortars and pestles for food processing, thick pottery, bone spear points, and composite fish hooks (Ambrose et al ., 1997, Amesbury et al., 1991). Previous investigations show that the Chamorro dentition is some of the largest in the world, intermediate between the larger Melanesia n dentition and the smaller Polynesia dentition (Brace et al., 1981; Brace et al., 199 0; Hanihara and Ishida, 2005). However, these studies pool samples of varying time periods and island samples together in their investigations. The large Naton Beach skeletal assemblage allows for within site diachronic comparisons to be made of the earl ier Pre Latte settlers to the later prehistoric inhabitants. This study therefore represents the first large within site diachronic study of dental size reduction in Guam. While the aforementioned accounts have shown that dental changes can occur over s hort evolutionary time spans, the Pre Latte and Latte periods span an even narrower time range, around 1500 years, than those previously investigated. However, two studies comparing immigrant dental dimensions in contemporary populations showed significan t differences between parents and offspring dental sizes due to

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88 differential access to nutritional resources (Goose, 1967; Goose and Lee, 1973). Thus, change in tooth size can occur over relatively short periods of time. The null hypothesis is as follows : H o : There is no significant reduction in the dental dimensions between the Pre Latte to Latte time periods. I hypothesize that there will be dental reduction between the Pre Latte and Latte due to selective pressures during odontogenesis as a result of an increase in population size (followed by decrease in health and competition for food resources) and change i n diet and food procurement strategies. Expected Results If a change is found between the Pre Latte and Latte samples the trends should follow one of the proposed models for dental reduction and its associated assumptions (Pinhasi et al., 2008). With the PME, a change in cultural practices specifically related to food preparation, such as tools and techniques for cooking, allow for a relaxation of selection pressures in maintaining large teeth. Given the state of relaxed selection, mutations begi n to accumulate (Brace, 1964; 1967; 1978; Brace and Mahler, 1971; Wolpoff, 1971; mutation pressure, which suggests that mutations lead to a reduction in structures, Brace and co lleagues posit that dental size decreases following selection relaxation and accumulation of mutations. If the PME is the model by which reduction occurs, reduction should occur equally in both the mesiodistal and buccolingual dimensions from one time per iod to the next (Christensen, 1998; Pinhasi et al., 2008).

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89 In SCE, there is a compromise between selection for large, complex teeth with thick enamel and small teeth with thin enamel and less morphological complexity. Large complex teeth provide a larger surface area for carious lesions and can contribute to dental crowding. However, a large surface area and thick enamel are well adapted to populations ingesting a coarse diet. S maller teeth are more resistant to dental caries and crowding but are less re sistant against the high biomechanical demands of a course diet (Calcagno, 1986; 1989). Additionally, untreated carious lesions would have been problematic and may have lead to decreased health in the population. Thus, in populations who have smaller for ces placed on the dentition and are more prone to dental caries due to a softer diet, selection would be for smaller teeth and an overall reduction in the masticatory apparatus (Calcagno, 1986; 1989). If SCE is at work in the temporally dispersed Chamorro populations, the buccolingual and mesiodistal dimensions and tooth types should be differentially affected, with a constant amount of variation between the time periods (Christensen, 1998; Pinhasi et al., 2008). Premolars and molars are most likely to sh ow greater dental reduction due to increased complexity and higher prevalence for caries. The MF model falls under the SCE theory (Calcagno, 1989). It suggests, in maxilla will undergo apposition or resorption due to the biomechanical stressors placed on them. Since teeth are more genetically controlled, a decrease on functional demands placed on the masticatory apparatus would result in a decrease in jaw size but not tooth size, thus causing dental crowding. Selective pressures would then restore harmony by reducing the size of the dentition for a better fit within the maxilla and

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90 mandible (Calcagno, 1989). If dental reduction follows the MF model, similar results as what is expected from the SCE model would be expected. The varying effect on the two dimensions does not suggest that a full sweep reduction of the masticatory apparatus is at work. Instead, the dental dimensions are affected differently due to changing func tional demands placed on the skull and are not the result of overall decrease of body size. Tooth classes are expected to be affected equally as a result of overall dental reduction. The IPDE model suggests that tooth reduction is a byproduct of smaller b ody size as a result of an increase of population density. Thus, new adaptive pressures, such as environmental stress due to a decrease in nutritional resources, trigger selection for a reduction of body size and as such, dentition also becomes smaller (M achiarelli and Bondioli, 1986; Pinhasi et al., 2008). If IPDE is at work, all teeth and both dimensions should be affected equally. Further, dental reduction should correspond to increase in population size and carious lesions (or other pathological indi cators) (Christensen, 1998; Pinhasi et al., 2008). The data presented in this study will be analyzed to determine if there is a trend in dental reduction. If reduction is observed, the aforementioned models will be examined to determine which best fits the data of the current study Methods Two standardized measurements were taken on each available tooth using Mitutoyo Digital Extended Pointed Jaw calipers following Moorees (1957) and Mayhall (1992). These fine tipped calipers allow for precise measurem ents of both isolated teeth and those within the alveolar process. The mesiodistal diameter (MD), or the length of the crown, was obtained by measuring the greatest distance between the

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91 mesial and distal portion of the tooth, as expected in proper anatomi cal position. The buccoli n gual diameter (BL), or the width of the crown, was obtained by measuring the width of the tooth, perpendicular to the mesiodistal plane. All measurements were taken to the clos est 0.01 mm and were not taken o n teeth with moderat e to extreme amounts of wear. Following Brace (1979, 1980), the mesiodistal and buccolingual diameters were multiplied to attain the cross sectional area (CX) of each tooth class. The left and right antimeres of each tooth are representative of the same genotype, thus two sides were averaged to best express the cross sectional area of that tooth class (Brace, 1990). Tooth summaries (TS) were also calculated following Brace (1978). The tooth summary is the sum of the upper and lower mean cross sectional areas of each tooth category. This number allows for a quick comparison of mean tooth size between groups and represents an approximation of the total occlusal area of the population. The data were tested statistically and visually for normality using t he Kolmogorov Smirnov test and normal Q homogeneity of variance was also performed on each measurement. A two way factorial Analysis of Variance (ANOVA) was performed to evaluate the effect of time pe riod and sex on each measurement using the following model: Tooth measurement = Time Period + Sex + Time*Sex + error where the tooth measurement is the dependent variable, time period and sex are the independent variables (fixed factors), and Time*Sex is t he interaction between those two variables.

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92 In order to prevent Type I error, in where a null hypothesis is rejected when in actuality, it should not be rejected, a sensitive post hoc Bonferonni correction was applied to prevent inflation of the alpha leve l when conducting multiple tests (Abdi, 2007). The Bonferonni corrected alpha was calculated by taking the desired alpha (0.05) divided by the number of ANOVAs run (n = 65). Thus, the Bonferonni alpha is 0.00078. Significance levels of the models were a nalyzed using the Bonferonni alpha to determine which measurements are significant. Once these measurements were identified, significance levels of the time period, sex, and interaction between the two were evaluated using a 0.05 alpha level. Results A to tal of 215 individuals permanent dentitions were analyzed. From these, 1242 individual teeth were measured. Appendix A presents the overall descriptive statistics of the dental metrics and the cross sectional areas of the teeth divided by sex and populat ion. The Pre Latte sample is composed of 633 teeth. Female teeth (n = 299) are represented more frequently than male teeth (n = 189). The Latte sample is composed of 609 teeth. Again, female teeth (n = 243) are represented more often than male dentition (n = 177). The tooth summary for the Pre Latte population is very large, 1423.2 mm (Table 4 1). As expected the male dentition is relatively larger than the female teeth (TS = 1470.9 mm and 1383.6 mm, respectively). The largest measurement of the dent al arcade is the mesiodistal diameter of the mandibular first molars in both males and females (male: L = 12.7 mm; R = 12.9 mm and female: L = 12.5 mm; R = 12.3 m m) (see Appendix A Table A 3). However, in terms of overall cross sectional area, maxillary

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93 first molar is the largest tooth in both males (AVG CX 1 = 147.5 mm 1 ) and females (AVG CX 1 = 138.2 mm 2 ) (see Appendix A, Table A 5). The overall tooth summary for the Latte sample is large, 1307.1 mm, however, it is significantly smaller than the Pre Latte tooth summary size. As expected, the tooth summary for the male dentition (TS = 1377.0 mm) is larger than that of the female dentition (TS = 1250.2 mm). Following the trend seen in the Pre Latte dentition, both the mandibular first molar is the largest measurement (males: L = 12.7 mm; R = 12.6 mm and females: L = 12.1 mm; R = 12.1 mm) (see Appendix A, Table A 3). In males, the largest tooth is the mandibular first molar with a cross sectional area of 139.1 mm 2 However, the maxillary first molar is the largest tooth in females with a cross sectional area of 128.1 mm 2 (see Appendix A, Table A 5). A comparison of group measurements and cross sectional area shows the direction of the differences between the groups (Appendix A 6). All of the variables except LMax C BL and LMax C CX have a larger mean value in the Pre Latte group versus the Latte, showing reduction has occurred in the dentition. The percent of change shows which teeth have the highest and lowest rat es of reduction. Overall, the percent of change in cross sectional areas are larger than in the buccolingual than in the mesiodistal diameters. The biggest difference is seen in the cross sectional area of the mandibular third molars: R mandibular molar (20%) and L mandibular molar (17%). The following teeth display a 10% or more decrease in cross sectional areas: left and 1 This cross sectional area is the average measurement of t he left and right sides.

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94 right maxillary molar (14%), right fourth premolar (11%), left mandibular second molar (10%) and, left and right mandibular first inc isors (10%). Kolmogorov Smirnov tests of normality showed that the majority (52 of 64) of measurements were normally distributed (Appendix A 7). Likewise, the data are tested for homoscedasticity using L have equal variance ( Appendix A 8). A two way factorial ANOVA was run on each of the 64 measurements ( Appendix B, Tables B 1 through B 64). Thirty nine of the 64 models were found to be statistically significant using the Bonferonni corrected alpha of 0.00078 (labeled red in Appendix B, Tables B 1 through B 64). Of these significant models, 28 have statistically significantly differences between the time periods, 29 had significant sex differences, and four had a significant interaction between time and sex, at a 0.05 alpha l evel (labeled green in Appendix B, Tables B 1 through B 64). Pre Latte and Latte Differences Very few measurements of the anterior dentition (incisors and canines) are significantly different between the two time periods. The only anterior teeth with stat istically significant differences are the left maxillary incisor BL, left mandibular incisor MD, and left mandibular canine BL. The remaining 25 measurements with significant differences are in the premolars and molars. Comparing the maxillary and mandi bular dentition shows that a little less than a quarter of the measurements with significant differences are located in the upper jaw (15 of 64; 23.4%) and half are from the lower jaw (13 of 64; 20.3%), with the maxillary measurements represented at a sli ghtly higher rate.

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95 The number of significant differences of the buccolingual measurements far outweighs the mesiodistal measurements. Seventeen buccolingual measurements (26.6%) are significantly different between the time periods while only eleven of 64 (17.2%) of the mesiodistal measurements are significantly different between the time periods. Male and Female Differences As was seen in the time period differences, the majority of the measurements that display significant differences between the sexes c ome from the posterior dentition. Only seven anterior teeth display significant differences between the sexes and only one of these measurements comes from the left maxillary lateral incisor BL. The left and right maxillary MD as well as both the buccoli ngual and mesiodistal measurements for the right and left mandibular canines are also significantly different between the sexes. The remaining 22 measurements with significant differences are on the premolars and molars. Significant sex differences betwee n the maxillary and mandibular dentition are found in approximately 20% of the upper and lower teeth ( 21.9% and 23.4%, respectively). Sex differences between the buccolingual and mesiodistal measurements are not as pronounced as was seen when comparing ti me periods. Significant differences in dental dimensions are nearly equal, with 25.0% (16 of 64) of the buccolingual measurements and 20.3% (13 of 64) mesiodistal measurements being significantly different. Interaction b etween Time and Sex Only six measur ements have a significant interaction between time period and sex: left maxillary lateral incisor BL, left mandibular fourth premolar MD, right

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96 mandibular canine BL, right mandibular second molar MD, right mandibular fourth premolar BL, and right mandibula r first molar MD. Two of the six measurements are in the anterior dentition, while the remaining measurements belong to the posterior teeth. All measurements, except the upper incisor dimension, come from the mandible. The buccolingual and mesiodistal m easurements are represented equally. Discussion Tooth Summaries and Rate of Change In both samples the male tooth summaries are larger than the female tooth summaries, indicating some degree of dimorphism between males and females. However, there is a sl ightly larger discrepancy in male to female tooth size in the Latte sample (1377 mm and 1250 mm, respectively) than in the Pre Latte sample (1471 mm and 1384 mm, respectively). Thus, the Latte sample displays a higher degree of sexual dimorphism than the Pre Latte sample, with a 10% difference between sexes compared to a Pre Latte difference of 6%. Overall, the Pre Latte tooth summary is much larger than the Latte tooth summary (1423 mm vs. 1307 mm). Both overall tooth summaries and those separated by sex, the Pre Latte tooth summaries are much larger than the Latte tooth summaries. The s e data points to a small amount of dental reduction over time. Overall, the dentition has reduced in size by 8% between the Pre Latte and Latte time periods. Previous studies in the prehistoric Chamorro populations in Guam, as shown in Table 4 2 (see table for references) have shown tooth summaries ranging from 1034 mm to 1487 mm (Bath, 1986; Pietrusewsky, 1986; Douglas and Ikehara, 1992; Tre mbly, 1999; Pietr usewsky et al., 2003 ). The Leo Palace sample, recovered from Tumon Bay, south of the current Naton Beach sample, represents the smallest tooth size in the

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97 range T he largest tooth summary comes from the Fujita sample, which were also rec overed from Tumon Bay. The Naton Beach tooth summaries from both time periods fall within this range. The Pre Latte sample summaries fall on the high end of the range, closest to the Fujita sample with a tooth summary of 1423 mm. Interestingly, the Fuji ta sample is the only other Pre Latte sample available for comparison. The Latte Period sample from Naton Beach has the largest tooth summary size (1307 mm) in comparison to the rest of the dentition from the Latte Period and is followed closely by the S an Vitores, Right of Way sample (1294 mm). In the remaining comparative samples, the tooth summaries fall far below 1300 mm. A comparison of the Pre Latte and Latte samples show that the Pre Latte teeth, from both Naton Beach and Fujita, are much larger than all of the Latte groups analyzed. When comparing the Chamorro dentition to other samples in the Asia Pacific region, the Pre Latte Fujita sample (1487 mm) has the largest tooth summary and is even larger than the oft reported largest Australian (1486 mm) and Tasmanian (1429 mm) dentitions (Table 4 3 see table for references ). The tooth summary size of the Pre Latte Naton Beach sample of the current study (1423 mm) follows closely behind the Tasmanian tooth summary size. The Latte Naton Beach sample (1307 mm) is relatively smaller and falls between the Hornbostle Thompson Collection in Guam (1309 mm) and the Vanuatu sample (1295 mm). Analysis of the group comparison of mean measurements and cross sectional areas shows the difference in mean measureme nts over time as well as the percent of change. In all of the variables, except for two, the measurements and cross sectional areas show a reduction in mean size over time. Two variables left maxillary canine, BL

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98 and CX, show a slight increase in size fr om Pre Latte to Latte time periods. The buccolingual measurement shows a rate of change of 0.62% while the cross sectional area variable has a 1.26% of change. Regardless of direction of change, these numbers are quite small and may not have any biolog ical significance or may simply largest amount of dental reduction occurred in the lower third molars followed by the upper second molars, upper right fourth premol ar, and lower first incisors. Hypothesis Testing When looking at the Pre Latte tooth summaries from both Naton Beach and Fujita, it is apparent that the Pre Latte teeth are larger than all other reported tooth sizes from the Latte Period. Hypothesis testi ng, using two way factorial ANOVA on 64 variables, was performed to determine the significance of these observed differences. Of the variables, 61% of these models are significant. Tooth size is significantly affected by time period in 44% of the cases, by sex in 45% of the cases, and the interaction between time period and sex in only 6% of the cases. While sex seems to have a slightly higher effect on the size of the dentition, there is not a very large interaction between time period and sex. Thus, t here does appear to be a significant decrease in size between the Pre Latte and Latte samples in some measurements and not others. The null hypothesis of no difference in tooth size between the time periods can be rejected. Time Period Differences The maj ority of the significant differences between the Pre Latte and Latte are relegated to the buccal dentition with only 5% of the significant differences belonging to

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99 the incisors and canines. Thus, over time the premolars and molars are reducing at a signif icant rate while the anterior teeth are much more stable between the time periods. There does not appear to be a large discrepancy in change in tooth size between the maxillary and mandibular arcades. Approximately 20% of the significant measurements ar e found in both the upper and the lower jaw, with the maxillary dentition reducing at a 3% higher rate. The data show that both the maxillary and mandibular dentition are reducing at approximately the same rate over time. The buccolingual measurements red uce at a much higher rate than do the mesiodistal measurements Approximately one quarter, 27%, of the buccolingual variables are found to be significantly different between the time periods, while only 17% of the mesiodistal measurements are affected by time. This finding is line with the results from craniofacial data 2 which show a higher decrease in width measurements, in both the cranium and mandible, in comparison with length measurements over time (see detailed discussion below) (Walth, pers. comm) Comparison of dental, craniofacial, and postcranial changes across time 2 In the following section, caution must be taken due to the fragmentary nature of the skeletal remains and small sample sizes. Very few intact crania and long bones were found and many of the estimates are taken on reconstructed bone. Thus, the data presented may not represent the amount of variation seen in the population. Nonetheless, limited data were collected and analyzed in an attempt to better understand the differences in cranial size and height between the time periods. 2 All cranial and postcranial measurements were taken on the Naton Beach sample by Cherie Walth. Her data, in conjunction with the data from the current study, were collected as part of a larger study on the Naton Beach Burial Com plex, which is currently being compiled for submission to the Guam Historic Preservation Office.

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100 Table 4 4 presents the cranial and mandibular measurements collected by Walth (pers. comm.). Cranial length, cranial breadth, bizygomatic breadth, bimaxillary breadth, and maximum alveolar breadth all show a decrease over time (Walth, pers. comm.) The largest difference between the samples is seen in width measurements: cranial breadth and bimaxillary breadth. Both males (Pre Latte: 144.3 mm; Latte: 133.7 mm) and females (Pre Latte: 136.0 mm; Latte: 128. 8 mm) display a decrease of nearly 10 mm in cranial breadth (Walth, pers. comm.) While there are no data on bimaxillary breadth in the Pre Latte females, the males show a large reduction of maxilla width (Pre Latte: 123.0 mm; Latte: 106.0 mm) (Walth, per s. comm.) Likewise, mandibular width has also undergone a decrease in size over time, particularly in the bigonial and bicondylar measurements. Bigonial breadth in females is unchanging over time (approximately 97 mm in both time periods); however, mal es show a large decrease from the Pre Latte (114.8 mm) to the Latte periods (106.5 mm) (Walth, pers. comm.) The bicondylar breadth also displays differences between the time periods in both males (Pre Latte: 136.0 mm; Latte: 126.1 mm) and females (Pre La tte: 122.5 mm; Latte: 113.9 mm) (Walth, pers. comm.) The cranial length trends differ between males and females. While the male length decreases slightly over time (Pre Latte: 187.3 mm; Latte: 183.5 mm), the female length actually increases, albeit ver y little, over time (Pre Latte: 176.3 mm; Latte: 179.0 mm) (Walth, pers. comm.) Comparably, mandibular length values remain fairly constant in both males and females over time (Pre Latte: M: 79.5 mm, F: 76.5 mm; Latte: M: 79.2 mm, F: 75.4 mm) (Walth, per s. comm.)

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101 The cranial and dental findings are complementary. Over time, there is an overall trend in the reduction of the width of the craniofacial complex coupled with a significant decrease in many buccolingual dental measurements. On the other hand, variables that correlate with craniofacial length, such as cranial an d mandibular length, as well as mesiodistal diameter, remain fairly stable. Stature estimates serve as a proxy for overall body size; however, as stated above, caution must be taken with the following approximations as taphonomic factors leading to severe fragmentation of the bone resulted in small sample sizes. The Pre Latte group, in particular, has a sample size of 12 for males and three for females. (pers. comm.) 2 show very little difference in stature between the time periods. The Pre e individuals are shorter than the Pre Latte group, with the mean stature differences varying by approximately one inch. While the sample size for stature estimates is too small to make any definitive conclusions, nonetheless, the observed decrease in sta ture is so minimal that it is unlikely that such a drastic decrease in the size of the dentition is the result in overall body size reduction Carious lesions The data on carious lesions are not discussed in detail in this section, as they will be fully ad dressed in Chapter 6. However, a brief overview of the carious lesion data will be presented to evaluate the modes of dental reduction. The data show a drastic drop in the overall frequencies of carious lesions between the Pre Latte and Latte periods. L ooking at carious lesion frequency per individual, the male frequencies drop from 74% to 28% while the female frequencies decrease from 76% to 25%. However,

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102 this drastic decrease in infectious disease may have more to do with a the introduced cultural pra ctice of betel nut chewing, which has cariostatic properties, as opposed to an selection for smaller and less complex teeth Further, physiological stress as expressed by linear enamel hypoplasias, becomes more frequent in the Latte period (see Chapter 5 for a more detailed discussion), serving as additional evidence that health status did not between the Pre Latte and Latte periods Mechanism for Dental Reduction Various mechanisms for dental reduction have been previously discussed. To reiterate, thes e theories are: Probable Mutation Effect (PME) Relaxation of selective forces due to a change in food processing techniques Increasing Population Density Effect (IPDE) Overall reduction of body size (and thus tooth size) due to increased population den sity, sedentism, and reduced nutritional requirements Selective Compromise Effect (SCE) Increase of fitness concomitant with decrease in carious lesions due to reduction of tooth complexity as a result of the reduction of tooth size Masticatory Functiona l Hypothesis (MF) Decreased functional demands of mastication as a result of a shortened craniofacial complex. These mechanisms are not necessarily mutually exclusive and it is possible that a combination of these may be at work and the result of dental reduction over time. Probable Mutation Effect Brace and Hinton, 1981; Brace, 1987) as an explanation dental reduction is based on nd accumulation. Relaxation of selection occurs after the advent of new cultural practices involved with cookery followed by an accumulation of mutations. Archaeological data suggest that there are

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103 changes in the exploitation of certain foods and also a n increased amount of food processing associated with the Latte p eriod populations ( Graves and Moore, 1985; Hunter Anderson and Butler, 1991 ; Moore, 2005 ) If PME were to work, it would not be advantageous for one dental dimension to decrease more than another ; thus, both the mesiodistal and buccolingual dimensions would decrease at the same rate. The data show, however, a greater proportion of buccolingual measurements rather than mesiodistal dimensions have significant differences. As such, PME can b e rejected as the mechanism of dental reduction, regardless of the observed differences in food production techniques. Increasing Population Density Effect Between the Pre Latte and Latte periods there was an increase in population density (Hunter Anderson and Butler, 1991) The IPDE model suggests that smaller body size, and as a result smaller tooth size, is concomitant with an increase in population size. Smaller body size reduce s environmental stress due to competition for nutritional resources (Machi arelli and Bondioli, 1986). A ll teeth and both dimensions should be affected equally. Specifically, in an island environment where space is limited, this seems like a plausible explanation. However, analysis of height between the Pre Latte and Latte peri ods shows little variation between the two, in both males and females. Further, the data show that each tooth type is affected differently with the posterior dentition reducing at a greater rate than the anterior dentition. Likewise, the mesiodistal and buccolingual dimensions are not affected equally. As such, dental reduction cannot be explained solely by smaller body size, as hypothesized in the IPDE model.

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104 Selective Compromise Effect Calcagno (1986; 1989) suggests that a highly cariogenic diet will lead towards selection for smaller dentition, which are more resistant to dental disease and crowding. The increased dependence on agriculture, particularly taro yams, and rice during the Latte Period would suggest a diet that is highly cariogenic. If SCE is at work, the mesiodistal and buccolingual diameters should be affected differently, as malocclusion would lead to selection for smaller mesiodistal diameters; whereas, selection for smaller buccolingual dimensions would occur with large amounts of c arious lesions (Greene, 1970; Sofaer, 1973; Christensen, 1998). Additionally, the posterior dentition would show greater amount of dental reduction to reduce occlusal complexity and prevalence for carious lesions. There is a significant difference in th e frequency of carious lesions between the Pre Latte and Latte with a drastic decrease in carious lesions in the later population. This finding could be the result of decreased size and complexity of the dentition as well as the culturally introduced prac tice of betel nut chewing which is known to have cariostatic properties. T he current study also demonstrates that the buccolingual and mesiodistal dimensions are not equally affected. A predilection for reduction of the buccolingual diameter is expected given the high number of carious lesions in the early population. Further, the anterior dentition shows a much smaller rate of dental reduction, which is primarily restricted to, and most prominent in the premolars and molars. Given these findings, the S CE is a likely model to explain the dental reduction observed in the prehistoric Chamorro.

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105 Masticatory Functional Hypothesis result of decreased functional demands placed on th e craniofacial apparatus, which in turn leads to a decrease in the masticatory complex (Carlson and Van Gerven, 1977; Calcagno, 1989). The archaeological data show significant changes in food processing techniques, as well as change in diet, associated wi th an increase in agricultural processes and cookery that result in softer diets. These cultural changes may have had such an impact that they led, indirectly, to biological changes, such as a decrease in the craniofacial complex, which subsequently affec ted the size of the dentition. Cranial and mandibular data show a decrease in size over time, particularly in the width dimensions. As was previously discussed, this finding is not associated with an overall decrease in body size, as stature remains fairly constant between the time per iods. Thus, it can be assumed that a decrease in the craniofacial complex is associated with reduced biomechanical demands due to dietary and food processing shifts. Therefore, the MFH model can also be utilized to explain the decrease in dental size ove r time in Guam. Conclusions This study analyzed diachronic trends in dental dimensions of the prehistoric Chamorro population on the island of Guam. An 8% reduction in tooth size is observed from the Pre Latte to Latte Periods. The results showed that s ignificant reduction occurred in 28 of the 64 analyzed dimensions. Reduction occurred most frequently in the buccolingual dimensions and at a greater rate in the posterior dentition. The maxilla and mandible were equally affected.

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106 These findings may be best explained by a combination of the Selective Compromise Effect and Masticatory Functional Hypothesis. An increased reliance on taro based agriculture would have led to a highly cariogenic diet. In this situation, small teeth would have been ideal, g reatly reducing the complexity of the occlusal surface and thus preventing formation of carious lesions. Additionally, increased food processing techniques, such as the use of mortar and pestles and cooking, minimize the force necessary to break down toug h food, which lead to decreased functional demands of the masticatory apparatus. These shifts occur over a relatively short period of time where dynamic transitions in cultural practices may have been the catalyst for biological change.

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107 Table 4 1. Tooth summary d ata Time Period Male Female Indet. Total Pre Latte 1470.9 1383.6 1463.9 1423.2 (n) (189) (299) (145) (633) Latte 1377.0 1250.2 1318.8 1307.1 (n) (177) (243) (189) (609) Table 4 2. Tooth summaries of prehistoric Chamorro p opulations from Guam Study Sample Subsample Time Period Male Female Total Sample References San Vitores Road Fujita Drainfield Pre Latte 1412 .0 1487 .0 Bath, 1986; Pietrusewsky, 1986 Naton Beach Pre Latte 1470.9 1383.6 1423.2 Current Study Naton Beach Latte 1377.0 1250.2 1307.1 Current Study San Vitores Road Right of Way Latte 1303 .0 1281 .0 1294 .0 Bath, 1986; Pietrusewsky, 1986 Apurguan N/A Latte 1259.5 1236.9 1247.9 Pietrusewsky, Douglas, Ikehara Quebral, 2003 Hyatt Hotel N/A Latte 1193.8 1138 .0 1161.7 Trembly, 1999 Leo Palace N/A Latte 1058.5 985.6 1034.5 Douglas and Ikehara, 1992

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108 Table 4 3. Tooth summaries of Pacific and circum Pacific s amples Study Sample Sample Tooth Summary References Guam, Fujita 1487 Pietrusewsky, 1986 Australia 1486 Brace, 1980 Tasmania 1429 Brace, 1980 Guam, Naton Beach (Pre Latte) 1423 Current Study Papua New Guinea, East Highlands 1395 Brace and Hinton, 1981 Tonga 1371 Brace and Hinton, 1981 Bougainville 1359 Brace and Hinton, 1981 Tumon Bay 1347 Pietrusewsky 1986a Northern Marianas 1341 Pietrusewsky and Batista, 1980 Fiji 1338 Brace and Hinton, 1981 New Britain 1334 Brace and Hinton, 1981 Papua New Guinea, Sepik River 1321 Brace and Hinton, 1981 Samoa 1311 Brace and Hinton, 1981 Guam, Hornbostle Thompson Collection 1309 Brace and Hinton, 1981 Guam, Naton Beach (Latte) 1307 Current Study Vanuatu 1295 Brace and Hinton, 1981 Guam, Right of Way 1294 Pietrusewsky 1986a Phillippines, Visayas 1288 Brace and Hinton, 1981 Pohnpei, Nan Mandol 1287 Pietrusewsky and Douglas, 1985 Papua New Guinea, North Coast 1286 Brace and Hinton, 1981 New Ireland 1266 Brace and Hinton, 1981 New Caledonia 1256 Brace and Hinton, 1981 Java 1240 Brace and Hinton, 1981 Northern Marianas 1238 Pietrusewsky 1986b Thailand 1233 Brace and Hinton, 1981 Marquesas 1204 Brace and Hinton, 1981 Hawaii 1200 Brace and Hinton, 1981 Japan 1200 Brace and Hinton, 1981 Borneo 1190 Brace and Hinton, 1981 Chatham Islands 1181 Brace and Hinton, 1981 China 1157 Brace and Hinton, 1981

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109 Table 4 4. Mean cranial and mandibular measurements associated with masticatory apparatus Pre Latte Latte Measurement Male (n) Female (n) Male (n) Female (n) Cranial length 187.3 (3) 176.3 (3) 183.5 (4) 179.0 (9) Cranial breadth 144.3 (3) 136.0 (3) 133.7 (3) 128.8 (10) Bizygomatic breadth 139.0 (1) 136.0 (1) 119.8 (4) Biauricular breadth 123.0 (2) 110.0 (4) 123.7 (3) 112.5 (4) Bimaxillary breadth 123.0 (1) 106.0 (1) 94.1 (4) Max alveolar breadth 59.0 (3) 56.6 (3) 57.5 (6) Max alveolar length 56.0 (3) 51.1 (2) Biogonial width 114.8 (4) 97.6 (3) 106.5 (15) 97.0 (13) Bicondylar breadth 136.0 (1) 122.5 (1) 126.1 (6) 113.9 (8) Mandibular length 79.6 (5) 76.5 (5) 79.2 (12) 75.4 (12) a. Data were compiled from unpublished measurements released to the author by Cherie Walth

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110 CHAPTER 5 DEVELOPMENTAL INSTABILITY: ENAMEL HYPOPLASIAS steady state of an organism. During periods of environmental stress, an organism undergoe s neural and endocrinal responses to activate homeostatic mechanisms to alleviate chronic stress events, which can leave lasting and permanent markers in the body, and specifically in the skeleton (Selye, 1973; Goodman et al., 1988; Goodman and Armelagos; 1989; Larsen, 1997; Goodman and Martin, 2002). Goodman and Armelagos (1984) present a stress model applicable to skeletal populations in which health is the fundamental variable in examining the adaptive processes of a population (Larsen, 1997; Goodman an d Martin, 2002). In this model, markers of skeletal stress develop when new cultural systems fail to buffer environmental stressors, such as disease or access to resources, therefore resulting in physiological disruption in the skeletal remains. Linear en amel hypoplasias (LEH) can be analyzed to estimate periods of non specific stress events in an individual or population. The current study investigates the use LEH as indicators of physiological disruptions and stress in two temporally disparate populatio ns from the Western Pacific island of Guam, in an attempt to understand the relationship between developmental instabilities and subsistence change in a population undergoing agricultural intensification. Enamel Hypoplasias Enamel hypoplasias are deficienc ies in enamel thickness that appear as horizontal linear grooves or pits and are the result of a disruption in amelogenesis (Pindborg,

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111 1970; Goodman and Armelagos, 1985). These defects are a permanent record of developmental disturbances that occurred dur ing prenatal and early childhood development due to physiological stress and are a direct response to an anomaly in matrix secretion that arrests ameloblastic activity (Pindborg, 1970; Duray, 1992). Enamel hypoplasias reflect a nonspecific physiological d isruption that occurred during growth and development of the tooth. Thus, disruption of enamel formation may be related to physiologic and metabolic stress where activation of the sympathetico adrenal medullary and pituitary adrenal cortical axes induces stress (Rose et al., 1985). These axes control output of hormones, which when elevated, will decrease the amount of protein made throughout the body and in turn interrupt the process of amelogenesis (Rose et al., 1985). The problem with understanding en amel defects lies not in how the developmental process of hypoplasias occurs, but in what external factors initiate the disruption (Goodman and Rose, 1990). Experimental studies in rats have reported hypoplasia formation as a result of exposure to various stressors such as malnutrition (Becks and Furata, 1941), infectious agents (Kreshover and Clough, 1953; Kreshover et al., 1953), and fever (Kreshover and Clough, 1953). These early studies solidified the current notion that linear enamel hypoplasias are the result of non specific physiological disturbances, which could be attributed to many different external stressors. As such, frequencies of enamel hypoplasias are often used to assess the relative health and nutritional status of prehistoric, historic, and contemporary human populations (Massler et al., 1941 ; Schiulli, 1978; Cook and Buikstra, 1979; Goodman et al., 1980; Cohen and Armelagos, 1984; Larsen, 1997; Zhou and Corruccini, 1998; Cucina, 2002; Pechenkina

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11 2 et al., 2002; Steckel and Rose, 2002; Hil lson, 2005; King et al., 2005; Buzon, 2006; Pechenkina and Delgado, 2006; Boldsen 2007; Berbesque and Doran, 2008), as well as in Neandertals (Ogilvie et al., 1989; Guatelli Steinberg et al., 2004), hominin ancestors (Skinner, 1996; Guatelli Steinberg, 200 3; Cunha et al., 2004), and non human primates (Guatelli Steinberg and Lukacs, 1999; Lukacs, 1999; Lukacs, 2001; Skinner and Hopwood, 2004). Health and Disease in the Shift to Agriculture The agricultural revolution has long been thought of as one of the major ad vances in human culture that le d to the rise of civilizations (Braidwood, 1960; Cohen, 1989). isease in prehistoric populations increased in the shift from hunter gathering to agriculture. An increase in infectious skeletal (yaws and tuberculosis) and dental diseases (carious lesions), non specific markers of stress (linear enamel hypoplasias) co u pled with decrease in stature and life expectancies were reported as evidence for declining changed the paradigm of health and agriculture, it was faulty in that it was skewed tow ards North American populations practicing the domestication and intensification of maize agriculture and it provided an overly simplified interpretation of the agricultural transition. Additionally, in the mid 1980 s, techniques and diagnoses had not yet been standardized (e.g. Buikstra and Ubelaker, 1994; Steckel and Rose, 2002). In the last two decades, vast improvements in methodologies have been made and research linking decreased health with dietary shifts and agricultural intensification has been c onducted on nearly every continent: North America (Larsen et al., 2002),

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113 South America (Ubelaker and Newson, 2002), Europe (Cucina, 2002), Africa (Keita and Boyce, 2001; Keita, 2003; Starling and Stock, 2007), Asia (Yammamoto, 1988; Lukacs, 1992; Lukacs an d Walimbe, 1998) Temple and Larsen, 2007; Temple, 2007) and Australia (Webb, 1984; Seow et al., 1991). These studies have shown that declining health cannot be explained by a single external stimulus and is multifactorial in nature. In populations underg oing a shift to agriculture, reduced health is related to a combination of increased population density and concomitant rise of infectious disease associated with sedentism and reduced access to nutritional requirements and food shortage due to drought or animal infestation related to the adoption of agriculture (Pietrusewsky and Douglas, 2002). However, the pattern of declining health with agricultural intensification is not universal may have no effect or may have an opposite trend of improved health (Hod ges, 1987; Neves and Wesolowski, 2002; Pietrusewsky and Douglas, 2002; Eshed et al., 2004; Douglas and Pietrusewsky, 2007). In some cases, p aleodemographic studies have sh own an increase in fertility after agricultural intensification, suggesting improved health status in the population (Bocquet Appel and Naji, 2006). For example, research from South East Asia does not appear to follow the trends observed in other parts of the world. A skeletal se ri es from northeastern Thailand spanning two millennia and including a period of agricultural intensification, shows mixed results (Pietrusewsky and Douglas, 2002; Douglas and Pietrusewsky, 2007). The earlier Non Nok Tha (5000 BP 1700 BP) displays a decr ease in linear enamel hypoplasias and cribra orbitalia and increase in stature despite the reliance on starchy foods (Douglas and Pietrusewsky, 2007); while the later Ban Chiang population (4100

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114 BP 1800 BP) has a decrease in carious lesions while LEH rem ain constant, demonstrating little evidence for health decline (Pietrusewsky and Douglas, 2002; Douglas and Pietrusewsky, 2007). Other skeletal assemblages in northeast Thailand demonstrate a similar trend. Domett and Tayles (2007) observe an improvement of health over time with an increase in height and similar rates of carious lesions in Bronze Age (Ban Lum Khao: 3400 2500 BP) and Iron Age (Noen U Loke: 2300 1600 BP) individuals The pattern of health and disease in Thailand is contrary to what is seen with the North American populations. Much of these results are attributed to the adoption of rice based agricultural practices instead of maize agriculture, which is prevalent in North America (Tayles et al., 2000). Further, rice, and other starchy crops, may have more nutritional value than other cultigens. Thus, many factors should be taken into account when attempting to interpret health differences in the shift to agricultural intensification. Environmental and genetic patterns may also come i nto play and affect the way cultural shifts in a population are expressed biologically. As such, varying regions should be analyzed independently to observe population specific trends that accompany intensification of agriculture. Current Study The shift between the Pre Latte ( 3 500 BP to 1000 CE ) and the Latte (1000 to 1521 CE ) time periods in Guam involves changes in population size, diet, and food procurement/preparation strategies (Hunter Anderson and Butler, 1991; Moore, 2005 ; Amesbury, 2007 ). The Pr e Latte population is a small semi nomadic foraging population that subsisted on bivalves, shellfish, and reef and pelagic fishing (Amesbery et al., 1991; McGovern and Wilson, 1996). Horticultural practices were likely brought to

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115 Guam by its earliest inh abitants from island Southeast Asia (Bellwood, 1978). Data from paleoenvironmental sediment cores suggest small scale gardening and resource collecting by initial settlers (Athens and Ward, 2004), and analyses of pollen and starch residues on the interior of Pre Latte pottery found evidence of taro, cabbage tree, fish, and shellfish (Loy, 2001a,b; Loy, 2002; Loy and Crowther, 2002). Charcoal becomes increasingly present in paleoenvironmental sediment cores around 1,800 BP, which corresponds to the Pre Lat te/Latte transition, and is suggestive of intensified land use (Athens and Ward, 2004). Latte period archaeology has identified a wide array of food processing artifacts, such as stone mortars and pounders, scrapers, knives, blades, and adzes, which are i ndicative of a more sedentary population with an increased reliance on agriculture (Hunter Anderson and Butler, 1991; Moore, 2005). Starch grain residues on Latte period potsherds identified cabbage tree, rice, sugarcane, and a taro ( Loy, 2001a,b; Loy, 20 02; Loy and Crowther, 2002). Taro was identified more than any other plant, suggesting a preference for this introduced tuber. Pollen and phytolith analyses have also found evidence of betel palm, breadfruit, coconut, bananas, and pandanus (Hunter Anders on and Butler, 1991; Cumming s and Puseman, 1998; Dixo n et al., 1999; Pearsall and Collins, 2000; Ward, 2000; Cummings, 2002; Athens and Ward, 2004). Stable isotope data reveal that the majority of the Latte diet was composed primarily of terrestrial C 3 resources, such as rice, root crops, and vegetables and that marine foods are merely a supplement, comprising approximately 30% of the diet (Hanson, 1991; Ambrose et al., 1997). Higher 13 N values are suggestive of preferential exploitation of reef and/ or lagoon fish over pelagic resources (Hanson,

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116 1991; Ambrose, 1997). Ethnohistoric and archaeological evidence points to exploitation of some larger pelagic fish such as tuna, dolphin and marlin (Driver 1993; Freycinet, 1996; Amesbury and Hunter Anderso n, 2008), however, deep sea fishing is more dangerous and thus not used as a reliable subsistence base (Russell, 1998). Thus, the favored reliance of marine based foods was abandoned for terrestrial resources and agricultural crops in the Latte period. T he stable isotope findings are further supported by archaeological units demonstrating a greater proportion of fishing gear and fishing related debris in the Pre Latte period in comparison with the Latte Period (Butler, 1995). Additionally, intensificatio n of agriculture, in the Latte period, would have permitted an expansion in population size. Population growth is evident with more varied habitation areas ranging from the preferred coastal locale to more marginal and inland environments, such as the int erior uplands (Hunter Anderson and Butler, 2001). Underwood (1973) and Hezel (1982) suggest that the prehistoric Latte population, at its largest, ranged between 30,000 to 40,000. The shift to agricultural intensification coupled with newly adapted food p rocessing tools is likely to be accompanied by biological changes, such as an increase in stress levels associated with population growth, limited access to resources, malnutrition, and increased prevalence of disease This study is restricted to the dent ition due to the poor quality of skeletal preservation that is typical in island and coastal environments such as Guam (Hanson and Butler, 1997) However, analyse s conducted by Cherie Walth over a year and a half period, will be considered to assist in th e construct ion a better health profile of the Chamorro population.

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117 The current study compares the frequency of linear enamel hypoplasias between the Pre Latte and Latte time periods in Guam. Decreased health, as evidenced by increased linear hypoplasias, is predicted concomitant with agricultural intensification, increased population size, and increase of infectious disease. The null hypothesis tested in this study is that there is no significant difference in the frequency of linear enamel hypoplasias between the Pre Latte and Latte time periods. Materials and Methods From 2006 to 2008, 376 prehistoric Chamorro burials were excavated during an archaeological mitigation project for the renovation of the Guam Aurora Villas & Spa in the Naton Beach Site on the northern end of Tumon Bay, Guam (DeFant, 2008) Of these, approximately 177 are associated with the Pre Latte Period and 190 belong to the Latte Period (DeFant, 2008) affiliation was based on the st ratigraphic location, associated artifacts, and radiocarbon dates (DeFant, 2008). Dating to roughly 2,500 BP, the Pre Latte Naton Beach sample represents some of the earliest settlers in Guam and is the largest Pre Latte mortuary sample discovered to date (DeFant, 2008). There are no radiocarbon dates for the Latte sample; however, archaeological materials associated the remains are consistent with those relegated to the Latte period, between AD 1000 to 1521. This study looked at the horizontal grooves or linear type hypoplasias, most commonly referred to in the literature as linear enamel hypoplasias (LEH) and 1982). The LEH were analyzed on the labial or buccal surfaces of the permanent dentition in individuals older than 10 years of age. Age estimates were determined skeletally using standards recommended by Buikstra and Ubelaker (1994) by Cherie

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118 Walth 1 Frequencies of LEH were examined in 197 individuals, 97 from the Pre Latte period and 100 from the Latte period. Distribution of sex is fairly even across the samples with females being represented more frequently than males in both the Pre Latte (38 and 33, respectively) and Latte (39 and 29, respectively) periods. L EH were scored macroscopically by both individual and tooth count methods, in all available and unmodified teeth, as suggested by Lukacs (1989). LEH were not sc ored in dentitions displaying moderate to extreme wear which was defined as teeth missing 75% or more of crown height Using the individual count method, LEH was scored as a discrete trait as present or absent. In the tooth count method, LEH were analyzed in each tooth individually and scored as an ordinal trait using the following scale: 0 = abs ence of LEH ; 1 = one hypoplastic defect observed in a single tooth; 2 = more than one LEH in a single tooth (Figures 5 1 & 5 2). The effects of labial abrasion, betel nut staining, and dental incising may have biased the analysis of LEH. The Pre Latte po pulation displays a unique pattern of dental modification that is not seen in the Latte period. Abrasion of the labial surface of adult maxillary teeth, from the central incisors to the fourth premolar, ranges from slight to extreme, and thus may oblitera te evidence of linear enamel hypoplasias (Figure 5 3). Additionally, central and lateral incisors are modified to a higher extent than the more posterior teeth (Parr and Walth, 2011), further skewing the data analysis since these teeth are also most likel y to display LEH (Goodman and Armelagos, 1985). The Latte dentition, on the other hand, displays dark reddish brown staining on the teeth due to 1 Data analysis on the cranial and postcranial skeletal remains of the Naton Beach sample was conducted by Cherie Walth. Her data, in conjunction with the dat a from the current study, were collected as part of a larger study on the Naton Beach Burial Complex, which is currently being compiled for submission to the Guam Historic Preservation Office.

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119 the cultural practice of betel nut chewing (Figure 5 4). This staining made LEH observation more difficult. Thus, LEH were scored as present if they were a palpable indention, through use of a fingernail, on the enamel surface. Intentional dental modification of the anterior dentition, in the form of incising cross hatched and oblique patterns on the enamel sur face, is also practiced in the Latte period (Ikehara Quebral and Douglas, 1997; Parr and Walt, 2012). Thus, LEH was not scored in teeth with modified surfaces. Frequency data by tooth and individual are reported. Multivariate statistical testing was utilized to determine if observed frequency differences were statistically significant. Differences between adult and subadult frequencies sex, and time period were tested using a Pearson Chi Square test Goodman and Armelagos (1985) demonstrated that the maxillary central incisors and mandibular canines are the most hypoplastic teeth in the dental arcade, as these teeth develop the earliest, tend to be more often affected. As such, this study compared the LEH frequencies of the maxillary central incis ors and mandibular canines, using a Pearson Chi Square test to see if significant differences exist between the time periods in these teeth. Results The mandibular canines and maxillary central incisors displayed the highest fre quencies of linear enamel h ypoplasias in both periods (Table 5 1 and Figure 6). In the Pre Latte, the mandibular canine has the highest frequency of occurrence ( n = 11; 29%), followed by the central maxillary incisor ( n = 5; 13%). Likewise, in the Latte period, LEH is highest in th e mandibular canine ( n = 27; 17%) and the maxillary central incisor ( n = 21; 14%).

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120 Age Differences Frequency differences of LEH in the juvenile and adult populations are apparent between the periods; however, caution must be taken due to the small sampl e size of the juvenile individuals, particularly in the Pre Latte population (n=5) (Table 5 2). In both periods, the juveniles have higher frequencies of hypoplastic teeth than the adults. In the Pre Latte, 40% ( n = 2) of the juveniles display LEH, while 15.2% ( n = 14) of the adults have LEH The LEH frequencies increase for both the juveniles ( n = 8; 61.5%) and adults ( n = 36; 41.4%) in the Latte period. The LEH frequencies were tested using the Pearson Chi Square test to determine if the observed diffe rences between subadults and adults w ere statistically significant; significant differences in age were found in both the Pre Latte or Latte time periods (Pre Latte: p = 0. 001 ; Latte: p = 0. 017 at the 0.05 alpha level) ( Table 5 6 ). Sex Differences Sex spe cific differences for LEH expression are noted in both the Pre Latte and the Latte periods, by individual and tooth count (Table 5 3 and 5 4). Even with the low frequencies of LEH in the Pre Latte, there are still major differences between male and female expression, where males exhibit LEH almost twice as frequently ( n = 8; 24.2%) as females ( n = 5; 13.2%). In the Latte period, the opposite trend is observed. Almost half ( n = 19; 48.7%) of the female population displays at least one or more LEH, while just over a quarter of the males exhibit LEH ( n = 8; 27.6%). Looking at tooth count incidence between the sexes, males from the Pre Latte period display LEH more frequently than fema les ( 14 /634 ; 2.2% and 13 /825 ; 1.6%; respectively); while in the Latte period, the female dentition has a slightly higher incidence of LEH ( 54 /670 ; 8.1%) when compared to males ( 33 /450 ; 7.3%).

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121 Sex differences (with and without indeterminate sex category inc lusion) were tested to determine significance of linear enamel hypoplasia expression both between and within each time period. Inclusion of indeterminate sex category did not affect the outcome, however the results discussed are based on tests on only mal es versus females. A Pearson Chi Square test found no significance sex differences between male and female LEH occurrence in either the Pre Latte (p = 0. 228 ) or Latte (p = 0. 078 ) time periods, at a 0.05 alpha level ( Table 5 7 ). Time Period Differences Lin ear e namel hypoplasia frequencies are discussed both in terms of overall individual frequency per time period (Table 5 3) as well as tooth count frequency (Table 5 4). Discrepancies are seen with expression of LEH between the time periods. The Pre Latte individuals have a low frequency of LEH, with only 16.5% (n = 16) affected (Table 5 3), while LEH frequencies are much higher in the Latte where nearly half of the population, 45.0% (n = 45) displays at least one or more LEH (Table 5 3). Likewise, in ter ms of tooth count, the Latte dentition (155/1693; 9.2%) is much more highly affected by LEH than the Pre Latte dentition (38/1925; 2%) (Table 5 4). A Pearson Chi Square test was performed to determine if the observed frequency differences of LEH expression between time periods were statistically significant ( Table 5 8 ). The Pearson Chi Square test demonstrates that there is a significant differ ence, at a 0.05 alpha level (p < 0.001 ), in expression of linear enamel hypoplasias between the Pre Latte and Latte time periods. Goodman and Armelagos (1985) showed that the maxillary central incisors and mandibular canines display the highest frequencies of linear enamel hypoplasias. Thus, these teeth were tested independently across time periods with a Pearson Chi Square

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122 test Results indicate significant differences in LEH expression in the maxillary central incisors and mandibular canines between the time periods at a 0.05 alpha level: left I 1 : p = 0.008 ; right I 1 : p = 0.031 ; left C 1 : p = 0.0 1 2; right C 1 : p = 0.038 ( Table 5 8 ). Discussion The current study found that the maxillary central incisor and the mandibular canines display the highest frequency of LEH. These findings are consistent with those reported in other studies (Goo dman and Armelagos, 1985; Hillier and Craig, 1992; Stodder, 1997). Age Differences No significant differences in subadult and adult expression of linear enamel hypoplasias were found between the time periods; however, when looking at the raw frequency dat a both the Pre Latte and Latte individuals displaying increase of LEH with age Both groups display similar rates of LEH increase, with subadults increasing by 22% and adults increasing by 26% between the Pre Latte and Latte periods. These f indings sugge st that there is an increase in LEH expression from the Pre Latte to the Latte periods in both subadults and adults and that the Latte population was exposed to significantly higher amounts of physiologi cal stress than the Pre Latte. T he raw frequency dat a show that juveniles have a higher frequency of hypoplastic teeth than the adults in each time period. In populations with abrasive diets this finding could be attributed to high rates of attritional wear, however, skeletal populations in the Marianas display minimal wear until approximately age 40 (Leigh, 1929; Stodder, 1993). Further, LEH data was not co llected on individuals with moderate to extreme amounts of wear. The disparate hypoplastic rates between juveniles and adults suggest that individuals surviving to adulthood may have been healthier and less susceptible to physiological

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123 stress compared to the greater frequency of individuals with LEH who died as subadults. While not statistically significant, this finding is similar to others who found significant differences between subadult and adult LEH frequencies in Guam (Stodder, 1997 ), Croatia ( Slau s, 2000) and North America (Cook and Buikstra, 1979; Duray, 1996) and may indicate a correlation between linear enamel hypoplasias and life expectancy (Slaus, 2000). Sex Differences As was noted in other studies in the Western Hemisphere ( Lanphear, 1990; Duray, 1996; Malville, 1997; Berbesque and Doran, 2008) as well as within Guam (Douglas et al., 1997; Pietrusewsky, 1997) sex is not found to be a statistically significant factor. However, Guatelli Steinberg and Lukacs (1999) found that expression of LEH is highly variable between the sexes. Female frequencies, by individual count (13%), are fairly low in the Pre Latte and increase dramatically to 49% in the Latte, whereas male LEH frequencies increase by a mere 5%. Sex specific differences were also no ted in LEH occurrence, by tooth count, in the Leo Palace and Hyatt Latte dentition, also located in Tumon Bay (Douglas and Ikehara, 1992; Trembly, 1999 ). The largest discrepancy is seen in the Leo Palace sample where the 38% of the male dentition display LEH, while only 5% of the female dentition displays LEH. Likewise, in the Hyatt sample, the males (10.2%) display LEH almost twice as frequently as the female dentition (6.2%). However, the findings of the Leo Palace (Douglas and Ikehara, 1992) and Hyatt (Trembly, 1999) sites are more in line with the Pre Latte sample of the current study, with females having smaller rates of LEH than men. In the Apurguan study, however, males have smaller LEH frequencies than females (Pietrusewsky et al., 1997;

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124 Pietruse wsky et al., 2003) and do not explain the increase of female LEH in the Latte population of the current study. The greater frequency of LEH in the femal e Latte population s is in contrast to the findings from the Leo Palace and Hyatt hotel sites which show more developmental disturbances in men in comparison to women (Douglas and Ikehara, 1992; Trembly, 1999) While, the sex differences in the Naton Beach sample were statistically significant, the drastic switch in female susceptibility to physiologica l stress between the Studies analyzing divi sion of labor in hunter gather ing and farming populations have shown significant dietary differences between the sexes (Hill a nd Hurtado, 198 9; Walker and Hewlett, 1990). Pre Latte w omen may have had better access to nutritionally rich resources. This trend appears to have shifted in the Latte period, as the LEH frequencies in women greatly exceed those of men. Thus, men in th e Latte likely had better access to resource thus making them less susceptible to physiological stress. Time Period Differences Overall, the average of both the Pre Latte and Latte tooth count frequencies in the current study (1.9% and 7.7%, respectively ) are much lower than all other LEH tooth count frequencies reported for Guam, which range from 13% in the Hyatt sample ( Trembly, 1999) to 22% in the Leo Palace sample (Douglas and Ikehara, 1992). When looking solely at the incisors and canines, an even h igher frequency of LEH is observed in Guam. Pietrusewsky and colleagues (1997) surveyed LEH expression in incisors and canines from six prehistoric Chamorro populations in Guam and report a combined tooth count frequency of 31%, however, this number combi nes both Pre Latte and Latte

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125 inclusion of all observable teeth in the dental arcade, including third molars. Additionally, the Pre Latte cultural practice of labial abras ion (Parr and Walth, 2011), which is most prominent in the anterior dentition (Figure 5 3 ), precludes examination of LEH in a large number of Pre Latte individuals. The current study shows an increase of LEH expression over time from 17% to 45% between the Pre Latte and Latte individuals and is supported by Pearson Chi Square test which demonstrates statistically significant differences in LEH expression between the time periods. This finding suggests that the Latte population were more susceptible to ext ernal stressors than the Pre Latte. Analysis of infectious disease demonstrates that the Latte population also had a greater degree of infectious disease (Walth, pers. comm.) Hanson and Butler (1977) report that treponemal infection and non specific pe riostitis are the most common infectious diseases in the Marianas Islands during the Latte period. In the current sample, evidence of periostitis, endemic yaws, and leprosy were observed, the majority of which occurred almost entirely in the Latte populat ion (Walth, pers. comm.). Non specific periostitis is the most commo n infectious disease occurring in primarily the Latte sample and s even Latte individuals displayed periosteal inflammation and infection indicativ e of yaws (Walth, pers. comm.). Yaws was also found in 19% of the adjacent Gongna Gun Beach population (Rothschild and Heathcote, 1993) and 20% of the Hyatt Hotel individuals (Trembly, 1999), both relegated to the Latte period and also located Tumon Bay. The Apurguan sample of Agana Bay, jus t south of Tumon Bay, has a much lower incidence, 9%, of treponemal infection (Pietrusewsky et al., 1997). A survey of remains from Guam, Saipan, and

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126 Tinan, housed at the Bishop Museum in Honoulu, reported very high rates of treponemal infection, 77%, (Su zuki, 1986). However, these remains were collected in the early 1900 s, are not well provenienced, and may have been retained specifically for interesting anatomical or pathological variants. None of the individuals from the Pre Latte sample were observed with yaws like infections (Walth, pers. comm.). Leprosy was observed at a lesser extent in both periods, with only one individual from the Pre Latte displaying lesions that may have been due to Mycobacterium leprae infection and two individuals from the Latte display lesions characteristic of leprosy (Walth, pers. comm.). These findings suggest that the Pre Latte individuals were much healthier overall with decreased rates of infectious disease as well as linear enamel hypoplasias, than their Latte count erparts. Carious lesions, on the other hand, reveal an opposite trend and are more prominent in the Pre Latte period and decrease in frequency in the Latte period. This finding is opposite of what had been expected and is likely due to the cultural practi ce of betel nut chewing of the Latte adults, which has cariostatic properties. Additionally, the Latte were dependent on other starchy crops, such as rice, taro, and yams, which may not be as cariogenic as maize (see Chapter 6 for more detailed discussion ). The current study found a significant increase of LEH frequencies over time, thus, the null hypothesis of no significant difference in the frequency of linear enamel hypoplasias between the two time periods can be rejected. It can be inferred that an increase of LEH over time is correlated with the intensification of agriculture concomitant with reduced nutritional values, population increase, and subsequent increase in infectious disease. These findings are consistent with studies, showing a

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127 positiv e correlation between an increase of hypoplastic defects with a shift in subsistence economies and agricultural intensification. This trend is best illustrated in the Dickinson Mo unds burial complex of Illinois where Goodman and colleagues (1980; 1984) do cumented an increase in LEH from the semi sedentary Late Woodland hunter gatherers (45%), to the Mississippian Acculturated Late Woodland transitional population (60%), and Middle Mississippian population (80%) with a large sedentary population reliant on intensified maize agriculture. Changing rates of LEH over time associated with dietary transitions were also found in North America with the adoption of a primarily maize subsistence economy (Sciulli, 1977; Sciulli, 1978; Cook, 1984; Larsen, 1995 ); as wel l as in the late Paleolithic to Mesolithic shift to early agriculture in the Levant (Smith et al., 1984); intensified agriculture in Ecuador (Ubelaker, 1984); improved food processing technologies associated with intensified agriculture in India (Lukacs, 1 992); shift to protein deficient and carbohydrate rich agricultural diet in the Early Bronze Age of Italy (Cucina, 2002); transition to early agriculture in the Nile Valley (Starling and Stock, 2007); and subsistence shifts directed by environmental change in Jomon foragers (Temple, 2007). Broader Implications An increased prevalence of linear enamel hypoplasias between the Pre Latte and Latte periods suggest a period of higher levels of physiological stress in the Latte period. Large scale environmental oscillations occurred during the Pre Latte/Latte transition which corresponds to the shift from the Medieval Warm Period (AD 800 to AD where rapid cooling temperatures, de cline in sea levels, and increased storminess lead to greater climatic variability (Bridgman, 1983; Hanson, 1991; Nunn and Britton, 2001;

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128 Nunn, 2007; Nunn et al., 2007; Hunter Anderson, 2010). During the Little Ice Age, rainfall in the Western Pacific was more erratic with more frequent and prolonged would have been associated with socie tal disruption, subsistence change, and move ment to inland habitation areas throughout the Pacific Basin (Nunn, 2000; Nunn and Britton, 2001; Nunn, 2007). Archaeological studies have demonstrated the predicted subsistence change with differential exploitation of marine resources and increased reliance on agriculture. A shift from bivalve to gastropod consumption, coupled with a shift from pelagic to reef/l agoon fish exploitation accompanied the Pre Latte/Latte transition as a result of ecosystem and sea level changes (Leidemann, 1980; Graves and Moore, 1985; Hanson, 1991; Amesbury et al., 1996; Ambrose, 1993; Ambrose et al 1997; Amesbury, 2007). Addition ally, reduction of fishing gear (Butler, 1995) and increased diversification of artifacts associated with farming (Moore, 2005) between the Pre Latte and Latte period indicates that there was a decreased emphasis on fishing and an increased reliance on agr icultural crops. Further evidence of agricultural intensification include technological advances in food preparation techniques with the advent of the lusong the Chamorro stone mortar, used to process rice, taro, and yams ( Moore, 2005), and production of thickened pottery for cooking (Loy, 2002; Moore, 2002). The archaeological data is supported by stable isotope analysis, which demonstrates that marine resources account for only 30% of the Latte diet with most of the sustenance derived from terrestrial resources, such as rice, root crops, and vegetables (Hanson,

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129 1991; Ambrose et al., 1997), adding further evidence of the importance of agricultural crops as a mainstay in the Latte period diet. Thus, the cultural and dietary shifts between the Pre Latte t o Latte period were likely brought on by climatic variation during the AD 1300 Event, which resulted in a transition from mixed horticultural subsistence to an increased reliance on agriculture and subsequent a population growth, as well as a decreased rel iance on marine resources, specifically pelagic fish. The above evidence points to a greater prevalence of developmental instabilities and infectious disease in the Latte period coupled with climate change and agricultural intensification, and subsequent malnutrition. Similar studies in the prehistoric North American Southwest (Stodder et al., 2002) and Japan (Temple, 2007), as well as in mid 1900 s China (Zhou and Corruccini, 1998), have also shown a correlation between seasonal resource depletion and sys (2007) of carious lesions in the Middle to Late Jomon time periods of Japan suggests: The presence of a dietary shift after a significant climate change follow the model of culturally induced stress of Goodman and Armelagos (1989), where behavioral decision in response to environmental constraint often carry biological consequences (p. 1043). The current study predicts a similar scenario in Guam, where environmental constraints due to climatic variability and i nstability, lead to dietary transitions and thus greater levels of stress as evidenced by increased linear enamel hypoplasias over time. Conclusions There were no significant differences in the rates of juvenile and adult LEH ; however, in terms of raw freq uency data, juveniles displayed higher hypoplastic defects than adults. This may suggest a correlation between life expectancy and physiological disruptions. Male and female differences in LEH expression were not significant within

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130 each time period, but looking at the raw frequency data provides interesting insight into differential access to resources. In the Pre Latte sample the females were exposed to less physiological stressors and as such, may have had better access to nutrition rich foods compared to the men. However, this trend shifted significantly in the Latte where women became much more highly susceptible to physiological stressors than men. Differences in gender status does not explain the gap between male and female LEH rates, as the Latte peoples followed matrilineal kinship system where the women were powerful and respected in all aspects of society (Souder, 1992). Thus, division of labor, with women collecting and gathering versus males hunting or fishing may be the r eason for sex discrepancies in frequencies of physiological stress. The Pre Latte peoples were horticulturalists and supplemented food with marine resources. Women may have gathered food as well as collected bivalves for consumption along the coast, whil e the men explored beyond the reefs to collect the larger pelagic fish. With the intensification of agriculture in the Latte period, more time would have been spent tending to and cultivating crops. Ethnohistoric accounts suggest that job was delegated t o women (Driver, 1993), who likely had less time to collect coastal marine foods. Men are reported as being the primary fishermen (Driver, 1993; Russell, 1998) who abandoned dangerous exploitation of the pelagic fish for the more easily obtainable reef fi sh and gastropods. In this scenario, men would have had greater access to protein content than women, and making them less susceptible to physiological disruptions. The Pre Latte people, overall, appear to have been healthier than the Latte individuals wi th less frequency of LEH and infectious disease. The increased

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131 susceptibility of the Latte population to linear enamel hypoplasias is associated with seasonal variability, such as typhoons, droughts, and increased aridity, which would have damaged crops a nd surrounding reef systems resulting in a reduction of agricultural productivity, depletion and /or destruction of staple foods (Stodder, 1997; Hunter Anderson, 2010), and spread of endemic disease ( Pietrusewsky et al., 1997; Stodder, 1997) Such climati c variability and decreased access to nutritional resources would have been detrimental to the overall health of the Latte peoples especially in developing children when the physiological disturbances that affect enamel formation are taking place The current study provides a diachronic analysis of the Chamorro health profile. Previous studies have described the Latte people as having higher susceptibility to linear enamel hypoplasias compared to Hawaiians as a result of nutritional deficiencies and infectious disease (Pietrusewsky et al., 1997). This study validates that claim and expands on the current knowledge of prehistoric health in the Chamorro by demonstrating that the earlier Pre Latte inhabitants were healthier than the later Latte population as a result of climatic instability and subsequent dietary transitions.

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132 Table 5 1. Percentage of teeth with one or more linear enamel hypoplasia a Pre Latte Latte n % n % Maxillary I1 5 13.2 21 13.5 I2 3 7.9 9 5.8 C 3 7.9 13 8.4 P3 2 5.3 6 3.9 P4 2 5.3 4 2.6 M1 0 0.0 6 3.9 M2 0 0.0 11 7.1 M3 1 2.6 2 1.3 Mandibular I1 2 5.3 11 7.1 I2 3 7.9 15 9.7 C 11 28.9 27 17.4 P3 1 2.6 12 7.7 P4 3 7.9 9 5.8 M1 2 5.3 4 2.6 M2 0 0.0 5 3.2 M3 0 0.0 0 0.0 a. Left and Right Sides Combined Table 5 2. Individual occurrence of Pre Latte and Latte linear enamel hypoplasias by age grouping Pre Latte Latte n/N % n/N % Juvenile 2/5 40.0 8/13 61.5 Adult 14/92 15.2 36/87 41.4 Total 16/97 16.5 44/100 44.0 n = number of individuals with linear enamel hypoplasias N = number of individuals examined Table 5 3. Individual occurrence of linear enamel hypoplasias Pre Latte Latte n/N % n/N % Males 8/33 24.2 8/29 27.6 Females 5/38 13.2 19/39 48.7 Indeterminate 3/26 11.5 18/32 56.3 Total 16/97 16.5 45/100 45.0 n = number of individuals with at least one linear enamel hypoplasia present N = number of individuals examined

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133 Table 5 4. Tooth count of linear enamel hypoplasias Pre Latte Latte n/N % n/N % Males 14/634 2.2 33/450 7.3 Females 13/825 1.6 54/670 8.1 Indeterminate 11/466 4.1 68/573 11.9 Total 38/1925 2.0 155/1693 9.2 n = number of teeth with at least one linear enamel hypoplasia N = number of teeth examined

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134 Table 5 5. LEH frequencies in comparative populations Tooth Count Individual Count Study Sample Subsample Time Period Male % Female % Male % Female % References Naton Beach Pre Latte Pre Latte 2.2 1.6 23.5 13.2 Current Study San Vitores Road Fujita Drainfield Pre Latte 19.5 Bath, 1986; Pietrusewsky, 1986 Naton Beach Latte Latte 7.3 8.1 28.6 48.7 Current Study Apurguan N/A Latte 20.6 25.1 Douglas et al., 1997; Pietrusewsky et al., 2003 Hyatt Hotel N/A Latte 10.2 6.2 Trembly, 19 99 Leo Palace N/A Latte 38.0 5.0 Douglas and Ikehara, 1992 Fiesta Resort N/A Latte 11.8 Defant et al 2008

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135 Table 5 6. Age differences of LEH using a Pearson Chi Square Test Value Df Asymp. Sig (2 sided) Age within Pre Latte 15.547 3 0.001* Age within Latte 12.089 4 0.017* Indicates statistically significant differences at the 0.05 alpha level Table 5 7. Sex differences of LEH u sing a Pearson Chi Square Test Value Df Asymp. Sig (2 sided) Exact Sig. (2 sided) Exact Sig. (1 sided) Age within Pre Latte 1.451 1 0.228 0.357 0.185 Age within Latte 3.102 1 0.078 0.087 0.065 a. Indeterminate sex not included + Indicates no statistical relationship at the 0.05 alpha level T able 5 8. Pre Latte and Latte d ifferences in LEH using a Pearson Chi Square Test Value Df Asymp. Sig (2 sided) Exact Sig. (2 sided) Exact Sig. (1 sided) Between Time Periods 18.716 1 0.000 0.000 0.000 LMAX I1 9.785 2 0.008 RMAX I1 8.889 3 0.031 LMAND C 10.894 3 0.012 RMAND C 8.444 3 0.038 Indicates statistically significant differences at the 0.05 alpha level

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136 Figure 5 1. Single linear enamel hypoplasia in the mandibular lateral incisor, canine, and third premolar (Photo by author) Figure 5 2. Multiple linear enamel hypoplasias in a single tooth (Photo by author)

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137 Figure 5 3. Labial abrasion in a Pre Latte Period individual (Photo by author) Figure 5 4. Betel nut staining in a Latte Period individual (Photo by author)

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138 Figure 5 5 Dental incising in a Latte Period Individual (Photo by author) Figure 5 6. Frequency of linear enamel hypoplasias by tooth type (right and left sides combined) 0 5 10 15 20 25 30 I1 I2 C P3 P4 M1 M2 M3 I1 I2 C P3 P4 M1 M2 M3 Maxillary 38 38 38 38 38 38 Mandibular 38 38 38 38 38 38 38 LEH Frequency Tooth Type Pre-Latte Latte

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139 CHAPTER 6 CARIOUS LESIONS Patterns of dental disease have long been studied by anthropologists interested in the health, diet, and lifestyle of prehistoric populations. Aristotle was the first to record the correlation between diet and dental disease in antiquity when he observed that sweet figs adhering to the teeth caused dental caries (Powell, 1985). Carious lesions, in particular, have been of interest because they are a direct indication of an infectious disease process and are easily observable in the dentition, which is mor e likely to survive in an archaeological setting than other indicators of skeletal infection. As such, carious lesions are the most common dental disease (Bunting, 1933) and are the most frequently reported dental pathology found in archaeological populat ions (Roberts and Manchester, 1995). The current study examines carious lesion rates between the horticultural Pre Latte ( 3500 BP to 1000 CE ) and early agricultural Latte ( 1000 to 1521 CE ) populations in Guam to investigate what effect intensified agricu lture and increased reliance on starchy foods such as rice, taro, and yams have on the oral health status of the population. The excavation of the Naton Beach Burial Complex allows for the first diachronic assessment health in prehistoric Chamorro populat ion using large sample sizes. Decreased health, as evidenced by increased carious lesions, is predicted concomitant with agricultural intensification. This study tests the null hypothesis that there is no significant difference in the frequency of cariou s lesions between the Pre Latte and Latte time periods.

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140 Formation of Carious Lesions Dental caries can take on many different forms such as pit and fissure caries, smooth surface caries, root caries, and deep dentin caries (Newbrun, 1982). Larsen (1997:65 demineralization of dental hard tissues by organic acids produced by bacterial of particular importance as it varies based on amounts of protein and carbohydrates in the diet. Lactic acid is produced when plaque bacteria metabolize carbohydrates, while alkaline waste products are produced by metabolization of protein. Thus, the formation of a ca rious lesion occurs when periods of acidity outweigh periods of alkalinity and mineral destruction of the enamel occurs (Hillson, 1979). The epidemiological literature has shown that sugar is one of the major factors of carious lesions (Stoppelaar et al., 1970; Newbrun, 1982) and that foods with high amounts of dietary sucrose, which correlate to acidic periods in plaque, are more likely to result in carious lesion formation (Hillson, 1979; Larsen, 1983 ). Carious Lesions and Agricultural Intensification Carious lesion etiology is not fully understood, however, many factors have been associated with their development, including genetic predisposition, salivary flow and chemistry, tooth size and morphology, diet, fluoride component in drinking water (Newbru n, 1982; Rowe, 1982), as well as the physical properties of food and the way it is prepared (Larsen, 1997). Turner (1979) reports carious lesion frequencies drawn from populations with different subsistence patterns from around the world, and suggests that caries prevalence increase from hunter gatherer societies to agricultural ones. He further

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141 proposes frequency ranges, in terms of tooth count frequencies, for different subsistence patterns: hunter gatherers: 0.0% to 5.3%, mixed economies: 0.44% to 10.3% and agricultural: 2.3% to 26.9%. These values are often used in studies to assist in reconstructing dietary subsistence patterns (e.g. Delgado Darias et al., 2005; Paleopathology at the O rigins of Agriculture lesions increase in frequency in the shift from hunter gatherer populations to agricultural ones. An increase in carious lesion prevalence and agricultural intensification has been repeatedly demonstrated world wide, with various food crops such as maize in North America (Larsen, 1981; Cook, 1984; Larsen, 1984; Larsen et al.,1991) and Ecuador (Ubelaker, 1980; Ubelaker, 1984; Ubelaker and Newson, 2002); wheat and barley in Egypt (Hillson, 1979), the Levant (Smith et al., 1984), and Pakistan (Lukacs, 1992); millet in North Africa (Martin et al., 1984) and Northern China (Pechenkina et al., 2002); and rice in Southeast Asia (Krigbaum, 2007) and Japan (Temple, 2007; Temple and Larse n, 2007). However, some recent studies have shown no significant relationship between carious lesion frequency and intensification of agriculture (Oxenham et al., 2000; Tayles et al., 2000; Pietrusewsky and Douglas, 2002; Eshad et al., 2006; Domett and Tay les, 2007; Douglas and Pietrusewsky, 2007; Lanfranco and Eggers, 2010). Specifically, trends in Southeast Asia show either homogeneity in carious lesion rate or decline in caries associated with the intensification of agriculture and reveal little evidenc e for decreased health status over time (Oxenham et al., 2000; Tayles et al., 2000; Pietrusewsky and Douglas, 2002; Oxenham et al., 2006; Douglas and Pietrusewsky,

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142 2007; Domett and Tayles, 2007). This geographical trend has been attributed to the transiti on to rice (1984) volume. However, other studies have found an increase of carious lesions with agricultural intensification of rice in present day Malaysia (Krigbaum, 2007) and Japan (Temp le and Larsen, 2007). Temple and Larsen (2007) suggest, in support of their findings, that rice, while not as cariogenic as other starches, has more cariogenic properties than originally proposed by Tayles and colleagues (2000). In a follow up article, T ayles and colle a gues (2009:163) suggest that the variability in rice cariogenicity groups reliant on rice may be due to minimal processing of the starchy crop (Talyes et al., 2009). The above studies depict the difficulty of interpreting carious lesions within the realm of dietary transitions, specifically in regards to rice agricultu re. Thus, the development of carious lesions within an individual or population must be understood as a process that is multifactorial in nature and cannot be attributed to a single variable. As such, analysis of carious lesions must be undertaken with a careful examination into diet, food processing, and other cultural factors that may be in effect. Culture History of Guam The shift between the Pre Latte ( 3500 BP to 1000 CE ) and the Latte (1000 to 1521 CE ) time periods in Guam involves changes in populat ion size, diet, and food procurement/preparation strategies ( Hunter Anderson and Butler, 1991 ; Moore, 2005 ; Amesbury, 2007 ). The Pre Latte population is a small semi nomadic foraging population that subsisted on bivalves, shellfish, and reef and pelagic fishing (Amesbery

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143 et al., 1991; McGovern and Wilson, 1996). Horticultural practices were likely brought to Guam by its earliest inhabitants from island Southeast As ia (Bellwood, 1978). Data from paleoenvironmental sediment cores suggest small scale garde ning and resource collecting by initial settlers (Athens and Ward, 2004), and analyses of pollen and starch residues on the interior of Pre Latte pottery found evidence of taro, cabbage tree, fish, and shellfish (Loy, 2001a,b; Loy, 2002; Loy and Crowther, 2002). Charcoal becomes increasingly present in paleoenvironmental sediment cores around 1,800 BP, which corresponds to the Pre Latte/Latte transition, and is suggestive of intensified land use (Athens and Ward, 2004). Latte period archaeology has identi fied a wide array of food processing artifacts, such as stone mortars and pounders, scrapers, knives, blades, and adzes, which are indicative of a more sedentary population with an increased reliance on agriculture ( Hunter Anderson and Butler, 1991 ; Moore, 2005). Starch grain residues on Latte period potsherds identified cabbage tree, rice, sugarcane, and a taro ( Loy, 2001a,b; Loy, 2002; Loy and Crowther, 2002). Taro was identified more than any other plant, suggesting a preference for this introduced tub er. Pollen and phytolith analyses have also found evidence of betel palm, breadfruit, coconut, bananas, and pandanus ( Hunter Anderson and Butler, 1991 ; Cummings and Puseman, 1998; Dixo n et al., 1999; Pearsall and Collins, 2000; Ward, 2000; Cummings, 2002; Athens and Ward, 2004). Stable isotope data reveal that the majority of the Latte diet was composed primarily of terrestrial C 3 resources, such as rice, root crops, and vegetables and that marine foods are merely a supplement, comprising approximately 30% of the diet (Hanson, 1991; Ambrose et al., 1997). Higher 13 N values are suggestive of

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144 preferential exploitation of reef and/or lagoon fish over pelagic resources (Hanson, 1991; Ambrose, 1997). Ethnohistoric and archaeological evidence points to exploi tation of some larger pelagic fish such as tuna, dolphin and marlin (Driver 1993; Freycinet, 1996; Amesbury and Hunter Anderson, 2008), however, deep sea fishing is more dangerous and thus not used as a reliable subsistence base (Russell, 1998). Thus, t he favored reliance of marine based foods was abandoned for terrestrial resources and agricultural crops in the Latte period. The stable isotope findings are further supported by archaeological units demonstrating a greater proportion of fishing gear and fishing related debris in the Pre Latte period in comparison with the Latte Period (Butler, 1995). Additionally, intensification of agriculture, in the Latte period, would have permitted an expansion in population size. Population growth is evident with more varied habitation areas ranging from the preferred coastal locale to more marginal and inland environments, such as the interior uplands (Hunter Anderson and Butler, 2001). Underwood (1973) and Hezel (1982) suggest that the prehistoric Latte populati on, at its largest, ranged between 30,000 and 40,000. The cultural modifications associated with intensification of agriculture, such as food processing, are likely to be accompanied by biological changes, such as an increase in stress levels associated with population growth, limited access to resources, malnutrition, and increased prevalence of disease. This study is restricted to the dentition due to the poor quality of skeletal preservation that is typical in island and coastal environments such as Guam (Hanson and Butler, 1997). However, analyses conducted by Cherie Walth, will be utilized to augment data from the current study to construct a health profile of the Chamorro population.

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145 Materials and Methods From 2006 to 2008, 376 prehistoric Chamorr o burials were excavated during an archaeological mitigation project for the renovation of the Guam Aurora Villas & Spa in the Naton Beach Site on the northern end of Tumon Bay, Guam (DeFant, 2008). Of these, approximately 177 are associated with the Pre Latte Period and 190 belong to on the stratigraphic location, associated artifacts, and radiocarbon dates (DeFant, 2008). Dating to roughly 2,500 BP, the Naton Beac h Pre Latte sample represents some of the earliest settlers in Guam and is the largest Pre Latte mortuary sample discovered to date (DeFant, 2008). There are no radiocarbon dates for the Latte sample; however, archaeological materials associated the remai ns are consistent with those relegated to the Latte period, between AD 1000 to 1521. Carious lesions were analyzed macroscopically on all fully formed permanent teeth, by individual and by tooth count methods, as recommended by Lukacs (1989). LEH by i ndi vidual count was scored as present or absent of carious lesions. Carious lesions, by tooth count, were scored using an ordinal scale: 0 = absence of carious lesions ; 1 = presence of carious lesions in a single tooth ; 2 = presence of one or more caries in a single tooth The Pre Latte sample consists of 99 individuals, while 108 individuals from the Latte period were analyzed (Table 6 1). A tot al of 3,666 teeth were analyzed: 1,930 from the Pre Latte and 1,736 from the Latte (Table 6 2). Females 1 are repr esented 1 Demographic data, as well as postcranial pathological conditi ons, were recorded by Cherie Walth. Her data, in conjunction with the data from the current study, were collected as part of a larger study on the Naton Beach Burial Complex, which is currently being compiled for submission to the Guam Historic Preservati on Office.

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146 more o ften than males in both samples; however, sex is distributed fairly evenly in the Pre Latte sample (females: n = 38; males: n = 33). The distribution within the Latte sample is more heterogeneous (females: n = 37; males: n = 29). Age groups were also analyzed separately, however, the children and juvenile samples are very small, particularly in the Pre Latte period (Table 6 3). Raw frequency data is reported by time period, sex, and age. Inflated carious rates are often reported in the bio archaeological literature when tooth class is not taken into account, as the molars, followed by the premolars, are more susceptible to caries and are also the best preserved in archaeological conditions and less vulnerable to postmortem loss (Hillson, 199 6). Therefore carious lesions were also analyzed in terms of tooth position. Multivariate statistical testing was performed to determine statistical significance of differences in carious lesion frequency between groups. A Pearson Chi Square test was used to test for significant differenc es between the time periods, the sexes, and the age groups. Results Tooth Position As expected, the molars have a greater frequency of carious lesions than rest of the dentition, with the mandibular second molar displaying the highest frequency of carious lesions in both groups (Pre Latte: n = 37; 29.6%, Latte: n = 14; 10.2%) (Table 6 4). The mandibular incisors and canines of the Pre Latte had higher than expected carious lesions prevalence (12% or greater), esp ecially when compared to the Latte, which had relatively no carious lesions in the mandibular incisors (Mand I1: n = 0; 0%; Mand I2: n = 1; 0.9%) and only 3.6 % (n = 4) in the mandibular canine.

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147 When anterior teeth (incisors and canines) and posterior teeth (premolars and molars) are combined, the data show that the posterior teeth have greater incidence of carious lesions (Pre Latte: n = 168; 13.8 % ; Latte: n = 60; 5.4 % ) than the anterior teeth (Pre Latte: n = 77; 10.7 % ; Latte: n = 17; 2.4 % ) (Table 6 5). Ag ain, as was noted earlier, the Pre Latte anterior dentition has a greater frequency of carious lesions in comparison to the Latte period. Age Differences Carious lesions were compared between age groups to see if the frequency increased with age (Table 6 3 ). However, caution must be taken, as the sample sizes of both children and juveniles are smal l. The Pre Latte population shows a n incre ase of carious lesion frequencies from 40% (n = 2) in juveniles to 78.1% (n = 50) in young adults, which then decrease d to 73.9% (n = 17) in middle aged ad ults. The Latte, however, display a decrease with increased maturity with 31% (n = 4) of the juveniles and 25% (n = 10) of the young adults displaying carious lesions; however, with increasing senility, caries increase in frequency with 31.1% (n = 10) of the middle aged adults affected The disparities between the age groups are slight, as is demonstrated with a Pearson Chi Square test which found no significant age differences in either the Pre Latte (p = 0. 093 at the 0.05 alpha level) or Latte the (p = 0. 741 at the 0.05 alpha level) time periods (Table 6 6). Sex Differences No sex specific differences are apparent in the distribution of carious lesions in either the Pre Latte or Latte individuals (Table 6 1) By individual count, both males and females, of the Pre Latte period, display high frequencies of carious lesions ( n = 25; 75.8% and n = 29; 76.3%, respectively) with females being slightly more prone to

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148 display carious lesions. Similarly, in the Latte p eriod, males and females exhibit low frequencies of carious lesions ( n = 7; 24.1% and n = 11; 29.7%, respectively). However, this may be due to sampling bias as females are more often represented than males in both groups. When looking at carious lesion frequency by tooth count method, males ( 110/644; 17.1%) have a higher frequency than females ( 94/858; 11%) in the Pre Latte period (Table 6 2). Thus, while more women display slightly more carious lesions than men, men are more prone to have multiple inf ected teeth. A Pearson Chi Square test found no signi ficant differences between male and female expression of carious lesions in the Pre Latte (p = 0. 956 at a 0.05 alpha level) or the Latte (P = 0. 613 at a 0.05 alpha level) time periods (Table 6 6). Tim e Period Differences Large differences in carious lesion frequencies are apparent between the Pre Latte and Latte populations. Carious lesions, by individual count, were found in the majority, 72.7% (n = 72) of the Pre Latte population, while only 24.1% ( n = 26) of the Latte individuals displayed carious lesions (Table 6 1). In terms of too th count, the Pre Latte incidence ( 246/1930; 12.7%) of carious lesions is triple what is seen in the Latte period ( 76/1736; 4.4 % ) ( Table 6 2). These findings are supported using a Pearson Chi Square test which found s ignificant differences (p < 0.000 at a 0.05 alpha level) in carious lesion frequencies between the Pre Latte and Latte time periods (Table 6 6). Discussion Tooth Position At first glance, the location of carious lesion, in terms of tooth type, follows the characteristic pattern of an increasing gradient in frequency from the anterior to posterior dentition (Watt et al., 1997; Vodanovic et al., 2005; Han et al., 201 0; Meng et

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149 al., 2011). However, departure for the norm occurs in the Pre Latte sample with uncharacteristically high levels of caries in the anterior teeth The expression of carious lesions in the anterior dentition of the Pre Latte is higher than seen the observed rates in premolars, which are usually more prone to bacterial infection (Hillson, 1996). The location of carious lesions on the ant erior dentition typically occur s at the cemento enamel junction, and at times, the root surface was also involv ed (Figure 1). Similar finding s of high rates of carious lesions in the ante rior dentition was noted in one individual and isolated teeth from the Chelechol ra Orrak cemetery in Palau (Nelson and Fitzpatrick, 2005) and in the Longshan p eriod Kangjia of No rthern China. Nelson and Fitzpatrick (2005) suggest the carious lesions may be related to betel hut chewing ; however, this interpretation seems unlikely given the cariostatic properties of betel nut. N o attempt was made to explain the anterior caries in Northern China (Pechenkina et al., 2002). This type of carious lesion is often associated with periodontal disease where the receding alveolus exposes the root and lesions develop circumferentially around the cemento enamel junction (Hillson, 1996). In a survey of skeletal reports from around the Marianas, Pietrusewsky and colleagues (1997) note a higher frequency of alveolar resorption in two Pre Latte samples, Matapang (44.3%) and Fujita (36.8%), whose rates of resorption exceed that of all other Latte s ites in Guam, except one. Another factor leading to the higher rate of carious lesions in the anterior teeth is the extreme dental crowding noted in the Pre Latte remains (Figure 2), which due to the fragmentary nature was not systematically analyzed. I n the few individuals where maxillary or mandibular reconstruction was possible, dental crowding was evid ent in the

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150 Pre Latte dentition and could increase the amount of food trapped interstitially between the incisors and canines Dental crowding was not observed in the Latte individuals and thus did not likely contribute to carious defects in the anterior teeth. Age Differences No significant difference in carious lesion frequency was found between the age groups in either per iod. The data were re analyzed grouping the adult sub groups (i.e. young adult, middle aged adult, and old adult) and comparing them to the juveniles; likewise, adult sub groups were analyzed separately to see if frequencies varied with senility; however, no significant difference were found in either case (data not shown). The raw frequency data indicates a decrease between the young adult and middle aged adult categories in the Pre Latte sample, whereas in the Latte, there is an increase between those t wo groups. There were no older adults in the Pre Latte individuals and only four available for analysis in the Latte, none of which had carious lesions. The decrease in caries rate with age, in the Pre Latte sample, is surprising given the progressive na ture of carious lesion development, which usually occurs at higher frequencies in older individuals (Thylstrup and Fejerskov, 1994). Dental attrition could explain decrease in caries with age, where the complex morphology of the occlusal surface of the de ntition is worn away, leaving less possibility for infection. The Latte pattern, on the other hand, follows the traditional trend with increase of caries prevalence with age. The effect of age on carious lesion formation has not been addressed in other s tudies from Guam. Sex Differences The current study found no significant differences in caries frequencies between males and females, in either the Pre Latte or the Latte groups, suggesting that both

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151 sexes were eating foods with similar levels of cariogeni city. Analysis of carious lesion frequencies in Taiwan also failed to show significant differences in caries rate between the sexes (Pietrusewsky and Tsang, 2003). Douglas and colleagues (1997), on the other hand, found significant sex differences in cari ous lesion prevalence in the Latte period Apurguan site, located south of the Naton Beach in Tumon Bay. They report a greater frequency of carious lesions, by tooth count, in young adult males in comparison to young adult females and suggest a differentia l access to sweet or sticky foods. However, when combining the age groups, males and females of the Apurguan sample show similar caries frequencies (males: 2.8%, females: 2.1%), which are lower than that found in the current study (males: 3.5%, females: 6 .6%). While not statistically significant, the Latte females of the Naton Beach sample have a higher prevalence of carious lesions than males. This finding is more in line with other studies that have shown sex specific with higher caries rates in femal es (Larsen et al., 1991; Kelley et al., 1991; Lukacs, 1996; Temple and Larsen, 2007). These difference sex specific differences have been attributed to differential access to dietary resources between the sexes due to division of labor. It is likely that this may also be the case in the current sample and is plausible given the linear enamel hypoplasia findings, which show greater incidence of hypoplasias in females. Time Period Differences The Naton Beach sample displays significant differences in cariou s lesion frequencies between the two time periods, with the Pre Latte displaying higher prevalence of caries than the Latte period. In their survey of carious lesions frequencies in various sites in Guam, Pietrusewsky and colleagues (1997) hint at the pos sibility of temporal changes in health and disease between the periods with 14.3% of the Pre

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152 Latte Fujita sample displaying carious lesions, which is higher than the Latte groups that range from 1.3% to 12.3% (Table 6 7 ). However, Pietrusewsky and colleag ues (1997) resist from making any definitive conclusions until larger sample sizes became available. The current study provides a large Pre Latte sample that can be used for comparative purposes. The tooth count frequency of carious lesions in the Pre L atte Naton Beach sample is 12.7%, which is comparable to the Fujita frequency. Likewise, the Latte prevalence of 4.4% falls in line with the other Latte samples whose caries rates range from 1.3% in the Leo Palace Sample to 12.3% in the Right of Way sampl e (see Table 6 x for references). Overall, the caries prevalence in the Pre Latte is higher than the Latte when comparing the various sites across Guam. A significant difference in carious lesion frequency of 8% was found between the Pre Latte and Latte groups; thus, the null hypothesis of no differences in carious lesion frequencies between time periods can be rejected. This study confirms the hypothesis of different caries rates between the Pre Latte and Latte periods; however, the cause of this dispa rity needs to be evaluated in concert with the varying dietary and cultural practices between the populations. The effects of diet The above findings are contrary to the expected results of an increase of carious lesions associated with intensification of agriculture, as was found in many studies throughout the world (Hillson, 1979; Ubelaker, 1980; Larsen, 1981; Cook, 1984; Larsen, 1984; Martin et al., 1984; Smith et al., 1984; Ubelaker, 1984; Larsen et al.,1991; Lukacs, 1992; Pechenkina et al., 2002; Ubel aker and Newson, 2002; Pechenkina et al., 2002; Krigbaum, 2007; Temple, 2 007; Temple and Larsen, 2007). In the current study,

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153 the opposite trend is demonstrated, thus a re analysis of the dietary differences between the groups is warranted. Pre Latte: The Pre Latte population displays much higher rates of carious lesions than would be expected for a pre agricultural society and dental crowding alone may explain an increase in bacterial infection; h owever, other causative factors will also be explored. Tu rner (1979) proposed increasing range of carious lesion prevalence from hunter gatherers (0.0% to 5.3%) to populations with mixed economies (0.44% to 10.3%) and agricultural populations (2.3% to 26.9% ). The combined male and female frequency of Pre Latte sample, 13% falls well within the range of agriculturalists; however, the rates are unexpected ly high for a horticultural population. Pollen and starch analyses found evidence of taro on Pre Latte pottery which has cariogenic properties. Yet, the high prevalence of carious lesions in the Pre Latte would not be expected unless taro was being intensively cultivated and was relied on as a staple crop on a broader scale. Charcoal presence, dating to the Pre Latte in paleoenvironmental cores is indicative o f repeated burning of forest patches (Athens and Ward, 2004). This practice may have been used as a method to clear land for small scale gardening. Thus, it is possible that the Pre Latte were more reliant on taro than had been previously expected and ma y have had participated more in incipient agricultural activities than has been observed through the archaeological record. An alternative explanation for the high rates of cario us lesions may be sugarcane consumption as a positive correlation between long term sugarcane chewing and carious lesions has been established (Frencken et al., 1968). Phytolyth analysis of one Latte period pottery sherd (Hunter Anderson and Moore, 2002) and stable isotope

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154 analyse s (Ambrose e t al., 1997) indicate that sugarcane may have been an important part of the diet in the Latte population. While presence of sugarcane in the Pre Latte period has not been supported through paleoenvironment sediment cores (Athens and Ward, 2004) or other a rchaeological evidence, it is possible that it may have been brought to Guam by the early colonizers (Moore, 2002) as it was domesticated in the southwest Pacific prior to expansion into Remote Oceania (Daniels and Daniels, 1993 ; Grivet et al., 2004 ) La tte: Guam is unique in that it is the only tropical Pacific island to have cultivated rice prior to European colonization (Hunter Anderson et al., 1995). Rice impressions were found on pottery sherds from Tumon Bay that were radiocarbon dated to the Latte 1425) (Moore et al., 1993) and subsequently, rice impressed Latte period pottery sherds were also found in other areas around Guam (Moore, 1994; Moore and Hunter Anderson, 1994). Presence of rice in the Latte Period is further established through identification of rice through phytolith analysis on four potsherds (Loy, 2001a). Given the archaeological evidence for the presence of rice in the Latte period, the low prevalence of caries is unexpected. T he cariogenicity of rice in prehistoric populations is currently being debated (e.g. Temple and Larsen, 2007; Tayles et al., 2000; Talyes et al., 2009). Some studies have shown an increase of carious lesions with the adoption of rice agriculture (Krigb aum, 2007; Temple and Larsen, 2007), while others have shown the op p osite (Oxenham et al., 2000; Tayles et al., 2000; Pietrusewsky and Douglas, 2002; Oxenham et al., 2006; Douglas and Pietrusewsky,

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155 2007; Domett and Tayles, 2007), and a study in living Thai children found a decrease in caries rates in children eating a primarily rice based diet (Kedjarune et al., 1997). Experimental studies have also produced varied results. Rats fed diets of wheat, corn, rice, and oats (independently) developed carious lesions, however, wheat and corn were determined to be the most cariogenic agents (Dodds, 1960). Krasse (1985) showed that the cariogenic potential of rice is low and Sreebny (1983) found no correlation of rice consumption and carious lesions. Grenby (1997) and Lingstrom and colleagues (2000) conclude that the cariogenicity of food starches can be greatly altered by cooking and food processing, and that a mix of processed starches and sugars are more cariogenic than starch alone (Grenby, 1997). Besides rice, the Latte people were known to have consumed other cariogenic foods. Starch grain residues have identified cabbage tree, sugarcane, and taro (Loy, 2001a,b; Loy, 2002; Loy and Crowther, 2002), while pollen and phytolith analyses found evidence of betel palm, breadfruit, coconut, bananas and pandanus ( Hunter Anderson and But ler, 1991 ; Cummings and Puseman, 1998; Dixon et al., 1999; Pearsall and Collins, 2000; Ward, 2000; Cummings, 2002; Athens and Ward, 2004). Further, yam cultivation in the interior of Guam (Moore, 2005) and a preponderance of taro residues on the interior of clay pots (Loy 2001a, 2001b, Low and Crowther, 2002) suggest a preference for these tubers. These data indicate that the prehistoric diet of the Latte period was more diversified than other populations experiencing intensification of agricultural practi ces who usually relied on one staple crop. Hunter Anderson and colleagues (1995) suggest that rice may have been used ceremonially as a prestige food and thus, did contribute signif icantly to the prehistoric

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156 diet. Even if this is the case and rice was no t a dietary staple many of the other primary foods such as sugarcane, breadfruit, bananas, yams, and taro are highly cariogenic. As Grenby (1997) demonstrated, a diet combining starchy foods, such as rice, taro, and yams with sugary foods, such as sugarc ane, breadfruit, bananas would lead to a highly cariogenic diet with a prevalence of dental caries. This finding lends further support to the multifactorial nature of carious etiology that cannot be explained by diet alone. Thus, the low prevalence of ca rious lesions in the Latte sample, in light of demonstrated exploitation of highly cariogenic foods, cannot be explained by dietary choices. The effects of betel nut chewing Another possibility, and the most likely explanation for the low levels of cario us les ions in the Latte period is betel nut chewing, which has been practiced in antiquity to modern times throughout South Asia, Southeast Asia, and the Western Pacific (Strickland, 2002; Zumbroich, 2007). Paleoenvironmental data from Guam have establish ed that betel nut ( Areca catechu ) is indigenous to the Marianas and pre dates human settlement (Athens and Ward, 2004). In the Marianas Islands, the areca nut is usually combined with the betel leaf ( Piper betle ) and slaked lime (CaCO 3 ) (Figure 5) (Hanson and Butler, 1997) and increases stamina, reduces hunger, and creates a sense of euphoria when chewed (Chu, 2001, 2002). Over time, betel nut chewing results in a dark reddish brown stain on the dentition (see Chapter 5, Figure 5 4). The cariostatic eff ects of betel nut have been demonstrated in the epidemiological literature, where the high alkalinity of the areca nut neutralizes acid formation in the mouth, and thus creates an environment unsuitable for dental caries (Chandra and Desai, 1970; Howden, 1 984; Moller et al., 1977; Chatrchaiwiwatana, 2006). Hanson

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157 and Butler (1997:280) outline four mechanisms that may lead to cariostasis with betel chewing: 1. Cleansing of tooth surfaces and thus removing sites of potential carious lesions 2. Appetite reduction which reduces potential intake of cariogenic foods 3. Increase of salivary flow and proteins that inhibit bacterial activity 4. Increase alkalinity of the oral cavity 5. Provides a physical barrier against the spread of cariogenic agents. Thus, the physical and ch emical aspects associated with betel nut chewing have great potential in reducing the prevalence of carious lesions. In the Naton Beach sample, betel nut staining is relegated almost entirely to the Latte period with only three Pre Latte individuals displa ying betel nut staining. The majority of the adult Latte pop ulation displays betel nut stai ning (64%), while no juveniles under the age of 18 were found with staining (Parr, 2012). W hen analysis is restricted to middle aged and older adults, the frequency of betel nut staining rises to 82%. These findings fall within the reported frequency levels reported in the Apurguan sample, 58.7% (Douglas et al., 1997) and an island wide survey, 92% (Hanson and Butler, 1997), both of which also noted low frequencies of carious lesions in the population. These data, coupled with the paucity of betel nut chewing in the Pre Latte and high prevalence of caries suggest that formation of carious lesio ns, in Guam, have an inverse relationship with betel chewing. However, it is interesting to note that the areca nut tree was present in the Pre Latte period (Athens and Ward), and whether betel chewing was initiated with the purpose of combatting poor den tal health is not known. Nonetheless, the data of the current study indicate that the cultural practice of betel nut chewing in the Latte period stymied the effects of a highly cariogenic diet

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158 associated with intensification of agricultural practices, res ulting in better overall dental health. Conclusions This study investigates patterns of carious lesion frequency across time in the prehistoric Chamorro and supports and expands on initial findings of temporal change in relation to dental health reported i n an earlier study (Pietrusewsky et al., 1997). Neither sex specific nor age specific changes were noted, indicating uniformity in diet across the subgroups. The early Pre Latte population displays high rates of carious lesions and an unusual trend of e levated caries in the anterior dentition likely due to dental crowding. The overall caries rates in the Pre Latte are more consistent of a population with an agricultural economy than a h orticultural one (Turner, 1979). This finding may be suggestive inc reased reliance on taro than was previously expected or sugarcane consumption; however, neither of these hypotheses are supported by the archaeological record. The low levels of caries prevalence in the Latte individuals are surprising given their carbohyd rate rich and cariogenic diets. This decrease in carious lesion frequency observed over time is contrary to the expected results as agricultural intensification is usually associated with a higher degree of carious lesions. This study demonstrates that r elationship between diet and dental caries is not as simplistic as it is often reported and analysis of carious lesions needs to be performed with careful consideration of the other factors that may affect dental health such as food processing and other c ultural factors. In this study, cultural practice of betel nut chewing, restricted to the Latte period, had a beneficial effect on the dental health of the Chamorro and limited bacterial

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159 infection and enamel destruction through its physical and chemical p roperties, despite the highly cariogenic diet of the agricultural population.

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160 Table 6 1. Individual occurrence carious lesion frequencies Pre Latte Latte n/N a % n/N a % Males 25/33 75.8 7/29 24.1 Females 29/38 76.3 11/37 29.7 Indeterminate 18/28 64.3 8/42 19.0 Total 72/99 72.7 26/108 24.1 a. n = number of individuals with carious lesions, N = number of individuals analyzed Table 6 2. Tooth count of carious lesion frequencies Pre Latte Latte n/N a % n/N a % Males 110/644 17.1 18/508 3.5 Females 94/858 11.0 46/701 6.6 Indeterminate 42/428 9.8 12/527 2.3 Total 246/1930 12.7 76/1736 4.4 a n = number of individuals with carious lesions, N = number of individuals analyzed Table 6 3. Individual occurrence carious lesions by age Pre Latte Latte n/N % n/N % Child 1/2 50.0 0/11 0.0 Juvenile 2/5 40.0 4/13 30.8 YA 50/64 78.1 10/40 25.0 MA 17/23 73.9 10/32 31.3 OA ------0/4 0.0 n = number of individuals with carious lesions N = number of individuals examined

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161 Table 6 4. Frequency of carious lesions by tooth class Pre Latte Latte Tooth a n/N b % n/N b % Max I1 9/117 7.7 4/105 3.8 Max I2 10/107 9.3 4/111 3.6 Max C 9/121 7.4 4/115 3.5 Max P3 8/123 6.5 6/123 4.9 Max P4 10/140 7.1 3/117 2.6 Max M1 16/135 11.9 4/131 3.1 Max M2 16/128 12.5 5/115 4.3 Max M3 8/53 15.1 4/44 9.1 Mand I1 13/108 12.0 0/144 0.0 Mand I2 16/124 12.9 1/116 0.9 Mand C 20/140 14.3 4/110 3.6 Mand P3 6/147 4.1 3/116 2.6 Mand P4 13/148 8.8 7/122 5.7 Mand M1 27/142 19.0 10/144 6.9 Mand M2 37/125 29.6 14/137 10.2 Mand M3 19/72 26.4 6/70 8.6 a. Right and left sides combined b. n = number of individuals with carious lesions, N = number of individuals analyzed Table 6 5. Frequency of carious lesions by tooth position Anterior teeth Posterior Teeth Time Period n/N a % n/N a % Pre Latte 77/721 10.7 168/1213 13.8 Latte 17/701 2.4 60/1119 5.4 a n = number of individuals with carious lesions, N = number of individuals analyzed

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162 Table 6 6 Pearson Chi Square Test on carious lesion expression Value Df Asymp. Sig (2 sided) Exact Sig. (2 sided) Exact Sig. (1 sided) Time Periods 49.046 1 0.000 0.000* 0.000* Sex within Pre Latte 0.003 1 0.956 + 1.000 + 0.587 + Sex within Latte 0.256 1 0.613 + 0.782 + 0.412 + Age within Pre Latte 6.415 3 0.093 + Age within Latte 1.971 4 0.741 + Indicates statistically significant differences at the 0.05 alpha level + Indicates no statistical relationship at the 0.05 alpha level

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163 Table 6 7 Carious lesion frequencies of the prehistoric Chamorro populations a cross Guam Tooth Count Individual Count Study Sample Subsample Time Period Male % Female % Male % Female % References Naton Beach Pre Latte Pre Latte 17.1 11.0 75.8 76.3 Current Study San Vitores Road Fujita Drainfield Pre Latte 14.3* Bath, 1986; Pietrusewsky, 1986 Matapang Pre Latte 7.6* Right of Way Latte 12.3* Naton Beach Latte Latte 3.5 6.6 29.0 29.7 Current Study Apurguan N/A Latte 2.8 2.1 Douglas et al., 1997; Pietrusewsky et al., 2003 Hyatt Hotel N/A Latte 4.5 5.3 Trembly, 1999 Leo Palace N/A Latte 4.8 1.3 Douglas and Ikehara, 1992 Males and females combined

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164 Figure 6 1. Pr e Latte carious lesions in the anterior dent ition (Photo by author) Figure 6 2. Pre Latte dental crowding (Photo by author) Figure 6 3. Betel n ut with piper leaf and slacked lime (Photo by author).

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165 CHAPTER 7 SUMMARY This research is a diachronic investigation of the evol utionary dynamics of the prehistoric Chamorro population from the Naton Beach Burial Complex in Guam. Patterns of health and disease are analyzed with a focus on biological processes to see how they relate to biocultural and environmental change in the pr ehistoric society. In an island wide survey of health and disease in the prehistoric Chamorro, Pietrusewsky and colleagues (1997) hint at temporal changes in disease frequencies between the time periods and suggest that elevated frequencies of disease are indicative of higher stress levels in some of the earliest Chamorro populations. However, their samples sizes are small and restricted to two sites, thus they hesitate on making definitive conclusions until larger samples became available. The Naton Bea ch skeletal population represents the largest and earliest mortuary assemblage excavated in Guam (DeFant, 2008), and thus allows for a more detailed investigation into diachronic changes occurring over time. The transition between the Pre Latte and Latte time periods are accompanied by changes in population size, diet and subsistence strategies (Hunter Anderson and Butler, 1991; Moore, 2005; Amesbury, 2007). Agricultural intensification with an increased reliance on staple crops of taro, yam, and rice sup plemented by marine resources replace the earlier marine dependent, horticultural and forager subsistence strategies (Hunter Anderson, 1991; Ambrose et al., 1997; Hanson and Butler, 1997). These transitions occur concomitantly with large scale environment al and climatic fluctuations such as sea level decline and increased storminess, aridity, and drought (Hunter Anderson and Butler, 1991; Nunn, 2007, Hunter Anderson, 2010). Thus, it was

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166 predicted that cultural and environmental shifts are likely to be acc ompanied by biological ones, due to increased stress levels associated with malnutrition, limited access to resources, and increased prevalence of disease. The dentition of the Pre Latte sample is significantly larger than the Latte teeth, with an 8% decr ease in overall tooth size over time. Dental reduction did not occur uniformly throughout the dentition and the greatest amount of reduction was observed in the buccolingual dimensions and the posterior teeth. Odontometric trends were analyzed in conjunct ion with data on craniometrics, stature, and pathological differences between the periods (Walth, pers. comm.) to assess which of the proposed mechanisms of dental reduction best fits the data. The high rates of carious lesions and dental crowding in the Pre Latte followed by a significant decrease of caries and reduction in dental crowding in the Latte suggest that there may have been selection for smaller, less complex teeth that are more resistant to dental caries, particularly in populations with soft cariogenic diets. This soft, cariogenic diet would have occurred with a transition to agriculture and reliance on staple crops of taro, yam, and rice accompanied by advanced food processing methods, such as cooking and the use of mortar and pestles, whi ch minimize the force necessary to break down tough food. With decreased functional demands placed on the masticatory apparatus, the maxilla and mandible reduce in size followed by a subsequent decrease in tooth size, as is seen in the Latte population. These findings

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167 There were no significant differences in the rates of juvenile and adult hypoplasias; ho wever, in terms of raw frequency data, juveniles displayed a higher percentage of hypoplastic defects than adults. Thus, there may be a correlation between life expectancy and physiological disruptions. Likewise, male and female hypoplasia frequencies de monstrated no significant differences. Nonetheless, female susceptibility to physiological stress differs between the Pre Latte and Latte periods, where females display fewer hypoplasias than men in the Pre Latte and higher rates of hypoplastic defects in the Latte. This transition indicates there may have been differential access to resources based on gender roles and division of labor. Significant differences in hypoplasia frequencies are demonstrated between the Pre Latte and Latte populations. The Pr e Latte individuals are less prone to hypoplastic defects and thus may not have been exposed to high degrees of physiological stressors as the Latte. Climatic instability, such as typhoons, droughts, and increased aridity, was more common in the Latte per iod, resulting in destruction of crops and reef systems, and likely led to reduced access to nutritional resources and subsequent decrease in health status. Analysis of carious lesion frequencies indicates that the Pre Latte display an unusually elevated n umber of caries in the anterior dentition, which may be the result of dental crowding. No significant differences in carious lesion frequencies were noted between age groups or sex, but males have more infected teeth, overall, than females. A significant decrease in caries rates occurs from the Pre Latte to Latte periods. This pattern is contrary to the expected results as intensification of agriculture is often associated with a higher degree of caries, especially with reliance on highly cariogenic

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168 stap le crops of taro, yam, and rice. The practice of betel nut chewing, which is restricted almost entirely to the Latte period, could explain this disparity as it is known to have cariostatic properties and may lead to better dental health in the Latte despi te the highly cariogenic diet brought on by an agricultural transition. Finally, this study is a clear example of the multifactorial nature of carious lesions and demonstrates the importance of evaluating factors other than diet and warns against making b road generalizations about the relationship between subsistence strategies and carious lesions. Future Studies The majority of archaeological, linguistic, and biological studies have focused on settlement of Remote Oceania through the Lapi ta expansion tha t began around 3 200 BP and led to the colonization of Polynesia (Kirch, 2010). However, settlement of the Marianas Isla nds has been securely dated to 3 600 to 3420 BP (Carson, 2008; Clarke et al., 2010), approximately 400 years before the Lapita peoples ve ntured into the remote Pacific. Biodistance studies investigating settlement patterns of the Marianas Islands have been restricted almost entirely to skeletal remains from the Latte period (Pietru s ewsky, 1990a; 1990b; 1994; 2006; Ishida and Dodo, 1997). T he current study demonstrates that significant biological changes have occurred between the early and late prehistoric Chamorro populations. Thus, comparisons of the Pre Latte Naton Beach sample with circum Pacific skeletal populations may shed new light on settlement and migration patterns of the Marianas Islands and Remote Oceania.

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169 Archaeological, paleoenvironmental, and stable isotope data suggest intensification of agriculture with an increased reliance on terrestrial crops and decrease exploitation of marine resources (Hunter Anderson and Butler, 1991; Ambrose et al., 1997; Athens and Ward, 2004; Moore, 2005; Amesbury, 2007). However, only one stable isotope study from Saipan incorporates samples from the Pre Latte period (n = 5) (McGovern and Wilson, 1996). Analysis of carbon and nitrogen stable isotope ratios from the Pre Latte Naton Beach sample in combination with indicators of dental health will help clarify temporal variability in diet and health as it relates to subsistence change in the prehis toric Chamorro. This study utilizes a diachronic approach to evaluate changes in dental health between the Pre Latte and Latte periods and provides information on the health of some of the earliest settlers of Guam, which was previously unknown. A rchaeolo gical data was evaluated to see if biological disparities in the populations correlate to environmental and cultural changes shifts. Large scale environmental and ecosystem fluctuations necessitated cultural shifts in subsistence strategies, resulting in dental reduction and an increase in physiological disturbances. Carious lesion rates, however, did not increase with the subsistence shifts, and instead improved in the Latte as the result of bete l nut chewing. This temporal assessment of the dentition i dentifies cultural and environmental processes that led to biological change in the prehistoric Chamorro

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170 APPENDIX A DENTAL METRICS Table A 1. Descriptive statistics of dental measurements Measurement N Minimum Maximum Mean Std. Dev. LMax I1 MD 99 7.7 10.4 8.9 0.6 LMax I1 BL 100 6.9 8.9 7.7 0.4 LMax I2 MD 100 6.2 8.6 7.2 0.5 LMax I2 BL 98 6.1 8.7 7.1 0.4 LMax C MD 108 7.1 9.6 8.4 0.5 LMax C BL 111 7.2 10.1 8.8 0.5 LMax P3 MD 120 6.9 10.3 7.9 0.5 LMax P3 BL 121 7.5 11.6 10.1 0.6 LMax P4 MD 133 6.5 9.9 7.6 0.5 LMax P4 BL 130 7.1 12.0 10.1 0.6 LMax M1 MD 119 9.9 12.5 11.3 0.6 LMax M1 BL 127 9.6 13.6 11.9 0.6 LMax M2 MD 118 8.4 12.4 10.5 0.7 LMax M2 BL 124 9.6 14.3 11.9 0.8 LMax M3 MD 47 7.2 13.0 9.4 1.0 LMax M3 BL 50 9.1 13.0 11.4 0.8 RMax I1 MD 102 7.2 10.0 8.8 0.6 RMax I1 BL 107 6.9 8.6 7.7 0.4 RMax I2 MD 96 6.1 8.4 7.2 0.5 RMax I2 BL 97 6.0 8.1 7.1 0.4 RMax C MD 114 7.3 9.6 8.5 0.4 RMax C BL 111 6.7 10.0 8.7 0.5 RMax P3 MD 119 6.2 9.0 7.8 0.5 RMax P3 BL 121 8.5 11.5 10.2 0.6 RMax P4 MD 126 6.2 9.2 7.5 0.5 RMax P4 BL 126 8.5 12.0 10.1 0.6 RMax M1 MD 127 10.1 13.1 11.4 0.6 RMax M1 BL 137 10.4 13.7 11.8 0.6 RMax M2 MD 121 7.6 13.0 10.4 0.7 RMax M2 BL 128 10.0 13.6 11.7 0.8 RMax M3 MD 45 7.0 11.3 9.3 0.9 RMax M3 BL 45 9.6 13.5 11.3 0.8 LMand I1 MD 95 4.1 6.8 5.7 0.4 LMand I1 BL 88 5.4 7.6 6.3 0.4 LMand I2 MD 101 4.9 7.4 6.4 0.5 LMand I2 BL 104 5.6 7.9 6.7 0.4 LMand C MD 117 4.4 8.4 7.3 0.5 LMand C BL 120 6.8 9.6 8.1 0.6

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171 Table A 1. Continued Measurement N Minimum Maximum Mean Std. Dev. LMand P3 MD 134 6.5 9.1 7.9 0.5 LMand P3 BL 133 7.4 10.4 8.9 0.6 LMand P4 MD 132 6.8 9.2 8.0 0.5 LMand P4 BL 128 7.5 10.6 9.0 0.6 LMand M1 MD 126 11.2 13.9 12.5 0.6 LMand M1 BL 140 9.4 12.3 10.8 0.5 LMand M2 MD 122 10.1 13.8 11.9 0.8 LMand M2 BL 130 6.7 12.7 10.6 0.8 LMand M3 MD 62 8.3 14.1 11.0 1.1 LMand M3 BL 67 8.8 12.3 10.3 0.8 RMand I1 MD 87 4.7 6.7 5.7 0.4 RMand I1 BL 83 5.6 7.3 6.3 0.4 RMand I2 MD 104 5.5 7.5 6.5 0.4 RMand I2 BL 105 5.9 7.7 6.8 0.4 RMand C MD 115 6.2 8.4 7.3 0.4 RMand C BL 114 7.0 9.5 8.1 0.5 RMand P3 MD 126 6.1 9.1 7.8 0.5 RMand P3 BL 125 6.1 10.3 8.9 0.6 RMand P4 MD 126 6.6 9.0 7.9 0.5 RMand P4 BL 133 7.3 11.8 9.0 0.6 RMand M1 MD 133 10.5 13.8 12.4 0.6 RMand M1 BL 145 9.6 12.2 10.9 0.5 RMand M2 MD 113 7.9 13.6 11.8 0.8 RMand M2 BL 127 9.2 12.2 10.6 0.6 RMand M3 MD 58 8.8 14.0 11.2 0.9 RMand M3 BL 64 8.4 12.8 10.4 0.8

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172 Table A 2. Descriptive statistics of dental measurements by time period Time Period Measurement N Minimum Maximum Mean Std. Dev. Pre Latte LMax I1 MD 53 8.0 10.4 9.1 0.5 LMax I1 BL 53 7.0 8.9 7.8 0.4 LMax I2 MD 50 6.2 8.6 7.3 0.5 LMax I2 BL 48 6.6 8.7 7.2 0.4 LMax C MD 56 7.6 9.5 8.4 0.5 LMax C BL 53 8.0 10.1 8.8 0.4 LMax P3 MD 58 7.3 10.3 8.1 0.5 LMax P3 BL 60 7.5 11.6 10.4 0.7 LMax P4 MD 71 7.0 9.9 7.8 0.5 LMax P4 BL 71 7.1 12.0 10.3 0.7 LMax M1 MD 56 10.2 12.5 11.5 0.5 LMax M1 BL 63 11.2 13.6 12.2 0.5 LMax M2 MD 64 9.7 12.4 10.9 0.6 LMax M2 BL 63 9.6 14.3 12.3 0.7 LMax M3 MD 29 8.0 13.0 9.6 0.9 LMax M3 BL 29 10.5 13.0 11.6 0.7 RMax I1 MD 48 8.0 10.0 9.0 0.4 RMax I1 BL 53 6.9 8.6 7.8 0.4 RMax I2 MD 47 6.3 8.4 7.4 0.5 RMax I2 BL 48 6.2 8.1 7.2 0.4 RMax C MD 60 7.7 9.5 8.5 0.4 RMax C BL 59 7.8 10.0 8.8 0.5 RMax P3 MD 61 7.2 9.0 8.0 0.4 RMax P3 BL 62 9.4 11.5 10.4 0.5 RMax P4 MD 63 6.6 9.2 7.8 0.5 RMax P4 BL 65 9.2 12.0 10.4 0.6 RMax M1 MD 64 10.6 13.1 11.7 0.5 RMax M1 BL 71 10.6 13.7 12.0 0.6 RMax M2 MD 61 9.7 13.0 10.8 0.6 RMax M2 BL 65 10.0 13.6 12.1 0.7 RMax M3 MD 21 7.7 11.3 9.5 1.0 RMax M3 BL 22 10.4 13.5 11.4 0.7 LMand I1 MD 52 5.4 6.8 5.9 0.3 LMand I1 BL 47 5.6 7.6 6.3 0.4 LMand I2 MD 56 5.7 7.4 6.6 0.4 LMand I2 BL 59 5.8 7.9 6.8 0.4 LMand C MD 68 4.4 8.4 7.3 0.6 LMand C BL 70 6.8 9.4 8.1 0.5 LMand P3 MD 78 7.2 9.1 8.0 0.4 LMand P3 BL 77 7.9 10.4 9.0 0.5 LMand P4 MD 70 7.1 9.2 8.1 0.4

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173 Table A 2. Continued Time Period Measurement N Minimum Maximum Mean Std. Dev. LMand P4 BL 71 8.1 10.6 9.1 0.5 LMand M1 MD 63 11.5 13.9 12.6 0.5 LMand M1 BL 74 9.9 12.3 10.9 0.5 LMand M2 MD 58 10.1 13.8 12.1 0.7 LMand M2 BL 63 9.7 12.7 11.0 0.6 LMand M3 MD 26 10.6 14.1 11.8 0.9 LMand M3 BL 30 9.6 12.3 10.8 0.6 RMand I1 MD 51 5.2 6.7 5.9 0.3 RMand I1 BL 50 5.8 7.3 6.3 0.4 RMand I2 MD 60 5.6 7.4 6.6 0.3 RMand I2 BL 60 5.9 7.7 6.8 0.4 RMand C MD 64 6.5 8.4 7.4 0.4 RMand C BL 64 7.2 9.5 8.2 0.5 RMand P3 MD 69 6.1 9.1 8.0 0.5 RMand P3 BL 70 6.1 10.3 9.0 0.6 RMand P4 MD 65 7.1 9.0 8.1 0.4 RMand P4 BL 74 8.2 10.7 9.2 0.5 RMand M1 MD 62 11.3 13.8 12.6 0.5 RMand M1 BL 68 10.2 12.2 11.0 0.5 RMand M2 MD 48 11.0 13.4 12.1 0.7 RMand M2 BL 58 9.7 11.8 10.9 0.5 RMand M3 MD 30 10.4 14.0 11.8 0.7 RMand M3 BL 35 9.4 12.8 10.9 0.7 Latte LMax I1 MD 46 7.7 9.8 8.6 0.6 LMax I1 BL 47 6.9 8.4 7.6 0.4 LMax I2 MD 50 6.2 8.3 7.0 0.5 LMax I2 BL 50 6.1 7.9 7.0 0.4 LMax C MD 52 7.1 9.6 8.4 0.4 LMax C BL 58 7.2 9.9 8.8 0.5 LMax P3 MD 62 6.9 8.8 7.7 0.4 LMax P3 BL 61 8.8 11.1 9.9 0.5 LMax P4 MD 62 6.5 8.2 7.4 0.4 LMax P4 BL 59 8.7 11.4 9.8 0.5 LMax M1 MD 63 9.9 12.2 11.0 0.6 LMax M1 BL 64 9.6 13.0 11.7 0.6 LMax M2 MD 54 8.4 11.1 10.1 0.6 LMax M2 BL 61 10.3 12.9 11.4 0.7 LMax M3 MD 18 7.2 10.9 9.1 1.1 LMax M3 BL 21 9.1 12.5 11.1 0.9 RMax I1 MD 54 7.2 9.7 8.6 0.6 RMax I1 BL 54 6.9 8.5 7.6 0.4

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174 Table A 2. Continued Time Period Measurement N Minimum Maximum Mean Std. Dev. RMax I2 MD 49 6.1 8.0 7.0 0.5 RMax I2 BL 49 6.0 8.0 7.0 0.4 RMax C MD 54 7.3 9.6 8.4 0.5 RMax C BL 52 6.7 9.8 8.7 0.6 RMax P3 MD 58 6.2 8.8 7.6 0.5 RMax P3 BL 59 8.5 11.3 9.9 0.5 RMax P4 MD 63 6.2 8.0 7.3 0.4 RMax P4 BL 61 8.5 11.0 9.7 0.5 RMax M1 MD 63 10.1 12.6 11.2 0.5 RMax M1 BL 66 10.4 12.9 11.5 0.5 RMax M2 MD 60 7.6 12.3 10.0 0.6 RMax M2 BL 63 10.4 12.8 11.3 0.6 RMax M3 MD 24 7.0 10.4 9.1 0.9 RMax M3 BL 23 9.6 12.8 11.1 0.8 LMand I1 MD 43 4.1 6.2 5.5 0.4 LMand I1 BL 41 5.4 6.8 6.2 0.4 LMand I2 MD 45 4.9 7.3 6.2 0.5 LMand I2 BL 45 5.6 7.4 6.6 0.4 LMand C MD 49 6.0 8.2 7.2 0.5 LMand C BL 50 6.9 9.6 7.9 0.6 LMand P3 MD 56 6.5 8.6 7.6 0.5 LMand P3 BL 56 7.4 10.4 8.7 0.6 LMand P4 MD 62 6.8 8.9 7.8 0.5 LMand P4 BL 57 7.5 10.2 8.8 0.6 LMand M1 MD 63 11.2 13.9 12.4 0.6 LMand M1 BL 66 9.4 12.1 10.6 0.6 LMand M2 MD 64 10.1 13.8 11.7 0.8 LMand M2 BL 67 6.7 11.8 10.3 0.8 LMand M3 MD 36 8.3 12.8 10.5 1.0 LMand M3 BL 37 8.8 11.2 9.9 0.7 RMand I1 MD 36 4.7 6.1 5.5 0.4 RMand I1 BL 33 5.6 6.8 6.2 0.3 RMand I2 MD 44 5.5 7.5 6.3 0.4 RMand I2 BL 45 5.9 7.4 6.6 0.4 RMand C MD 51 6.2 8.1 7.2 0.4 RMand C BL 50 7.0 9.1 8.1 0.5 RMand P3 MD 57 6.7 8.8 7.6 0.5 RMand P3 BL 55 7.6 10.3 8.8 0.6 RMand P4 MD 61 6.6 8.8 7.8 0.5 RMand P4 BL 59 7.3 11.8 8.8 0.7 RMand M1 MD 71 10.5 13.8 12.3 0.6 RMand M1 BL 77 9.6 12.1 10.7 0.6 RMand M2 MD 65 7.9 13.6 11.6 0.9

PAGE 175

175 Table A 2. Continued. Time Period Measurement N Minimum Maximum Mean Std. Dev. RMand M2 BL 69 9.2 12.2 10.4 0.6 RMand M3 MD 28 8.8 11.9 10.6 0.7 RMand M3 BL 29 8.4 10.8 9.7 0.6

PAGE 176

176 Table A 3. Descriptive statistics of dental measurements by time period and sex Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. Pre Latte Male LMax I1 MD 17 8.2 10.1 9.2 0.5 LMax I1 BL 16 7.6 8.5 7.9 0.3 LMax I2 MD 13 6.2 8.4 7.3 0.7 LMax I2 BL 14 6.7 8.0 7.2 0.4 LMax C MD 17 7.8 9.5 8.7 0.5 LMax C BL 15 8.1 9.4 8.8 0.4 LMax P3 MD 21 7.4 10.3 8.2 0.6 LMax P3 BL 21 7.5 11.6 10.4 0.8 LMax P4 MD 22 7.0 9.9 7.9 0.7 LMax P4 BL 23 7.1 12.0 10.4 0.9 LMax M1 MD 15 11.0 12.5 11.8 0.4 LMax M1 BL 20 11.8 13.6 12.5 0.6 LMax M2 MD 20 9.7 11.9 10.9 0.5 LMax M2 BL 21 11.4 13.7 12.5 0.6 LMax M3 MD 10 8.9 13.0 9.9 1.2 LMax M3 BL 10 10.6 13.0 11.8 0.7 RMax I1 MD 14 8.0 9.8 9.1 0.5 RMax I1 BL 18 6.9 8.5 7.8 0.4 RMax I2 MD 14 6.8 8.4 7.5 0.5 RMax I2 BL 15 6.4 8.0 7.2 0.5 RMax C MD 20 8.0 9.5 8.7 0.4 RMax C BL 19 7.8 9.6 8.8 0.5 RMax P3 MD 19 7.4 8.9 8.0 0.4 RMax P3 BL 20 9.7 11.3 10.4 0.5 RMax P4 MD 17 7.3 9.2 7.9 0.6 RMax P4 BL 19 9.4 12.0 10.6 0.6 RMax M1 MD 20 10.6 12.9 11.8 0.5 RMax M1 BL 22 11.6 13.7 12.4 0.6 RMax M2 MD 24 9.9 11.9 10.9 0.6 RMax M2 BL 25 11.1 13.6 12.4 0.6 RMax M3 MD 6 8.4 11.3 9.9 1.2

PAGE 177

177 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMax M3 BL 7 10.8 13.5 12.0 0.8 LMand I1 MD 17 5.6 6.8 5.9 0.3 LMand I1 BL 16 5.6 7.6 6.4 0.5 LMand I2 MD 17 5.9 7.2 6.6 0.3 LMand I2 BL 20 6.3 7.6 6.8 0.3 LMand C MD 21 4.4 8.4 7.6 0.8 LMand C BL 24 7.2 9.2 8.4 0.5 LMand P3 MD 26 7.4 9.1 8.1 0.4 LMand P3 BL 26 8.1 9.9 9.0 0.4 LMand P4 MD 20 7.3 8.9 8.2 0.4 LMand P4 BL 22 8.5 10.6 9.3 0.6 LMand M1 MD 17 11.8 13.8 12.7 0.5 LMand M1 BL 22 10.4 12.0 11.1 0.4 LMand M2 MD 16 10.1 13.3 12.0 0.8 LMand M2 BL 18 10.1 12.0 11.2 0.6 LMand M3 MD 6 10.9 12.7 11.8 0.8 LMand M3 BL 9 9.8 11.6 11.0 0.6 RMand I1 MD 15 5.2 6.2 5.8 0.3 RMand I1 BL 15 5.9 6.9 6.3 0.3 RMand I2 MD 18 5.6 7.3 6.6 0.4 RMand I2 BL 21 6.1 7.6 6.8 0.4 RMand C MD 21 6.7 8.4 7.5 0.4 RMand C BL 21 7.2 9.0 8.2 0.5 RMand P3 MD 23 6.1 9.0 8.0 0.6 RMand P3 BL 24 6.1 10.1 8.9 0.8 RMand P4 MD 20 7.7 9.0 8.3 0.3 RMand P4 BL 24 8.5 10.7 9.4 0.6 RMand M1 MD 19 12.1 13.6 12.9 0.4 RMand M1 BL 22 10.6 12.2 11.3 0.4 RMand M2 MD 11 11.0 13.4 11.9 0.7 RMand M2 BL 15 10.3 11.6 11.1 0.4

PAGE 178

178 Table A 3 Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMand M3 MD 8 11.4 13.1 12.1 0.6 RMand M3 BL 9 10.2 11.6 11.0 0.5 Female LMax I1 MD 21 8.0 9.8 8.9 0.4 LMax I1 BL 23 7.0 8.3 7.7 0.3 LMax I2 MD 22 6.5 8.6 7.3 0.5 LMax I2 BL 20 6.6 7.5 7.0 0.2 LMax C MD 24 7.7 8.8 8.2 0.4 LMax C BL 24 8.0 9.5 8.7 0.4 LMax P3 MD 23 7.3 9.0 8.0 0.4 LMax P3 BL 26 9.2 11.5 10.4 0.5 LMax P4 MD 31 7.1 8.3 7.6 0.4 LMax P4 BL 33 9.2 11.4 10.3 0.4 LMax M1 MD 27 10.4 12.5 11.4 0.5 LMax M1 BL 29 11.3 13.2 12.0 0.4 LMax M2 MD 30 9.7 11.9 10.7 0.5 LMax M2 BL 30 11.4 14.0 12.2 0.6 LMax M3 MD 15 8.0 10.2 9.3 0.7 LMax M3 BL 15 10.5 12.6 11.4 0.6 RMax I1 MD 20 8.3 9.7 9.0 0.4 RMax I1 BL 22 7.0 8.3 7.7 0.4 RMax I2 MD 20 6.3 8.0 7.3 0.5 RMax I2 BL 20 6.2 8.1 7.1 0.4 RMax C MD 27 7.7 9.1 8.3 0.3 RMax C BL 27 8.0 9.4 8.7 0.4 RMax P3 MD 25 7.2 9.0 7.9 0.4 RMax P3 BL 26 9.5 11.4 10.4 0.4 RMax P4 MD 29 6.6 8.5 7.6 0.4 RMax P4 BL 29 9.2 11.1 10.2 0.5 RMax M1 MD 23 10.9 13.1 11.7 0.5 RMax M1 BL 29 10.6 13.0 11.8 0.5 RMax M2 MD 25 9.7 11.7 10.6 0.5

PAGE 179

179 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMax M2 BL 28 10.7 13.3 12.0 0.6 RMax M3 MD 12 7.7 10.7 9.4 1.0 RMax M3 BL 12 10.4 12.2 11.1 0.5 LMand I1 MD 23 5.4 6.6 5.9 0.3 LMand I1 BL 19 5.7 7.0 6.3 0.4 LMand I2 MD 25 5.7 7.4 6.5 0.4 LMand I2 BL 24 5.8 7.3 6.7 0.4 LMand C MD 30 6.5 7.8 7.2 0.3 LMand C BL 29 7.2 8.8 8.0 0.4 LMand P3 MD 32 7.2 9.0 7.9 0.4 LMand P3 BL 31 8.1 10.4 9.0 0.5 LMand P4 MD 33 7.1 9.2 8.0 0.4 LMand P4 BL 34 8.1 9.9 9.0 0.5 LMand M1 MD 27 11.9 13.4 12.5 0.4 LMand M1 BL 32 10.0 12.3 10.8 0.4 LMand M2 MD 24 10.8 13.1 12.0 0.6 LMand M2 BL 27 10.3 12.7 10.8 0.5 LMand M3 MD 13 10.6 13.3 11.4 0.8 LMand M3 BL 14 9.6 11.9 10.5 0.6 RMand I1 MD 25 5.5 6.7 5.9 0.3 RMand I1 BL 23 5.8 7.0 6.3 0.3 RMand I2 MD 24 5.8 7.4 6.5 0.3 RMand I2 BL 22 5.9 7.6 6.8 0.4 RMand C MD 27 6.5 8.2 7.2 0.4 RMand C BL 27 7.3 9.3 8.1 0.5 RMand P3 MD 28 7.3 8.6 7.9 0.3 RMand P3 BL 29 8.4 10.3 9.0 0.5 RMand P4 MD 30 7.1 8.5 7.9 0.4 RMand P4 BL 34 8.2 10.0 9.0 0.4 RMand M1 MD 24 11.3 13.0 12.3 0.4 RMand M1 BL 26 10.2 11.9 10.8 0.3

PAGE 180

180 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMand M2 MD 23 11.2 12.9 12.0 0.6 RMand M2 BL 27 10.2 11.7 10.8 0.4 RMand M3 MD 13 10.4 12.3 11.5 0.5 RMand M3 BL 16 9.4 12.8 10.7 0.8 Indet. LMax I1 MD 15 8.5 10.4 9.3 0.5 LMax I1 BL 14 7.1 8.9 7.9 0.6 LMax I2 MD 15 6.7 8.2 7.4 0.5 LMax I2 BL 14 6.7 8.7 7.4 0.5 LMax C MD 15 7.6 9.2 8.4 0.5 LMax C BL 14 8.0 10.1 8.8 0.6 LMax P3 MD 14 7.3 8.6 8.1 0.4 LMax P3 BL 13 9.0 11.4 10.2 0.7 LMax P4 MD 18 7.2 9.0 7.8 0.5 LMax P4 BL 15 9.2 11.3 10.4 0.6 LMax M1 MD 14 10.2 12.0 11.5 0.5 LMax M1 BL 14 11.2 12.9 12.1 0.4 LMax M2 MD 14 10.2 12.4 11.2 0.6 LMax M2 BL 12 9.6 14.3 12.2 1.1 LMax M3 MD 4 8.6 10.3 9.8 0.8 LMax M3 BL 4 11.3 12.5 11.8 0.6 RMax I1 MD 14 8.3 10.0 9.1 0.5 RMax I1 BL 13 7.2 8.6 7.9 0.5 RMax I2 MD 13 6.9 8.2 7.6 0.4 RMax I2 BL 13 6.5 8.0 7.2 0.4 RMax C MD 13 7.8 9.3 8.7 0.5 RMax C BL 13 7.9 10.0 8.8 0.7 RMax P3 MD 17 7.6 8.6 8.1 0.3 RMax P3 BL 16 9.4 11.5 10.4 0.6 RMax P4 MD 17 6.9 9.1 7.9 0.5 RMax P4 BL 17 9.2 11.1 10.3 0.6 RMax M1 MD 21 10.7 12.4 11.5 0.5

PAGE 181

181 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMax M1 BL 20 11.2 13.1 12.0 0.4 RMax M2 MD 12 9.9 13.0 11.0 0.8 RMax M2 BL 12 10.0 13.1 11.9 0.8 RMax M3 MD 3 8.8 9.7 9.3 0.5 RMax M3 BL 3 10.9 11.5 11.2 0.3 LMand I1 MD 12 5.5 6.6 5.9 0.4 LMand I1 BL 12 5.9 7.1 6.4 0.4 LMand I2 MD 14 6.1 7.2 6.7 0.3 LMand I2 BL 15 6.4 7.9 6.9 0.5 LMand C MD 17 6.6 8.2 7.3 0.5 LMand C BL 17 6.8 9.4 8.0 0.7 LMand P3 MD 20 7.3 9.0 8.1 0.5 LMand P3 BL 20 7.9 9.9 9.0 0.5 LMand P4 MD 17 7.7 9.0 8.2 0.4 LMand P4 BL 15 8.4 10.0 9.2 0.5 LMand M1 MD 19 11.5 13.9 12.7 0.6 LMand M1 BL 20 9.9 11.9 10.9 0.5 LMand M2 MD 18 11.0 13.8 12.4 0.8 LMand M2 BL 18 9.7 12.5 11.1 0.7 LMand M3 MD 7 11.5 14.1 12.4 0.8 LMand M3 BL 7 10.5 12.3 11.1 0.6 RMand I1 MD 11 5.5 6.6 6.0 0.4 RMand I1 BL 12 5.8 7.3 6.4 0.5 RMand I2 MD 18 6.0 7.1 6.5 0.3 RMand I2 BL 17 6.4 7.7 6.8 0.4 RMand C MD 16 6.8 8.3 7.4 0.5 RMand C BL 16 7.6 9.5 8.2 0.6 RMand P3 MD 18 7.3 9.1 8.1 0.5 RMand P3 BL 17 8.1 9.8 9.0 0.5 RMand P4 MD 15 7.4 8.7 8.1 0.4 RMand P4 BL 16 8.5 9.9 9.1 0.5

PAGE 182

182 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMand M1 MD 19 11.5 13.8 12.7 0.6 RMand M1 BL 20 10.4 12.0 10.9 0.4 RMand M2 MD 14 11.0 13.4 12.2 0.8 RMand M2 BL 16 9.7 11.8 11.0 0.6 RMand M3 MD 9 11.0 14.0 12.0 0.9 RMand M3 BL 10 10.2 12.2 11.0 0.6 Latte Male LMax I1 MD 13 7.8 9.8 8.7 0.6 LMax I1 BL 14 6.9 8.4 7.8 0.5 LMax I2 MD 15 6.4 8.1 7.0 0.5 LMax I2 BL 15 6.6 7.7 7.2 0.4 LMax C MD 15 8.1 9.4 8.6 0.4 LMax C BL 15 8.3 9.9 9.1 0.5 LMax P3 MD 17 7.1 8.8 7.8 0.4 LMax P3 BL 17 8.8 11.1 10.1 0.5 LMax P4 MD 16 7.0 8.1 7.6 0.4 LMax P4 BL 15 9.5 11.4 10.2 0.5 LMax M1 MD 15 10.5 12.0 11.3 0.5 LMax M1 BL 15 11.0 12.9 12.0 0.6 LMax M2 MD 10 9.7 11.1 10.4 0.5 LMax M2 BL 13 10.8 12.7 11.8 0.7 LMax M3 MD 8 8.1 10.9 9.7 0.9 LMax M3 BL 10 10.3 12.5 11.4 0.8 RMax I1 MD 15 7.6 9.7 8.7 0.6 RMax I1 BL 16 7.0 8.5 7.7 0.4 RMax I2 MD 13 6.3 7.9 6.9 0.4 RMax I2 BL 14 6.7 7.9 7.2 0.4 RMax C MD 16 8.0 9.5 8.7 0.4 RMax C BL 15 8.2 9.8 8.9 0.4 RMax P3 MD 20 7.2 8.8 7.8 0.4 RMax P3 BL 20 8.8 11.3 10.1 0.5 RMax P4 MD 17 6.8 8.0 7.5 0.3

PAGE 183

183 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMax P4 BL 16 9.4 11.0 10.1 0.5 RMax M1 MD 19 10.6 12.2 11.3 0.5 RMax M1 BL 19 11.0 12.9 11.7 0.5 RMax M2 MD 15 9.4 11.2 10.2 0.5 RMax M2 BL 16 10.6 12.8 11.6 0.7 RMax M3 MD 12 7.9 10.4 9.3 0.8 RMax M3 BL 11 10.1 12.8 11.3 0.9 LMand I1 MD 10 4.8 6.2 5.5 0.4 LMand I1 BL 10 5.5 6.7 6.2 0.4 LMand I2 MD 9 5.4 7.0 6.3 0.5 LMand I2 BL 10 6.4 7.4 6.9 0.3 LMand C MD 15 6.9 8.2 7.6 0.4 LMand C BL 16 7.5 9.6 8.2 0.6 LMand P3 MD 17 7.2 8.6 7.9 0.5 LMand P3 BL 18 8.0 9.8 8.9 0.5 LMand P4 MD 19 7.2 8.9 8.1 0.5 LMand P4 BL 18 8.4 10.2 9.1 0.5 LMand M1 MD 17 11.8 13.4 12.7 0.5 LMand M1 BL 17 10.1 12.1 11.0 0.5 LMand M2 MD 18 10.5 13.8 12.1 0.8 LMand M2 BL 19 10.0 11.8 10.7 0.6 LMand M3 MD 10 8.3 12.8 10.7 1.3 LMand M3 BL 10 9.1 10.8 10.2 0.6 RMand I1 MD 7 4.8 6.0 5.5 0.4 RMand I1 BL 8 5.6 6.8 6.2 0.4 RMand I2 MD 11 5.8 6.9 6.4 0.4 RMand I2 BL 13 5.9 7.4 6.7 0.4 RMand C MD 16 6.9 8.1 7.5 0.4 RMand C BL 16 7.6 9.1 8.4 0.5 RMand P3 MD 17 6.8 8.8 7.9 0.6 RMand P3 BL 16 7.8 10.3 9.0 0.7

PAGE 184

184 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMand P4 MD 19 7.2 8.8 8.0 0.5 RMand P4 BL 19 8.1 11.8 9.1 0.8 RMand M1 MD 20 10.5 13.7 12.6 0.7 RMand M1 BL 22 10.3 11.9 11.0 0.5 RMand M2 MD 18 10.6 13.6 12.0 0.8 RMand M2 BL 22 9.8 12.2 10.7 0.7 RMand M3 MD 10 9.8 11.9 10.8 0.6 RMand M3 BL 10 8.9 10.5 9.8 0.5 Female LMax I1 MD 18 7.7 9.6 8.6 0.6 LMax I1 BL 17 6.9 8.3 7.5 0.5 LMax I2 MD 18 6.3 7.7 7.0 0.4 LMax I2 BL 19 6.1 7.2 6.7 0.4 LMax C MD 19 7.1 8.8 8.2 0.4 LMax C BL 21 7.2 9.1 8.5 0.5 LMax P3 MD 23 6.9 8.4 7.5 0.4 LMax P3 BL 23 9.0 10.9 9.7 0.4 LMax P4 MD 26 6.6 8.0 7.3 0.4 LMax P4 BL 25 8.7 10.4 9.7 0.5 LMax M1 MD 27 10.1 12.0 11.0 0.6 LMax M1 BL 27 9.6 12.5 11.6 0.6 LMax M2 MD 26 8.4 11.1 9.9 0.6 LMax M2 BL 29 10.4 12.8 11.3 0.6 LMax M3 MD 7 7.2 10.0 8.5 0.9 LMax M3 BL 8 9.1 12.4 10.6 1.0 RMax I1 MD 21 7.2 9.6 8.4 0.7 RMax I1 BL 20 6.9 8.3 7.4 0.4 RMax I2 MD 16 6.3 8.0 6.9 0.5 RMax I2 BL 18 6.1 8.0 6.8 0.5 RMax C MD 21 7.3 8.9 8.2 0.4 RMax C BL 20 6.7 9.1 8.4 0.6 RMax P3 MD 24 6.2 8.3 7.5 0.5

PAGE 185

185 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMax P3 BL 24 8.5 10.6 9.8 0.6 RMax P4 MD 29 6.2 8.0 7.2 0.4 RMax P4 BL 28 8.5 10.5 9.5 0.5 RMax M1 MD 23 10.1 12.0 11.1 0.5 RMax M1 BL 26 10.4 12.3 11.4 0.5 RMax M2 MD 27 7.6 10.7 9.8 0.6 RMax M2 BL 29 10.4 12.4 11.2 0.6 RMax M3 MD 9 7.0 10.1 9.0 1.0 RMax M3 BL 9 9.6 12.4 10.9 0.8 LMand I1 MD 14 4.1 6.0 5.4 0.5 LMand I1 BL 13 5.4 6.5 6.0 0.3 LMand I2 MD 19 4.9 6.9 6.1 0.5 LMand I2 BL 18 6.0 7.1 6.5 0.3 LMand C MD 19 6.0 7.3 6.9 0.3 LMand C BL 20 7.0 8.2 7.6 0.3 LMand P3 MD 21 6.5 7.9 7.4 0.4 LMand P3 BL 20 7.4 9.2 8.5 0.5 LMand P4 MD 26 6.9 8.3 7.6 0.4 LMand P4 BL 22 7.7 9.6 8.6 0.5 LMand M1 MD 18 11.2 13.1 12.1 0.5 LMand M1 BL 21 9.4 11.3 10.4 0.4 LMand M2 MD 25 10.1 12.8 11.5 0.7 LMand M2 BL 26 6.7 11.1 10.0 0.9 LMand M3 MD 16 8.7 12.0 10.3 0.9 LMand M3 BL 17 9.0 10.6 9.7 0.5 RMand I1 MD 11 5.0 5.9 5.5 0.3 RMand I1 BL 9 5.8 6.5 6.1 0.3 RMand I2 MD 15 5.5 6.8 6.2 0.4 RMand I2 BL 15 6.2 7.1 6.6 0.3 RMand C MD 17 6.2 7.4 6.9 0.3 RMand C BL 18 7.0 8.2 7.7 0.3

PAGE 186

186 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMand P3 MD 22 6.7 7.8 7.3 0.3 RMand P3 BL 22 7.6 9.1 8.5 0.4 RMand P4 MD 23 6.6 8.2 7.5 0.4 RMand P4 BL 21 7.3 9.4 8.5 0.6 RMand M1 MD 21 11.2 13.0 12.1 0.5 RMand M1 BL 23 9.6 11.5 10.5 0.5 RMand M2 MD 26 7.9 12.7 11.3 1.0 RMand M2 BL 26 9.3 11.1 10.2 0.5 RMand M3 MD 11 8.8 11.5 10.3 0.7 RMand M3 BL 11 8.4 10.8 9.6 0.6 Indet. LMax I1 MD 15 7.7 9.4 8.5 0.5 LMax I1 BL 16 7.0 8.3 7.7 0.3 LMax I2 MD 17 6.2 8.3 7.1 0.5 LMax I2 BL 16 6.4 7.9 7.2 0.4 LMax C MD 18 7.6 9.6 8.6 0.5 LMax C BL 22 7.9 9.9 8.9 0.5 LMax P3 MD 22 7.2 8.7 7.7 0.4 LMax P3 BL 21 9.1 10.7 10.0 0.4 LMax P4 MD 20 6.5 8.2 7.3 0.4 LMax P4 BL 19 8.8 10.4 9.7 0.4 LMax M1 MD 21 9.9 12.2 11.0 0.6 LMax M1 BL 22 10.6 13.0 11.6 0.6 LMax M2 MD 18 9.3 10.9 10.1 0.4 LMax M2 BL 19 10.3 12.9 11.4 0.7 LMax M3 MD 3 7.9 10.0 9.1 1.1 LMax M3 BL 3 10.9 12.5 11.5 0.8 RMax I1 MD 18 7.7 9.7 8.7 0.6 RMax I1 BL 18 7.0 8.5 7.6 0.4 RMax I2 MD 20 6.1 8.0 7.0 0.5 RMax I2 BL 17 6.0 7.7 7.0 0.4 RMax C MD 17 7.6 9.6 8.5 0.5

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187 Table A 3. Continued RMax C BL 17 7.9 9.7 8.8 0.5 RMax P3 MD 14 7.0 8.4 7.7 0.4 RMax P3 BL 15 9.1 10.5 10.0 0.4 RMax P4 MD 17 6.7 8.0 7.4 0.4 RMax P4 BL 17 9.0 10.4 9.7 0.4 RMax M1 MD 21 10.1 12.6 11.1 0.6 RMax M1 BL 21 10.5 12.8 11.5 0.6 RMax M2 MD 18 9.0 12.3 10.1 0.7 RMax M2 BL 18 10.5 12.0 11.2 0.5 RMax M3 MD 3 8.3 9.1 8.7 0.4 RMax M3 BL 3 10.9 11.6 11.2 0.4 LMand I1 MD 19 4.5 6.2 5.6 0.4 LMand I1 BL 18 5.8 6.8 6.3 0.3 LMand I2 MD 17 5.3 7.3 6.2 0.5 LMand I2 BL 17 5.6 7.0 6.6 0.4 LMand C MD 15 6.6 7.9 7.3 0.4 LMand C BL 14 6.9 9.2 8.1 0.6 LMand P3 MD 18 6.9 8.4 7.7 0.5 LMand P3 BL 18 7.5 10.4 8.8 0.7 LMand P4 MD 17 6.8 8.7 7.8 0.5 LMand P4 BL 17 7.5 9.9 8.8 0.7 LMand M1 MD 28 11.4 13.9 12.3 0.6 LMand M1 BL 28 9.7 12.1 10.5 0.6 LMand M2 MD 21 10.3 12.9 11.5 0.7 LMand M2 BL 22 9.5 11.2 10.2 0.5 LMand M3 MD 10 9.2 12.0 10.6 0.9 LMand M3 BL 10 8.8 11.2 10.0 0.8 RMand I1 MD 18 4.7 6.1 5.5 0.4 RMand I1 BL 16 5.6 6.7 6.2 0.3 RMand I2 MD 18 5.6 7.5 6.3 0.5 RMand I2 BL 17 6.0 7.1 6.7 0.3 RMand C MD 18 6.4 8.0 7.3 0.4

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188 Table A 3. Continued Time Period Sex Measurement N Minimum Maximum Mean Std. Dev. RMand C BL 16 7.7 9.0 8.3 0.4 RMand P3 MD 18 6.9 8.5 7.8 0.5 RMand P3 BL 17 7.8 10.3 8.8 0.7 RMand P4 MD 19 7.1 8.7 7.8 0.4 RMand P4 BL 19 7.4 10.2 8.9 0.6 RMand M1 MD 30 11.3 13.8 12.3 0.6 RMand M1 BL 32 9.6 12.1 10.7 0.6 RMand M2 MD 21 10.3 13.3 11.7 0.7 RMand M2 BL 21 9.2 11.5 10.3 0.6 RMand M3 MD 7 9.6 11.5 10.8 0.6 RMand M3 BL 8 8.5 10.7 9.8 0.8

PAGE 189

189 Table A 4. Descriptive statistics of cross sectional area by time period Time Period Measurement N Minimum Maximum Mean Std. Dev. Pre Latte AVG Max I1 CX 38 61.6 81.7 71.0 5.9 AVG Max I2 CX 34 42.9 66.9 52.6 5.4 AVG Max C CX 37 61.6 93.0 74.1 6.4 AVG Max P3 CX 45 70.3 103.2 84.2 7.4 AVG Max P4 CX 50 64.5 102.7 80.3 8.4 AVG Max M1 CX 43 124.6 162.3 141.1 9.8 AVG Max M2 CX 47 111.4 157.2 131.3 10.4 AVG Max M3 CX 13 86.4 120.5 105.1 10.4 AVG Mand I1 CX 36 31.7 45.2 37.6 3.5 AVG Mand I2 CX 39 36.0 55.2 44.6 4.0 AVG Mand C CX 49 43.8 77.4 59.9 7.1 AVG Mand P3 CX 58 54.0 88.2 72.1 7.0 AVG Mand P4 CX 52 60.2 95.6 74.3 6.7 AVG Mand M1 CX 45 120.6 165.2 137.3 10.0 AVG Mand M2 CX 32 106.6 155.6 131.8 11.2 AVG Mand M3 CX 15 101.8 171.9 125.9 17.7 Latte AVG Max I1 CX 35 54.5 80.6 66.3 7.4 AVG Max I2 CX 32 39.2 63.3 49.4 5.5 AVG Max C CX 36 56.8 92.4 74.8 7.6 AVG Max P3 CX 47 61.1 98.9 76.3 7.6 AVG Max P4 CX 49 55.3 88.6 72.5 7.0 AVG Max M1 CX 52 106.8 157.6 129.3 11.9 AVG Max M2 CX 48 87.5 139.2 114.0 11.5 AVG Max M3 CX 11 73.2 127.1 101.2 17.1 AVG Mand I1 CX 26 26.6 40.9 33.6 3.8 AVG Mand I2 CX 31 34.0 51.6 42.0 4.0 AVG Mand C CX 33 44.7 72.3 58.2 7.5 AVG Mand P3 CX 45 53.8 87.5 66.6 8.1 AVG Mand P4 CX 44 52.8 87.5 68.4 7.8 AVG Mand M1 CX 53 111.6 164.3 131.4 12.1 AVG Mand M2 CX 50 94.6 162.8 119.7 14.7 AVG Mand M3 CX 17 85.4 123.3 103.3 10.9

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190 Table A 5. Descriptive statistics of cross sectional area by time period and sex Time Period Sex N Minimum Maximum Mean Std. Dev. Pre Latte Male AVG Max I1 CX 11 61.6 80.7 71.5 5.0 AVG Max I2 CX 10 44.3 66.9 52.9 7.4 AVG Max C CX 10 65.7 85.9 75.7 6.3 AVG Max P3 CX 15 74.4 98.8 85.0 6.7 AVG Max P4 CX 12 74.1 102.7 84.9 9.7 AVG Max M1 CX 12 131.3 161.7 147.5 9.9 AVG Max M2 CX 17 111.4 157.2 134.9 11.4 AVG Max M3 CX 3 100.1 113.2 106.2 6.6 AVG Mand I1 CX 11 32.7 43.0 37.7 3.3 AVG Mand I2 CX 12 41.7 55.2 45.9 3.8 AVG Mand C CX 15 43.8 74.0 63.2 7.6 AVG Mand P3 CX 21 54.0 88.2 72.7 8.3 AVG Mand P4 CX 17 69.6 95.6 77.9 6.9 AVG Mand M1 CX 12 132.1 161.3 144.0 8.2 AVG Mand M2 CX 8 115.0 153.5 130.6 11.0 AVG Mand M3 CX 3 130.6 145.6 140.2 8.4 Female AVG Max I1 CX 16 61.6 78.6 69.3 5.8 AVG Max I2 CX 14 42.9 57.5 51.3 3.8 AVG Max C CX 20 61.6 78.4 71.9 4.0 AVG Max P3 CX 20 70.3 103.2 83.8 7.8 AVG Max P4 CX 27 69.0 92.2 77.6 6.3 AVG Max M1 CX 19 124.6 162.3 138.2 8.9 AVG Max M2 CX 23 113.2 152.9 127.9 9.3 AVG Max M3 CX 9 86.4 120.5 104.1 12.1 AVG Mand I1 CX 17 31.7 43.5 37.4 3.4 AVG Mand I2 CX 15 36.0 50.2 43.4 4.0 AVG Mand C CX 22 48.0 67.9 57.0 4.7 AVG Mand P3 CX 25 62.4 87.9 71.3 6.3 AVG Mand P4 CX 27 60.2 83.0 71.6 5.7 AVG Mand M1 CX 20 120.6 151.7 133.6 7.1 AVG Mand M2 CX 16 119.7 144.6 129.7 7.5 AVG Mand M3 CX 9 101.8 127.0 115.4 7.9 Indet. AVG Max I1 CX 11 63.7 81.7 73.0 6.6 AVG Max I2 CX 10 45.2 61.2 54.1 5.1 AVG Max C CX 7 67.6 93.0 78.1 9.7 AVG Max P3 CX 10 73.2 95.0 83.6 8.1 AVG Max P4 CX 11 64.5 100.0 81.9 9.6 AVG Max M1 CX 12 126.0 154.1 139.3 8.5 AVG Max M2 CX 7 119.6 146.7 133.6 9.1

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191 Table A 5. Continued Time Period Sex N Minimum Maximum Mean Std. Dev. AVG Max M3 CX 1 111.0 111.0 111.0 0.0 AVG Mand I1 CX 8 32.8 45.2 37.8 4.4 AVG Mand I2 CX 12 39.4 52.2 44.7 4.2 AVG Mand C CX 12 49.3 77.4 60.8 8.6 AVG Mand P3 CX 12 59.1 85.1 72.8 6.5 AVG Mand P4 CX 8 70.1 84.5 75.9 5.6 AVG Mand M1 CX 13 121.1 165.2 136.8 12.7 AVG Mand M2 CX 8 106.6 155.6 137.2 16.3 AVG Mand M3 CX 3 126.4 171.9 143.1 25.0 Latte Male AVG Max I1 CX 13 55.4 80.6 67.6 7.5 AVG Max I2 CX 12 43.3 61.3 49.9 4.7 AVG Max C CX 12 68.4 92.4 78.0 7.1 AVG Max P3 CX 16 63.0 98.9 79.3 8.4 AVG Max P4 CX 12 67.4 88.6 77.1 7.0 AVG Max M1 CX 14 117.1 149.0 133.6 10.3 AVG Max M2 CX 8 107.3 139.2 122.1 12.9 AVG Max M3 CX 4 90.1 127.1 111.7 15.6 AVG Mand I1 CX 7 26.6 39.8 34.2 4.5 AVG Mand I2 CX 6 38.8 51.6 44.3 4.7 AVG Mand C CX 11 53.8 72.3 62.9 6.4 AVG Mand P3 CX 13 56.4 87.5 69.4 9.7 AVG Mand P4 CX 14 59.8 87.5 72.4 7.6 AVG Mand M1 CX 14 122.9 149.5 139.1 8.5 AVG Mand M2 CX 14 108.5 162.8 129.1 16.1 AVG Mand M3 CX 7 95.3 123.3 106.3 9.1 Female AVG Max I1 CX 12 54.5 79.2 64.5 8.6 AVG Max I2 CX 11 39.2 52.7 46.4 5.2 AVG Max C CX 14 56.8 76.4 69.9 6.0 AVG Max P3 CX 18 61.1 81.3 72.5 6.2 AVG Max P4 CX 23 55.3 81.9 70.8 6.6 AVG Max M1 CX 21 109.2 146.5 128.1 11.4 AVG Max M2 CX 23 87.5 132.0 111.1 10.7 AVG Max M3 CX 5 73.2 112.1 93.1 17.2 AVG Mand I1 CX 7 29.0 38.4 33.5 3.7 AVG Mand I2 CX 13 34.0 46.2 40.9 3.9 AVG Mand C CX 13 44.7 57.9 51.7 3.7 AVG Mand P3 CX 19 53.8 70.5 62.8 4.9 AVG Mand P4 CX 19 52.8 75.6 65.1 6.4 AVG Mand M1 CX 16 111.6 146.3 125.7 8.9

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192 Table A 5. Continued Time Period Sex N Minimum Maximum Mean Std. Dev. AVG Mand M2 CX 22 94.6 139.2 115.7 12.6 AVG Mand M3 CX 7 85.4 113.3 98.3 10.8 Indet. AVG Max I1 CX 10 60.1 78.6 66.8 5.9 AVG Max I2 CX 9 44.1 63.3 52.4 5.5 AVG Max C CX 10 66.4 88.8 77.6 7.2 AVG Max P3 CX 13 67.0 90.1 77.9 6.9 AVG Max P4 CX 14 61.0 84.3 71.4 6.3 AVG Max M1 CX 17 106.8 157.6 127.3 13.5 AVG Max M2 CX 17 100.2 137.2 114.2 10.7 AVG Max M3 CX 2 89.4 111.1 100.2 15.3 AVG Mand I1 CX 12 29.0 40.9 33.4 3.7 AVG Mand I2 CX 12 37.5 48.5 42.0 3.6 AVG Mand C CX 9 53.4 71.5 61.6 6.2 AVG Mand P3 CX 13 57.6 87.1 69.4 8.5 AVG Mand P4 CX 11 54.7 84.5 69.2 8.2 AVG Mand M1 CX 23 111.8 164.3 130.8 13.7 AVG Mand M2 CX 14 98.3 144.7 116.4 12.8 AVG Mand M3 CX 3 94.3 122.5 108.2 14.1

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193 Table A 6. Group comparisons of dental measurements Measurement Time Period N Mean Std. Dev. Std. Error Mean Difference a Percent of Change b LMax I1 MD Pre Latte 53 9.110 0.489 0.067 0.521 5.72 Latte 46 8.589 0.555 0.082 LMax I1 BL Pre Latte 53 7.824 0.392 0.054 0.179 2.23 Latte 47 7.646 0.427 0.062 LMax I1 CX Pre Latte 50 70.991 6.029 0.853 5.308 7.47 Latte 44 65.683 7.230 1.090 LMax I2 MD Pre Latte 50 7.349 0.526 0.074 0.308 4.19 Latte 50 7.040 0.471 0.067 LMax I2 BL Pre Latte 48 7.185 0.387 0.056 0.173 2.41 Latte 50 7.011 0.439 0.062 LMax I2 CX Pre Latte 47 52.983 5.526 0.806 3.371 6.36 Latte 46 49.612 5.593 0.825 LMax C MD Pre Latte 56 8.440 0.493 0.066 0.020 0.23 Latte 52 8.420 0.450 0.062 LMax C BL Pre Latte 53 8.763 0.448 0.062 0.022 0.26 Latte 58 8.785 0.531 0.070 LMax C CX Pre Latte 51 73.335 6.778 0.949 1.191 1.62 Latte 52 74.526 7.320 1.015 LMax P3 MD Pre Latte 58 8.131 0.490 0.064 0.456 5.61 Latte 62 7.675 0.413 0.052 LMax P3 BL Pre Latte 60 10.358 0.661 0.085 0.430 4.14 Latte 61 9.929 0.461 0.059 LMax P3 CX Pre Latte 57 84.160 7.806 1.034 7.732 9.19 Latte 60 76.427 7.320 0.945 LMax P4 MD Pre Latte 71 7.752 0.531 0.063 0.357 4.61 Latte 62 7.395 0.398 0.051 LMax P4 BL Pre Latte 71 10.347 0.650 0.077 0.511 4.94 Latte 59 9.835 0.524 0.068

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194 Table A 6. Continued Measurement Time Period N Mean Std. Dev. Std. Error Mean Difference a Percent of Change b LMax P4 CX Pre Latte 67 80.442 8.136 0.994 7.553 9.39 Latte 59 72.889 7.251 0.944 LMax M1 MD Pre Latte 56 11.513 0.502 0.067 0.464 4.03 Latte 63 11.049 0.575 0.072 LMax M1 BL Pre Latte 63 12.191 0.519 0.065 0.504 4.13 Latte 64 11.688 0.597 0.075 LMax M1 CX Pre Latte 55 140.416 10.928 1.474 11.066 7.88 Latte 62 129.350 12.511 1.589 LMax M2 MD Pre Latte 64 10.875 0.568 0.071 0.819 7.53 Latte 54 10.055 0.559 0.076 LMax M2 BL Pre Latte 63 12.316 0.719 0.091 0.900 7.31 Latte 61 11.416 0.661 0.085 LMax M2 CX Pre Latte 62 133.699 12.378 1.572 18.631 13.94 Latte 54 115.068 11.628 1.582 LMax M3 MD Pre Latte 29 9.609 0.933 0.173 0.471 4.60 Latte 18 9.138 1.083 0.255 LMax M3 BL Pre Latte 29 11.629 0.678 0.126 0.506 4.35 Latte 21 11.122 0.919 0.200 LMax M3 CX Pre Latte 29 111.867 13.421 2.492 8.727 7.80 Latte 18 103.139 17.202 4.054 RMax I1 MD Pre Latte 48 9.036 0.433 0.063 0.446 4.94 Latte 54 8.589 0.614 0.084 RMax I1 BL Pre Latte 53 7.753 0.430 0.059 0.172 2.21 Latte 54 7.581 0.407 0.055 RMax I1 CX Pre Latte 44 70.210 5.723 0.863 4.908 6.99 Latte 51 65.302 7.722 1.081 RMax I2 MD Pre Latte 47 7.404 0.458 0.067 0.427 5.76 Latte 49 6.977 0.481

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195 Table A 6. Continued Measurement Time Period N Mean Std. Dev. Std. Error Mean Difference a Percent of Change b RMax I2 BL Pre Latte 48 7.156 0.434 0.063 0.159 2.22 Latte 49 6.997 0.446 0.064 RMax I2 CX Pre Latte 45 52.908 5.569 0.830 3.861 7.30 Latte 44 49.047 5.362 0.808 RMax C MD Pre Latte 60 8.505 0.422 0.055 0.069 0.81 Latte 54 8.436 0.475 0.065 RMax C BL Pre Latte 59 8.751 0.480 0.063 0.092 1.05 Latte 52 8.658 0.558 0.077 RMax C CX Pre Latte 57 74.349 6.953 0.921 0.474 0.64 Latte 48 73.875 7.738 1.117 RMax P3 MD Pre Latte 61 7.988 0.372 0.048 0.344 4.30 Latte 58 7.644 0.469 0.062 RMax P3 BL Pre Latte 62 10.403 0.497 0.063 0.470 4.52 Latte 59 9.933 0.542 0.071 RMax P3 CX Pre Latte 60 83.107 7.281 0.940 6.982 8.40 Latte 58 76.126 8.317 1.092 RMax P4 MD Pre Latte 63 7.756 0.510 0.064 0.418 5.40 Latte 63 7.338 0.376 0.047 RMax P4 BL Pre Latte 65 10.354 0.556 0.069 0.609 5.88 Latte 61 9.744 0.523 0.067 RMax P4 CX Pre Latte 62 80.243 8.326 1.057 8.473 10.56 Latte 61 71.770 6.958 0.891 RMax M1 MD Pre Latte 64 11.674 0.493 0.062 0.485 4.16 Latte 63 11.189 0.540 0.068 RMax M1 BL Pre Latte 71 12.027 0.557 0.066 0.488 4.06 Latte 66 11.539 0.529 0.065 RMax M1 CX Pre Latte 62 140.422 10.585 1.344 10.924 7.78 Latte 63 129.498 11.577 1.459

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196 Table A 6. Continued Measurement Time Period N Mean Std. Dev. Std. Error Mean Difference a Percent of Change b RMax M2 MD Pre Latte 61 10.810 0.607 0.078 0.804 7.44 Latte 60 10.006 0.622 0.080 RMax M2 BL Pre Latte 65 12.120 0.665 0.082 0.795 6.56 Latte 63 11.325 0.612 0.077 RMax M2 CX Pre Latte 60 131.269 11.855 1.531 17.861 13.61 Latte 60 113.408 11.628 1.501 RMax M3 MD Pre Latte 21 9.503 0.999 0.218 0.374 3.93 Latte 24 9.130 0.852 0.174 RMax M3 BL Pre Latte 22 11.400 0.716 0.153 0.286 2.51 Latte 23 11.114 0.836 0.174 RMax M3 CX Pre Latte 21 108.557 17.013 3.712 6.796 6.26 Latte 23 101.761 13.657 2.848 LMand I1 MD Pre Latte 52 5.899 0.334 0.046 0.411 6.96 Latte 43 5.488 0.445 0.068 LMand I1 BL Pre Latte 47 6.349 0.429 0.063 0.183 2.89 Latte 41 6.165 0.359 0.056 LMand I1 CX Pre Latte 46 37.637 4.032 0.595 3.618 9.61 Latte 39 34.019 4.549 0.728 LMand I2 MD Pre Latte 56 6.570 0.374 0.050 0.365 5.56 Latte 45 6.204 0.496 0.074 LMand I2 BL Pre Latte 59 6.800 0.398 0.052 0.176 2.59 Latte 45 6.624 0.352 0.052 LMand I2 CX Pre Latte 53 44.739 4.472 0.614 3.632 8.12 Latte 44 41.107 4.493 0.677 LMand C MD Pre Latte 68 7.339 0.575 0.070 0.132 1.80 Latte 49 7.207 0.482 0.069 LMand C BL Pre Latte 70 8.146 0.545 0.065 0.220 2.70 Latte 50 7.925 0.557 0.079

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197 Table A 6. Continued Measurement Time Period N Mean Std. Dev. Std. Error Mean Difference a Percent of Change b LMand C CX Pre Latte 65 59.944 7.959 0.987 2.639 4.40 Latte 46 57.305 7.437 1.097 LMand P3 MD Pre Latte 78 8.041 0.421 0.048 0.394 4.90 Latte 56 7.647 0.490 0.065 LMand P3 BL Pre Latte 77 8.985 0.485 0.055 0.270 3.00 Latte 56 8.715 0.605 0.081 LMand P3 CX Pre Latte 77 72.281 7.083 0.807 5.363 7.42 Latte 54 66.917 8.034 1.093 LMand P4 MD Pre Latte 70 8.115 0.432 0.052 0.316 3.89 Latte 62 7.799 0.503 0.064 LMand P4 BL Pre Latte 71 9.145 0.513 0.061 0.344 3.76 Latte 57 8.802 0.595 0.079 LMand P4 CX Pre Latte 68 74.108 6.922 0.839 5.017 6.77 Latte 56 69.092 8.358 1.117 LMand M1 MD Pre Latte 63 12.593 0.518 0.065 0.217 1.73 Latte 63 12.376 0.604 0.076 LMand M1 BL Pre Latte 74 10.895 0.479 0.056 0.298 2.73 Latte 66 10.598 0.553 0.068 LMand M1 CX Pre Latte 63 137.227 10.985 1.384 5.937 4.33 Latte 62 131.290 12.264 1.558 LMand M2 MD Pre Latte 58 12.141 0.740 0.097 0.466 3.83 Latte 64 11.676 0.790 0.099 LMand M2 BL Pre Latte 63 10.998 0.583 0.073 0.745 6.78 Latte 67 10.252 0.751 0.092 LMand M2 CX Pre Latte 57 134.062 13.600 1.801 13.698 10.22 Latte 64 120.364 14.561 1.820 LMand M3 MD Pre Latte 26 11.764 0.860 0.169 1.265 10.75 Latte 36 10.499 1.012 0.169

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198 Table A 6.. Continued Measurement Time Period N Mean Std. Dev. Std. Error Mean Difference a Percent of Change b LMand M3 BL Pre Latte 30 10.782 0.640 0.117 0.852 7.90 Latte 37 9.931 0.658 0.108 LMand M3 CX Pre Latte 26 126.197 16.266 3.190 21.521 17.05 Latte 36 104.676 14.967 2.494 RMand I1 MD Pre Latte 51 5.913 0.330 0.046 0.416 7.04 Latte 36 5.496 0.370 0.062 RMand I1 BL Pre Latte 50 6.326 0.367 0.052 0.164 2.59 Latte 33 6.162 0.322 0.056 RMand I1 CX Pre Latte 49 37.495 3.846 0.549 3.706 9.88 Latte 32 33.789 3.613 0.639 RMand I2 MD Pre Latte 60 6.560 0.335 0.043 0.233 3.54 Latte 44 6.327 0.411 0.062 RMand I2 BL Pre Latte 60 6.827 0.396 0.051 0.178 2.60 Latte 45 6.649 0.353 0.053 RMand I2 CX Pre Latte 55 44.824 4.091 0.552 2.407 5.36 Latte 42 42.417 3.858 0.595 RMand C MD Pre Latte 64 7.360 0.430 0.054 0.125 1.70 Latte 51 7.235 0.438 0.061 RMand C BL Pre Latte 64 8.165 0.505 0.063 0.078 0.96 Latte 50 8.087 0.498 0.070 RMand C CX Pre Latte 61 60.216 6.657 0.852 1.595 2.64 Latte 47 58.621 6.754 0.985 RMand P3 MD Pre Latte 69 7.982 0.474 0.057 0.351 4.39 Latte 57 7.631 0.525 0.070 RMand P3 BL Pre Latte 70 8.971 0.589 0.070 0.217 2.42 Latte 55 8.754 0.597 0.080 RMand P3 CX Pre Latte 68 71.768 8.142 0.987 5.005 6.97 Latte 55 66.763 8.557 1.154

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199 Table A 6. Continued Measurement Time Period N Mean Std. Dev. Std. Error Mean Difference a Percent of Change b RMand P4 MD Pre Latte 65 8.074 0.411 0.051 0.308 3.81 Latte 61 7.766 0.477 0.061 RMand P4 BL Pre Latte 74 9.154 0.496 0.058 0.318 3.48 Latte 59 8.836 0.705 0.092 RMand P4 CX Pre Latte 64 74.145 7.185 0.898 5.331 7.19 Latte 56 68.814 7.829 1.046 RMand M1 MD Pre Latte 62 12.609 0.508 0.064 0.306 2.43 Latte 71 12.303 0.639 0.076 RMand M1 BL Pre Latte 68 10.993 0.453 0.055 0.261 2.37 Latte 77 10.732 0.555 0.063 RMand M1 CX Pre Latte 62 138.419 10.024 1.273 6.794 4.91 Latte 71 131.624 12.390 1.470 RMand M2 MD Pre Latte 48 12.054 0.675 0.097 0.463 3.84 Latte 65 11.591 0.876 0.109 RMand M2 BL Pre Latte 58 10.935 0.472 0.062 0.581 5.31 Latte 69 10.355 0.619 0.075 RMand M2 CX Pre Latte 48 131.708 12.162 1.755 11.882 9.02 Latte 64 119.826 14.788 1.849 RMand M3 MD Pre Latte 30 11.806 0.700 0.128 1.218 10.32 Latte 28 10.588 0.679 0.128 RMand M3 BL Pre Latte 35 10.859 0.654 0.111 1.119 10.30 Latte 29 9.740 0.615 0.114 RMand M3 CX Pre Latte 30 128.416 13.708 2.503 25.477 19.84 Latte 28 102.939 10.877 2.056 a. Calculated as Latte mean subtracted from Pre Latte mean b. Calculated as mean of Pre Latte subtracted from the mean of the Latte divided by 100 multiplied by 100

PAGE 200

200 Table A 7. Kolmogorov Smirnov a Test for n ormality Measurement Time Period n Statistic df Sig. LMax I1 MD Pre Latte 53 0.094 53 0.200* Latte 46 0.086 46 0.200* LMax I1 BL Pre Latte 53 0.080 53 0.200* Latte 47 0.075 47 0.200* LMax I2 MD Pre Latte 50 0.066 50 0.200* Latte 50 0.109 50 0.193 LMax I2 BL Pre Latte 48 0.109 48 0.200* Latte 50 0.055 50 0.200* LMax C MD Pre Latte 56 0.094 56 0.200* Latte 52 0.071 52 0.200* LMax C BL Pre Latte 53 0.072 53 0.200* Latte 58 0.060 58 0.200* LMax P3 MD Pre Latte 58 0.103 58 0.196 Latte 62 0.079 62 0.200* LMax P3 BL Pre Latte 60 0.130 60 0.013 Latte 61 0.113 61 0.050 LMax P4 MD Pre Latte 71 0.106 71 0.046 Latte 62 0.101 62 0.182 LMax P4 BL Pre Latte 71 0.106 71 0.046 Latte 59 0.101 62 0.182 LMax M1 MD Pre Latte 56 0.085 56 0.200* Latte 63 0.100 63 0.195 LMax M1 BL Pre Latte 63 0.120 63 0.024 Latte 64 0.074 64 0.200* LMax M2 MD Pre Latte 64 0.117 64 0.030 Latte 54 0.068 54 0.200* LMax M2 BL Pre Latte 63 0.091 63 0.200* Latte 61 0.126 61 0.018 LMax M3 MD Pre Latte 29 0.164 29 0.045 Latte 18 0.170 18 0.179 LMax M3 BL Pre Latte 29 0.091 29 0.200* Latte 21 0.100 21 0.200* RMax I1 MD Pre Latte 48 0.108 48 0.200* Latte 54 0.078 54 0.200* RMax I1 BL Pre Latte 53 0.089 53 0.200* Latte 54 0.101 54 0.200* RMax I2 MD Pre Latte 47 0.060 47 0.200* Latte 49 0.075 49 0.200* RMax I2 BL Pre Latte 48 0.081 48 0.200* Latte 54 0.076 49 0.200* RMax C MD Pre Latte 60 0.074 60 0.200*

PAGE 201

201 Table A 7. Continued Measurement Time Period n Statistic df Sig. Latte 52 0.073 54 0.200* RMax C BL Pre Latte 59 0.094 59 0.200* Latte 58 0.078 52 0.200* Rmax P3 MD Pre Latte 61 0.054 61 0.200* Latte 59 0.076 58 0.200* Rmax P3 BL Pre Latte 62 0.074 62 0.200* Latte 63 0.103 59 0.186 Rmax P4 MD Pre Latte 63 0.108 63 0.068 Latte 61 0.053 63 0.200* Rmax P4 BL Pre Latte 65 0.053 65 0.200* Latte 63 0.047 61 0.200* Rmax M1 MD Pre Latte 64 0.087 64 0.200* Latte 66 0.052 63 0.200* Rmax M1 BL Pre Latte 71 0.078 71 0.200* Latte 60 0.072 66 0.200* Rmax M2 MD Pre Latte 61 0.074 61 0.200* Latte 63 0.106 60 0.089 Rmax M2 BL Pre Latte 65 0.086 65 0.200* Latte 24 0.104 63 0.088 Rmax M3 MD Pre Latte 21 0.069 21 0.200* Latte 23 0.107 24 0.200* Rmax M3 BL Pre Latte 22 0.142 22 0.200* Latte 23 0.092 23 0.200* Lmand I1 MD Pre Latte 52 0.127 52 0.034 Latte 43 0.084 43 0.200* Lmand I1 BL Pre Latte 47 0.094 47 0.200* Latte 41 0.103 41 0.200* Lmand I2 MD Pre Latte 56 0.087 56 0.200* Latte 45 0.060 45 0.200* Lmand I2 BL Pre Latte 59 0.108 59 0.083 Latte 45 0.077 45 0.200* Lmand C MD Pre Latte 68 0.109 68 0.045 Latte 49 0.087 49 0.200* Lmand C BL Pre Latte 70 0.089 70 0.200* Latte 50 0.134 50 0.025 Lmand P3 MD Pre Latte 78 0.068 78 0.200* Latte 56 0.100 56 0.200* Lmand P3 BL Pre Latte 77 0.065 77 0.200* Latte 56 0.042 56 0.200* Lmand P4 MD Pre Latte 70 0.056 70 0.200* Latte 62 0.085 62 0.200*

PAGE 202

202 Table A 7. Continued Measurement Time Period n Statistic df Sig. LMand P4 BL Pre Latte 71 0.078 71 0.200* Latte 57 0.082 57 0.200* LMand M1 MD Pre Latte 63 0.056 63 0.200* Latte 63 0.067 63 0.200* LMand M1 BL Pre Latte 74 0.068 74 0.200* Latte 66 0.107 66 0.061 LMand M2 MD Pre Latte 58 0.096 58 0.200* Latte 64 0.112 64 0.044 LMand M2 BL Pre Latte 63 0.097 63 0.200* Latte 67 0.099 67 0.176 LMand M3 MD Pre Latte 26 0.133 26 0.200* Latte 36 0.092 36 0.200* LMand M3 BL Pre Latte 30 0.133 30 0.185 Latte 37 0.104 37 0.200* RMand I1 MD Pre Latte 51 0.127 51 0.040 Latte 36 0.103 36 0.200* RMand I1 BL Pre Latte 50 0.097 50 0.200* Latte 33 0.109 33 0.200* RMand I2 MD Pre Latte 60 0.099 60 0.200* Latte 44 0.070 44 0.200* RMand I2 BL Pre Latte 60 0.073 60 0.200* Latte 45 0.092 45 0.200* RMand C MD Pre Latte 64 0.069 64 0.200* Latte 51 0.052 51 0.200* RMand C BL Pre Latte 64 0.059 64 0.200* Latte 50 0.057 50 0.200* RMand P3 MD Pre Latte 69 0.094 69 0.200* Latte 57 0.116 57 0.055 RMand P3 BL Pre Latte 70 0.085 70 0.200* Latte 55 0.083 55 0.200* RMand P4 MD Pre Latte 65 0.062 65 0.200* Latte 61 0.071 61 0.200* RMand P4 BL Pre Latte 74 0.095 74 0.097 Latte 59 0.135 59 0.009 RMand M1 MD Pre Latte 62 0.055 62 0.200* Latte 71 0.073 71 0.200* RMand M1 BL Pre Latte 68 0.103 68 0.071 Latte 77 0.081 77 0.200* RMand M2 MD Pre Latte 48 0.143 48 0.015 Latte 65 0.092 65 0.200* RMand M2 BL Pre Latte 58 0.087 58 0.200*

PAGE 203

203 Table A 7. Continued Measurement Time Period n Statistic df Sig. Latte 69 0.070 69 0.200* RMand M3 MD Pre Latte 30 0.119 30 0.200* Latte 28 0.069 28 0.200* RMand M3 BL Pre Latte 35 0.093 35 0.200* Latte 29 0.131 29 0.200* a. Lilliefors Significance Correction *This is a lower bound of the true significance.

PAGE 204

204 Table A 8. Levene's Test of Homogeneity of Variance based on the mean Measurement Levene Statistic df1 df2 Sig. LMax I1 MD 0.674 1 97 0.414 LMax I1 BL 0.884 1 98 0.349 LMax I2 MD 0.427 1 98 0.515 LMax I2 BL 1.638 1 96 0.204 LMax C MD 1.952 1 106 0.165 LMax C BL 1.145 1 109 0.287 LMax P3 MD 0.448 1 118 0.504 LMax P3 BL 2.538 1 119 0.114 LMax P4 MD 1.869 1 131 0.174 LMax P4 BL 0.113 1 128 0.737 LMax M1 MD 1.520 1 117 0.220 LMax M1 BL 0.871 1 125 0.352 LMax M2 MD 0.405 1 116 0.526 LMax M2 BL 0.072 1 122 0.790 LMax M3 MD 2.212 1 45 0.144 LMax M3 BL 1.829 1 48 0.183 RMax I1 MD 6.234 1 100 0.014 RMax I1 BL 0.512 1 105 0.476 RMax I2 MD 0.302 1 94 0.584 RMax I2 BL 0.003 1 95 0.960 RMax C MD 0.368 1 112 0.545 RMax C BL 0.896 1 109 0.346 RMax P3 MD 1.952 1 117 0.165 RMax P3 BL 0.169 1 119 0.682 RMax P4 MD 4.465 1 124 0.037 RMax P4 BL 0.045 1 124 0.833 RMax M1 MD 0.704 1 125 0.403 RMax M1 BL 0.015 1 135 0.902 RMax M2 MD 0.286 1 119 0.594 RMax M2 BL 0.068 1 126 0.795 RMax M3 MD 0.859 1 43 0.359 RMax M3 BL 0.882 1 43 0.353 LMand I1 MD 3.333 1 93 0.071 LMand I1 BL 0.256 1 86 0.614 LMand I2 MD 4.023 1 99 0.048 LMand I2 BL 0.187 1 102 0.666 LMand C MD 0.008 1 115 0.929 LMand C BL 0.034 1 118 0.855 LMand P3 MD 2.809 1 132 0.096 LMand P3 BL 2.807 1 131 0.096 LMand P4 MD 2.733 1 130 0.101 LMand P4 BL 0.857 1 126 0.356 Table A 8. Continued

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205 Measurement Levene Statistic df1 df2 Sig. LMand M1 MD 1.438 1 124 0.233 LMand M1 BL 1.319 1 138 0.253 LMand M2 MD 1.026 1 120 0.313 LMand M2 BL 1.763 1 128 0.187 LMand M3 MD 0.956 1 60 0.332 LMand M3 BL 0.045 1 65 0.832 RMand I1 MD 0.481 1 85 0.490 RMand I1 BL 0.269 1 81 0.606 RMand I2 MD 3.097 1 102 0.081 RMand I2 BL 0.198 1 103 0.657 RMand C MD 0.021 1 113 0.884 RMand C BL 0.012 1 112 0.913 RMand P3 MD 3.359 1 124 0.069 RMand P3 BL 1.095 1 123 0.297 RMand P4 MD 1.086 1 124 0.299 RMand P4 BL 1.984 1 131 0.161 RMand M1 MD 2.773 1 131 0.098 RMand M1 BL 2.673 1 143 0.104 RMand M2 MD 0.076 1 111 0.783 RMand M2 BL 2.637 1 125 0.107 RMand M3 MD 0.048 1 56 0.828 RMand M3 BL 0.029 1 62 0.866

PAGE 206

206 APPENDIX B ANALYSES OF VARIANCE TESTS FOR D ENTAL MEASUREMENTS

PAGE 207

207 Table B 1. Two Way Factorial ANOVA for LMax I1 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.596 a 3 1.199 A.53 0.00604 Intercept 5239.096 1 5239.096 19797.321 0.00 TimePeriod 2.852 1 2.852 10.777 0.00 SexCombined 0.622 1 0.622 2.351 0.13 TimePeriod SexCombined 0.008 1 0.008 0.031 0.86 Error 17.201 65 0.265 Total 5429.299 69 Corrected Total 20.798 68 R Squared = 0.173 (Adjusted R Squared = 0.135) Table B 2. Two Way Factorial ANOVA for LMax I1 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 1.209 a 3 0.403 2.664 0.05503 Intercept 4046.654 1 4046.654 26742.88 0.00 TimePeriod 0.418 1 0.418 2.763 0.10 SexCombined 0.727 1 0.727 4.807 0.03 TimePeriod SexCombined 0.092 1 0.092 0.606 0.44 Error 9.987 66 0.151 Total 4189.417 70 Corrected Total 11.196 69 R Squared = 0.108 (Adjusted R Squared = 0.067)

PAGE 208

208 Table B 3. Two Way Factorial ANOVA for LMax I2 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 1.651 a 3 0.55 2.111 0.10751 Intercept 3342.829 1 3342.829 12821.516 0.00 TimePeriod 1.46 1 1.46 5.599 0.02 SexCombined 0.014 1 0.014 0.052 0.82 TimePeriod SexCombined 0.026 1 0.026 0.098 0.75 Error 16.686 64 0.261 Total 3501.821 68 Corrected Total 18.337 67 R Squared = 0.090 (Adjusted R Squared = 0.047)

PAGE 209

209 Table B 4. Two Way Factorial ANOVA for LMax I2 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.054 a 3 1.018 8.652 0.00007 a Intercept 3285.352 1 3285.352 27927.215 0.00 TimePeriod 0.494 1 0.494 4.202 0.04 b SexCombined 1.848 1 1.848 15.709 0.00 b TimePeriod SexCombined 0.573 1 0.573 4.875 0.03 b Error 7.529 64 0.118 Total 3350.988 68 Corrected Total 10.583 67 R Squared = 0.289 (Adjusted R Squared = 0.255) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 210

210 Table B 5. Two Way Factorial ANOVA for LMax C MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.989 a 3 1.33 8.015 0.00011 a Intercept 5170.027 1 5170.027 31161.041 0.00 TimePeriod 0.193 1 0.193 1.161 0.28 SexCombined 3.781 1 3.781 22.787 0.00 b TimePeriod SexCombined 0.008 1 0.008 0.048 0.83 Error 11.78 71 0.166 Total 5305.585 75 Corrected Total 15.769 74 R Squared = 0.253 (Adjusted R Squared = 0.221) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 211

211 Table B 6. Two Way Factorial ANOVA for LMax C BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.367 a 3 1.122 5.939 0.00113027 Intercept 5511.714 1 5511.714 29166.158 0.00 TimePeriod 0.006 1 0.006 0.034 0.85 SexCombined 2.056 1 2.056 10.881 0.00 TimePeriod SexCombined 1.387 1 1.387 7.337 0.01 Error 13.417 71 0.189 Total 5730.832 75 Corrected Total 16.785 74 R Squared = 0.201 (Adjusted R Squared = 0.167)

PAGE 212

212 Table B 7. Two Way Factorial ANOVA for LMax P3 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 6.572 a 3 2.191 9.997 0.00001 a Intercept 5168.768 1 5168.768 23586.009 0.00 TimePeriod 4.613 1 4.613 21.049 0.00 b SexCombined 1.462 1 1.462 6.673 0.01 b TimePeriod SexCombined 0.072 1 0.072 0.329 0.57 Error 17.532 80 0.219 Total 5268.44 84 Corrected Total 24.104 83 R Squared = 0.273 (Adjusted R Squared = 0.245) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 213

213 Table B 8. Two Way Factorial ANOVA for LMax P3 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 6.839 a 3 2.28 6.899 0.00033 a Intercept 8777.969 1 8777.969 26566.965 0.00 TimePeriod 4.814 1 4.814 14.571 0.00 b SexCombined 1.02 1 1.02 3.086 0.08 TimePeriod SexCombined 0.512 1 0.512 1.549 0.22 Error 27.424 83 0.33 Total 9033.594 87 Corrected Total 34.262 86 R Squared = 0.200 (Adjusted R Squared = 0.171) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 214

214 Table B 9. Two Way Factorial ANOVA for LMax P4 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.817 a 3 1.272 5.717 0.00125 Intercept 5169.992 1 5169.992 23228.853 0.00 TimePeriod 2.061 1 2.061 9.262 0.00 SexCombined 1.526 1 1.526 6.857 0.01 TimePeriod SexCombined 0.018 1 0.018 0.079 0.78 Error 20.254 91 0.223 Total 5496.688 95 Corrected Total 24.071 94 R Squared = 0.159 (Adjusted R Squared = 0.131)

PAGE 215

215 Table B 10. Two Way Factorial ANOVA for LMax P4 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 7.387 a 3 2.462 6.679 0.00040 a Intercept 9134.581 1 9134.581 24776.573 0.00 TimePeriod 3.061 1 3.061 8.303 0.00 b SexCombined 2.669 1 2.669 7.239 0.01 b TimePeriod SexCombined 0.918 1 0.918 2.491 0.12 Error 33.918 92 0.369 Total 9921.115 96 Corrected Total 41.305 95 R Squared = 0.179 (Adjusted R Squared = 0.152) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 216

216 Table B 11. Two Way Factorial ANOVA for LMax M1 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 6.835 a 3 2.278 8.657 0.00005 a Intercept 9960.838 1 9960.838 37847.924 0.00 TimePeriod 3.734 1 3.734 14.187 0.00 b SexCombined 2.77 1 2.77 10.524 0.00 b TimePeriod SexCombined 0 1 0 0 1.00 Error 21.054 80 0.263 Total 10771.033 84 Corrected Total 27.89 83 R Squared = .245 (Adjusted R Squared = .217) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 217

217 Table B 12. Two Way Factorial ANOVA for LMax M1 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 9.023 a 3 3.008 10.362 0.00001 a Intercept 12272.892 1 12272.892 42282.653 0.00 TimePeriod 4.685 1 4.685 16.141 0.00 b SexCombined 3.665 1 3.665 12.626 0.00 b TimePeriod SexCombined 0.099 1 0.099 0.34 0.56 Error 25.252 87 0.29 Total 13107.333 91 Corrected Total 34.275 90 R Squared = 0.263 (Adjusted R Squared = 0.238) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level Table B 13. Two Way Factorial ANOVA for LMax M2 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 13.957 a 3 4.652 15.346 0.00000 a Intercept 7942.252 1 7942.252 26198.798 0.00 TimePeriod 7.42 1 7.42 24.475 0.00 b SexCombined 2.185 1 2.185 7.208 0.01 b TimePeriod SexCombined 0.623 1 0.623 2.056 0.16 Error 24.859 82 0.303 Total 9494.712 86 Corrected Total 38.815 85 R Squared = 0.360 (Adjusted R Squared = 0.336) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 218

218 Table B 14. Two Way Factorial ANOVA for LMax M2 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 22.942 a 3 7.647 21.584 0.0000 a Intercept 11879.491 1 11879.491 33527.741 0.00 TimePeriod 14.303 1 14.303 40.368 0.00 b SexCombined 4.26 1 4.26 12.023 0.00 b TimePeriod SexCombined 0.165 1 0.165 0.466 0.50 Error 31.534 89 0.354 Total 13283.297 93 Corrected Total 54.477 92 R Squared = 0.421 (Adjusted R Squared = 0.402) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level Table B 15. Two Way Factorial ANOVA for LMax M3 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 9.775 a 3 3.258 3.705 0.02019 Intercept 3234.598 1 3234.598 3677.787 0.00 TimePeriod 2.543 1 2.543 2.892 0.10 SexCombined 7.939 1 7.939 9.027 0.00 TimePeriod SexCombined 1.06 1 1.06 1.205 0.28 Error 31.662 36 0.879 Total 3588.821 40 Corrected Total 41.437 39 R Squared = 0.236 (Adjusted R Squared = 0.172)

PAGE 219

219 Table B 16. Two Way Factorial ANOVA for LMax M3 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 6.851 a 3 2.284 3.934 0.01518 Intercept 5238.222 1 5238.222 9024.631 0.00 TimePeriod 3.945 1 3.945 6.796 0.01 SexCombined 3.695 1 3.695 6.366 0.02 TimePeriod SexCombined 0.427 1 0.427 0.736 0.40 Error 22.637 39 0.58 Total 5588.395 43 Corrected Total 29.488 42 R Squared = 0.232 (Adjusted R Squared = 0.173) Table B 17. Two Way Factorial ANOVA for RMax I1 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 4.297a 3 1.432 4.882 0.00398 Intercept 5248.785 1 5248.785 17890.567 0.00 TimePeriod 3.281 1 3.281 11.182 0.00 SexCombined 0.496 1 0.496 1.692 0.20 TimePeriod SexCombined 0.157 1 0.157 0.537 0.47 Error 19.363 66 0.293 Total 5410.721 70 Corrected Total 23.661 69 R Squared = 0.182 (Adjusted R Squared = 0.144)

PAGE 220

220 Table B 18. Two Way Factorial ANOVA for RMax I1 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 1.381 a 3 0.46 2.781 0.04712 Intercept 4379.103 1 4379.103 26459.067 0.00 TimePeriod 0.436 1 0.436 2.637 0.11 SexCombined 0.726 1 0.726 4.388 0.04 TimePeriod SexCombined 0.181 1 0.181 1.093 0.30 Error 11.916 72 0.166 Total 4447.095 76 Corrected Total 13.297 75 R Squared = 0.104 (Adjusted R Squared = 0.066) Table B 19. Two Way Factorial ANOVA for RMax I2 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.021 a 3 1.007 4.563 0.00609 Intercept 3129.101 1 3129.101 14179.744 0.00 TimePeriod 2.903 1 2.903 13.156 0.00 SexCombined 0.087 1 0.087 0.396 0.53 TimePeriod SexCombined 0.201 1 0.201 0.911 0.34 Error 13.02 59 0.221 Total 3238.189 63 Corrected Total 16.041 62 R Squared = 0.188 (Adjusted R Squared = 0.147)

PAGE 221

221 Table B 20. Two Way Factorial ANOVA for RMax I2 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 2.116 a 3 0.705 3.709 0.01598 Intercept 3287.914 1 3287.914 17291.212 0.00 TimePeriod 0.347 1 0.347 1.827 0.18 SexCombined 1.476 1 1.476 7.764 0.01 TimePeriod SexCombined 0.263 1 0.263 1.383 0.24 Error 11.979 63 0.19 Total 3353.617 67 Corrected Total 14.095 66 R Squared = 0.150 (Adjusted R Squared = 0.110)

PAGE 222

222 Table B 21. Two Way Factorial ANOVA for RMax C MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.435 a 3 1.145 8.345 0.00007 a Intercept 5804.658 1 5804.658 42306.754 0.00 TimePeriod 0.045 1 0.045 0.328 0.57 SexCombined 3.381 1 3.381 24.641 0.00 b TimePeriod SexCombined 0.047 1 0.047 0.34 0.56 Error 10.976 80 0.137 Total 5986.055 84 Corrected Total 14.411 83 R Squared = 0.238 (Adjusted R Squared = 0.210) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 223

223 Table B 22. Two Way Factorial ANOVA for RMax C BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.010 a 3 1.003 4.726 0.00443 Intercept 5859.032 1 5859.032 27594.638 0.00 TimePeriod 0.114 1 0.114 0.539 0.47 SexCombined 1.852 1 1.852 8.723 0.00 TimePeriod SexCombined 1.26 1 1.26 5.933 0.02 Error 16.349 77 0.212 Total 6118.622 81 Corrected Total 19.359 80 R Squared = 0.155 (Adjusted R Squared = 0.123) Table B 23. Two Way Factorial ANOVA for RMax P3 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.284 a 3 1.095 5.804 0.00118 Intercept 5279.585 1 5279.585 27987.8 0.00 TimePeriod 2.313 1 2.313 12.262 0.00 SexCombined 0.637 1 0.637 3.378 0.07 TimePeriod SexCombined 0.198 1 0.198 1.049 0.31 Error 15.846 84 0.189 Total 5358.863 88 Corrected Total 19.13 87 R Squared = 0.172 (Adjusted R Squared = 0.142)

PAGE 224

224 Table B 24. Two Way Factorial ANOVA for RMax P3 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 6.590 a 3 2.197 8.209 0.00007 a Intercept 9185.092 1 9185.092 34324.858 0.00 TimePeriod 4.824 1 4.824 18.026 0.00 b SexCombined 1.015 1 1.015 3.792 0.05 TimePeriod SexCombined 0.464 1 0.464 1.734 0.19 Error 23.013 86 0.268 Total 9326.817 90 Corrected Total 29.603 89 R Squared = 0.223 (Adjusted R Squared = 0.195) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level Table B 25. Two Way Factorial ANOVA for RMax P4 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 5.308 a 3 1.769 9.707 0.00001 a Intercept 4896.429 1 4896.429 26863.383 0.00 TimePeriod 3.521 1 3.521 19.32 0.00 b SexCombined 1.696 1 1.696 9.303 0.00 b TimePeriod SexCombined 0.039 1 0.039 0.217 0.64 Error 16.04 88 0.182 Total 5224.436 92 Corrected Total 21.348 91 R Squared = 0.249 (Adjusted R Squared = 0.223) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 225

225 Table B 26. Two Way Factorial ANOVA for RMax P4 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 13.591 a 3 4.53 17.023 0.00000 a Intercept 8849.841 1 8849.841 33254.095 0.00 TimePeriod 7.204 1 7.204 27.07 0.00 b SexCombined 4.575 1 4.575 17.19 0.00 b TimePeriod SexCombined 0.295 1 0.295 1.107 0.30 Error 23.419 88 0.266 Total 9386.412 92 Corrected Total 37.01 91 R Squared = 0.367 (Adjusted R Squared = 0.346) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level Table B 27. Two Way Factorial ANOVA for RMax M1 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 6.817 a 3 2.272 9.041 0.00003 a Intercept 11153.872 1 11153.872 44379.428 0.00 TimePeriod 6.286 1 6.286 25.011 0.00 b SexCombined 0.293 1 0.293 1.166 0.28 TimePeriod SexCombined 0.06 1 0.06 0.238 0.63 Error 20.358 81 0.251 Total 11257.388 85 Corrected Total 27.175 84 R Squared = 0.251 (Adjusted R Squared = 0.223) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 226

226 Table B 28. Two Way Factorial ANOVA for RMax M1 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 10.285 a 3 3.428 13.18 0.00000 a Intercept 13092.688 1 13092.688 50334.755 0.00 TimePeriod 5.236 1 5.236 20.132 0.00 b SexCombined 4.575 1 4.575 17.59 0.00 b TimePeriod SexCombined 0.53 1 0.53 2.037 0.16 Error 23.93 92 0.26 Total 13430.062 96 Corrected Total 34.215 95 R Squared = 0.301 (Adjusted R Squared = 0.278) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level Table B 29. Two Way Factorial ANOVA for RMax M2 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 16.801 a 3 5.6 18.605 0.0000 a Intercept 9320.397 1 9320.397 30962.461 0.00 TimePeriod 11.614 1 11.614 38.581 0.00 b SexCombined 2.76 1 2.76 9.168 0.00 c TimePeriod SexCombined 0.044 1 0.044 0.145 0.70 Error 26.189 87 0.301 Total 9869.333 91 Corrected Total 42.99 90 R Squared = 0.391 (Adjusted R Squared = 0.370) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 227

227 Table B 30. Two Way Factorial ANOVA for RMax M2 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 19.825 a 3 6.608 17.436 0.00000 a Intercept 12923.433 1 12923.433 34099.631 0.00 TimePeriod 13.13 1 13.13 34.645 0.00 b SexCombined 4.096 1 4.096 10.807 0.00 c TimePeriod SexCombined 0.001 1 0.001 0.002 0.97 Error 35.625 94 0.379 Total 13716.55 98 Corrected Total 55.45 97 R Squared = 0.358 (Adjusted R Squared = 0.337) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level Table B 31. Two Way Factorial ANOVA for RMax M3 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.030 a 3 1.01 1.057 0.37980 Intercept 3176.591 1 3176.591 3323.831 0.00 TimePeriod 1.985 1 1.985 2.077 0.16 SexCombined 1.799 1 1.799 1.882 0.18 TimePeriod SexCombined 0.052 1 0.052 0.055 0.82 Error 33.45 35 0.956 Total 3445.583 39 Corrected Total 36.48 38 R Squared = 0.083 (Adjusted R Squared = 0.004)

PAGE 228

228 Table B 32. Two Way Factorial ANOVA for RMax M3 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 4.869 a 3 1.623 2.617 0.06636 Intercept 4784.046 1 4784.046 7712.077 0.00 TimePeriod 2.036 1 2.036 3.282 0.08 SexCombined 3.336 1 3.336 5.377 0.03 TimePeriod SexCombined 0.56 1 0.56 0.903 0.35 Error 21.712 35 0.62 Total 4975.126 39 Corrected Total 26.581 38 R Squared = 0.183 (Adjusted R Squared = 0.113) Table B 33. Two Way Factorial ANOVA for LMand I1 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.498 a 3 1.166 8.023 0.00014 a Intercept 1881.394 1 1881.394 12944.752 0.00 TimePeriod 3.145 1 3.145 21.636 0.00 b SexCombined 0.153 1 0.153 1.054 0.31 TimePeriod SexCombined 0.029 1 0.029 0.202 0.66 Error 8.72 60 0.145 Total 2109.172 64 Corrected Total 12.219 63 R Squared = 0.286 (Adjusted R Squared = 0.251) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 229

229 Table B 34. Two Way Factorial ANOVA for LMand I1 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 1.021 a 3 0.34 1.996 0.12549 Intercept 2117.819 1 2117.819 12423.168 0.00 TimePeriod 0.805 1 0.805 4.724 0.03 SexCombined 0.147 1 0.147 0.86 0.36 TimePeriod SexCombined 0.029 1 0.029 0.17 0.68 Error 9.206 54 0.17 Total 2266.611 58 Corrected Total 10.226 57 R Squared = 0.100 (Adjusted R Squared = 0.050) Table B 35. Two Way Factorial ANOVA for LMand I2 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 2.137 a 3 0.712 3.621 0.01749 Intercept 2482.62 1 2482.62 12616.504 0.00 TimePeriod 1.588 1 1.588 8.072 0.01 SexCombined 0.147 1 0.147 0.747 0.39 TimePeriod SexCombined 0.032 1 0.032 0.161 0.69 Error 12.987 66 0.197 Total 2883.989 70 Corrected Total 15.125 69 R Squared = 0.141 (Adjusted R Squared = 0.102)

PAGE 230

230 Table B 36. Two Way Factorial ANOVA for LMand I2 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 1.000a 3 0.333 2.778 0.04775 Intercept 2933.052 1 2933.052 24446.84 0.00 TimePeriod 0.035 1 0.035 0.295 0.59 SexCombined 0.823 1 0.823 6.858 0.01 TimePeriod SexCombined 0.186 1 0.186 1.552 0.22 Error 8.158 68 0.12 Total 3256.129 72 Corrected Total 9.158 71 R Squared = 0.109 (Adjusted R Squared = 0.070) Table B 37. Two Way Factorial ANOVA for LMand C MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 7.250 a 3 2.417 9.571 0.00002 a Intercept 4254.588 1 4254.588 16849.096 0.00 TimePeriod 0.472 1 0.472 1.869 0.18 SexCombined 6.667 1 6.667 26.402 0.00 b TimePeriod SexCombined 0.53 1 0.53 2.101 0.15 Error 20.453 81 0.253 Total 4524.127 85 Corrected Total 27.704 84 R Squared = 0.262 (Adjusted R Squared = 0.234) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 231

231 Table B 38. Two Way Factorial ANOVA for LMand C BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 8.290 a 3 2.763 13.657 0.00000 a Intercept 5498.925 1 5498.925 27176.031 0.00 TimePeriod 1.806 1 1.806 8.926 0.00 b SexCombined 6.223 1 6.223 30.752 0.00 b TimePeriod SexCombined 0.35 1 0.35 1.731 0.19 Error 17.199 85 0.202 Total 5797.582 89 Corrected Total 25.49 88 R Squared = 0.325 (Adjusted R Squared = 0.301) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 232

232 Table B 39. Two Way Factorial ANOVA for LMand P3 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 7.152 a 3 2.384 14.168 0.00000 a Intercept 5576.195 1 5576.195 33138.75 0.00 TimePeriod 3.141 1 3.141 18.665 0.00 b SexCombined 3.193 1 3.193 18.977 0.00 c TimePeriod SexCombined 0.978 1 0.978 5.81 0.02 b Error 15.481 92 0.168 Total 5948.445 96 Corrected Total 22.633 95 R Squared = 0.316 (Adjusted R Squared = 0.294) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level Table B 40. Two Way Factorial ANOVA for LMand P3 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.386 a 3 1.129 4.522 0.00530 Intercept 7101.785 1 7101.785 28448.67 0.00 TimePeriod 1.873 1 1.873 7.501 0.01 SexCombined 1.085 1 1.085 4.348 0.04 TimePeriod SexCombined 0.58 1 0.58 2.325 0.13 Error 22.717 91 0.25 Total 7498.103 95 Corrected Total 26.103 94 R Squared = 0.130 (Adjusted R Squared = 0.101)

PAGE 233

233 Table B 41. Two Way Factorial ANOVA for LMand P4 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 5.820 a 3 1.94 10.709 0.00000 a Intercept 5933.765 1 5933.765 32751.291 0.00 TimePeriod 1.424 1 1.424 7.861 0.01 b SexCombined 3.369 1 3.369 18.594 0.00 b TimePeriod SexCombined 0.891 1 0.891 4.917 0.03 c Error 17.031 94 0.181 Total 6216.696 98 Corrected Total 22.851 97 R Squared = 0.255 (Adjusted R Squared = 0.231) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 234

234 Table B 42. Two Way Factorial ANOVA for LMand P4 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 6.441 a 3 2.147 8.575 0.00004 a Intercept 7381.609 1 7381.609 29484.726 0.00 TimePeriod 2.376 1 2.376 9.489 0.00 b SexCombined 3.874 1 3.874 15.473 0.00 b TimePeriod SexCombined 0.456 1 0.456 1.82 0.18 Error 23.033 92 0.25 Total 7816.277 96 Corrected Total 29.473 95 R Squared = 0.219 (Adjusted R Squared = 0.193) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 235

235 Table B 43. Two Way Factorial ANOVA for LMand M1 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 4.344 a 3 1.448 6.442 0.00061 a Intercept 11906.606 1 11906.606 52971.467 0.00 TimePeriod 0.538 1 0.538 2.392 0.13 SexCombined 3.361 1 3.361 14.951 0.00 b TimePeriod SexCombined 0.787 1 0.787 3.501 0.07 Error 16.858 75 0.225 Total 12364.952 79 Corrected Total 21.202 78 R Squared = 0.205 (Adjusted R Squared = 0.173) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level Table B 44. Two Way Factorial ANOVA for LMand M1 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 6.261 a 3 2.087 11.262 0.00000 a Intercept 10230.494 1 10230.494 55205.338 0.00 TimePeriod 1.255 1 1.255 6.774 0.01 b SexCombined 4.932 1 4.932 26.615 0.00 b TimePeriod SexCombined 0.348 1 0.348 1.876 0.17 Error 16.308 88 0.185 Total 10773.979 92 Corrected Total 22.569 91 R Squared = 0.277 (Adjusted R Squared = 0.253) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 236

236 Table B 45. Two Way Factorial ANOVA for LMand M2 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 5.804 a 3 1.935 3.727 0.01461 Intercept 11366.565 1 11366.565 21895.661 0.00 TimePeriod 0.875 1 0.875 1.685 0.20 SexCombined 2.399 1 2.399 4.621 0.03 TimePeriod SexCombined 1.841 1 1.841 3.546 0.06 Error 41.011 79 0.519 Total 11757.644 83 Corrected Total 46.815 82 R Squared = 0.124 (Adjusted R Squared = 0.091) Table B 46. Two Way Factorial ANOVA for LMand M2 BL. Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 16.986 a 3 5.662 13.04 0.00000 a Intercept 9889.247 1 9889.247 22776.433 0.00 TimePeriod 9.171 1 9.171 21.123 0.00 b SexCombined 6.205 1 6.205 14.291 0.00 b TimePeriod SexCombined 0.686 1 0.686 1.58 0.21 Error 37.34 86 0.434 Total 10185.178 90 Corrected Total 54.326 89 R Squared = 0.313 (Adjusted R Squared = 0.289) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level

PAGE 237

237 Table B 47. Two Way Factorial ANOVA for LMand M3 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 14.361 a 3 4.787 5.102 0.00430 Intercept 4826.761 1 4826.761 5144.283 0.00 TimePeriod 11.881 1 11.881 12.662 0.00 SexCombined 1.593 1 1.593 1.698 0.20 TimePeriod SexCombined 0.002 1 0.002 0.002 0.97 Error 38.469 41 0.938 Total 5423.724 45 Corrected Total 52.831 44 R Squared = 0.272 (Adjusted R Squared = 0.219)

PAGE 238

238 Table B 48. Two Way Factorial ANOVA for LMand M3 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 10.903 a 3 3.634 10.974 0.00001 a Intercept 5020.924 1 5020.924 15161.674 0.00 TimePeriod 7.384 1 7.384 22.297 0.00 b SexCombined 2.949 1 2.949 8.905 0.00 b TimePeriod SexCombined 0.004 1 0.004 0.011 0.92 Error 15.233 46 0.331 Total 5287.874 50 Corrected Total 26.136 49 R Squared = 0.417 (Adjusted R Squared = 0.379) a. Significant at an adjusted 0.00078 Bonferonni alpha level b. Significant at a 0.05 alpha level Table B 49. Two Way Factorial ANOVA for RMand I1 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 2.074a 3 0.691 6.384 0.00088 Intercept 1525.705 1 1525.705 14086.23 0.00 TimePeriod 1.681 1 1.681 15.524 0.00 SexCombined 0.011 1 0.011 0.105 0.75 TimePeriod SexCombined 0.07 1 0.07 0.644 0.43 Error 5.849 54 0.108 Total 1943.646 58 Corrected Total 7.923 57 R Squared = 0.262 (Adjusted R Squared = 0.221)

PAGE 239

239 Table B 50. Two Way Factorial ANOVA for RMand I1 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 0.313a 3 0.104 1.015 0.39384 Intercept 1800.943 1 1800.943 17491.938 0.00 TimePeriod 0.215 1 0.215 2.088 0.15 SexCombined 0.064 1 0.064 0.622 0.43 TimePeriod SexCombined 0.05 1 0.05 0.49 0.49 Error 5.251 51 0.103 Total 2165.768 55 Corrected Total 5.564 54 R Squared = 0.056 (Adjusted R Squared = 0.001) Table B 51. Two Way Factorial ANOVA for RMand I2 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 1.458a 3 0.486 3.851 0.01345 Intercept 2626.457 1 2626.457 20807.176 0.00 TimePeriod 1.09 1 1.09 8.632 0.00 SexCombined 0.291 1 0.291 2.304 0.13 TimePeriod SexCombined 0.021 1 0.021 0.165 0.69 Error 8.079 64 0.126 Total 2872.665 68 Corrected Total 9.537 67 R Squared = 0.153 (Adjusted R Squared = 0.113)

PAGE 240

240 Table B 52. Two Way Factorial ANOVA for RMand I2 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 0.696a 3 0.232 1.416 0.24585 Intercept 3056.193 1 3056.193 18653.853 0.00 TimePeriod 0.617 1 0.617 3.764 0.06 SexCombined 0.015 1 0.015 0.091 0.76 TimePeriod SexCombined 0.05 1 0.05 0.305 0.58 Error 10.977 67 0.164 Total 3239.594 71 Corrected Total 11.673 70 R Squared = 0.060 (Adjusted R Squared = 0.018) Table B 53. Two Way Factorial ANOVA for RMand C MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 4.763a 3 1.588 11.801 0.00000 a Intercept 4121.565 1 4121.565 30632.515 0.00 TimePeriod 0.424 1 0.424 3.151 0.08 SexCombined 4.405 1 4.405 32.741 0.00 b TimePeriod SexCombined 0.221 1 0.221 1.645 0.20 Error 10.36 77 0.135 Total 4308.722 81 Corrected Total 15.124 80 R Squared = 0.315 (Adjusted R Squared = 0.288) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 241

241 Table B 54. Two Way Factorial ANOVA for RMand C BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 5.060a 3 1.687 8.182 0.00008 a Intercept 5160.1 1 5160.1 25033.04 0.00 TimePeriod 0.379 1 0.379 1.84 0.18 SexCombined 3.81 1 3.81 18.481 0.00 b TimePeriod SexCombined 1.459 1 1.459 7.079 0.01 b Error 16.078 78 0.206 Total 5378.017 82 Corrected Total 21.138 81 R Squared = 0.239 (Adjusted R Squared = 0.210) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 242

242 Table B 55. Two Way Factorial ANOVA for RMand P3 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 6.317a 3 2.106 10.207 0.00001 a Intercept 5266.893 1 5266.893 25532.716 0.00 TimePeriod 2.49 1 2.49 12.073 0.00 b SexCombined 2.508 1 2.508 12.159 0.00 b TimePeriod SexCombined 1.202 1 1.202 5.826 0.02c Error 17.74 86 0.206 Total 5464.768 90 Corrected Total 24.057 89 R Squared = 0.263 (Adjusted R Squared = 0.237) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level Table B 56. Two Way Factorial ANOVA for RMand P3 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 3.900a 3 1.3 3.774 0.01344 Intercept 6819.722 1 6819.722 19797.856 0.00 TimePeriod 0.935 1 0.935 2.715 0.10 SexCombined 0.609 1 0.609 1.767 0.19 TimePeriod SexCombined 2.201 1 2.201 6.39 0.01 Error 29.969 87 0.344 Total 7183.181 91 Corrected Total 33.869 90 R Squared = 0.115 (Adjusted R Squared = 0.085)

PAGE 243

243 Table B 57. Two Way Factorial ANOVA for RMand P4 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 7.144a 3 2.381 15.137 0.00000 a Intercept 5637.778 1 5637.778 35835.681 0.00 TimePeriod 2.59 1 2.59 16.462 0.00 b SexCombined 4.774 1 4.774 30.344 0.00 b TimePeriod SexCombined 0.032 1 0.032 0.204 0.65 Error 13.844 88 0.157 Total 5810.048 92 Corrected Total 20.989 91 R Squared = 0.340 (Adjusted R Squared = 0.318) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 244

244 Table B 58. Two Way Factorial ANOVA for RMand P4 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 8.684a 5 1.736 5.319 0.00018 a Intercept 10197.624 1 10197.624 31229.701 0.00 TimePeriod 3.274 1 3.274 10.028 0.00 b SexCombined 5.262 2 2.631 8.058 0.00 b TimePeriod SexCombined 0.467 2 0.233 0.715 0.49 Error 41.470 127 0.326 Total 10854.678 133 Corrected Total 50.155 132 R Squared = 0.173 (Adjusted R Squared = 0.141) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 245

245 Table B 59. Two Way Factorial ANOVA for RMand M1 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 8.627a 5 1.725 5.646 0.00000 a Intercept 20146.669 1 20146.669 65924.701 0.00 TimePeriod 3.391 1 3.391 11.095 0.00 b SexCombined 5.407 2 2.704 8.847 0.00 b TimePeriod SexCombined 0.107 2 0.054 0.176 0.84 Error 38.811 127 0.306 Total 20649.328 133 Corrected Total 47.439 132 R Squared = 0.182 (Adjusted R Squared = 0.150) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level

PAGE 246

246 Table B 60. Two Way Factorial ANOVA for RMand M1 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 8.435a 3 2.812 15.045 0.00000 a Intercept 11000.381 1 11000.381 58862.344 0.00 TimePeriod 2.322 1 2.322 12.427 0.00 b SexCombined 6.328 1 6.328 33.859 0.00 b TimePeriod SexCombined 0 1 0 0.002 0.96 Error 16.633 89 0.187 Total 11058.707 93 Corrected Total 25.068 92 R Squared = 0.336 (Adjusted R Squared = 0.314) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level Table B 61. Two Way Factorial ANOVA for RMand M2 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 8.581a 3 2.86 4.426 0.00645 Intercept 9745.587 1 9745.587 15078.329 0.00 TimePeriod 2.103 1 2.103 3.254 0.08 SexCombined 1.378 1 1.378 2.132 0.15 TimePeriod SexCombined 2.81 1 2.81 4.348 0.04 Error 47.828 74 0.646 Total 10819.881 78 Corrected Total 56.41 77 R Squared = 0.152 (Adjusted R Squared = 0.118)

PAGE 247

247 Table B 62. Two Way Factorial ANOVA for RMand M2 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 10.469a 3 3.49 13.665 0.00000 a Intercept 9750.763 1 9750.763 38183.968 0.00 TimePeriod 6.657 1 6.657 26.068 0.00 b SexCombined 3.592 1 3.592 14.067 0.00 b TimePeriod SexCombined 0.216 1 0.216 0.844 0.36 Error 21.961 86 0.255 Total 10225.763 90 Corrected Total 32.43 89 R Squared = 0.323 (Adjusted R Squared = 0.299) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha leve

PAGE 248

248 Table B 63. Two Way Factorial ANOVA for RMand M3 MD Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 18.254a 3 6.085 16.108 0.00000 a Intercept 5082.833 1 5082.833 13456.099 0.00 TimePeriod 16.495 1 16.495 43.668 0.00 b SexCombined 2.826 1 2.826 7.48 0.01 b TimePeriod SexCombined 0.079 1 0.079 0.21 0.65 Error 14.354 38 0.378 Total 5234.769 42 Corrected Total 32.607 41 R Squared = 0.560 (Adjusted R Squared = 0.525) a Significant at an adjusted 0.00078 Bonferonni alpha level b Significant at a 0.05 alpha level Table B 64. Two Way Factorial ANOVA for RMand M3 BL Source Type III Sum of Squares df Mean Square F Sig. Corrected Model 14.067a 3 4.689 11.696 0.00001 Intercept 4651.429 1 4651.429 11602.26 0.00 TimePeriod 13.714 1 13.714 34.207 0.00 SexCombined 0.779 1 0.779 1.943 0.17 TimePeriod SexCombined 0.046 1 0.046 0.116 0.74 Error 16.838 42 0.401 Total 4925.682 46 Corrected Total 30.905 45 R Squared = 0.455 (Adjusted R Squared = 0.416)

PAGE 249

249 LIST OF REFERENCES Salkind, ed., Encyclopedia of Measurment and Statistics. Thousand Oaks: Sage. p 1 9. Ambrose SH, Butler BM, Hanson DB, Hunter Anderson RL, Krueger HW. 1997. Stable isotopic analysis of human diet in the Marianas Archipelago, Western Pacific. Am J of Phys Anthropol 104:343 361. Amesbury JR. 1999. Changes in species composition of archaeological marine shell assemblages in Guam. Micronesica 31:347 366. Amesbury JR. 2007. Mollusk collecting and environmental change during the prehistoric period in the Mariana Islands. Coral Reefs 26:947 95 8. Amesbury JR, Hunter Anderson RL, Moore DR. 1991. An archaeological study of the San Antonio Burial Trench and a report on the archaeological monitoring of road construction along Marine Drive between Rts. 8 and 4, Agana, Guam. A Report Prepared for Gove rnment of Guam Public Works, Division of Highway Engineering, and Black Construction Corporation, Agana, Guam. Amesbury JR, Hunter Anderson RL. 2003. Review of archaeological and historical data concerning reef fishing in the U.S. Flag Islands of Micrones ia: Guam and the Northern Mariana Islands. Prepared for Western Pacific Regional Fishery Management Council, Honolulu. Micronesian Archaeological Research Services, Guam. Amesbury JR, Hunter Anderson RL. 2008. An analysis of archaeological and historical d ata on fisheries for the pelagic species in Guam and the Northern Marianas Islands. A report prepared for Pelagic Fisheries Research Program, Joint Institute for Marine and Atmospheric Research, School of Ocean and Earth Science and Technology, University Services, Guam. Amesbury JR, Moore DR, Hunter Anderson RL. 1996. Cultural adaptations and late Holocene sea level change in the Marianas: recent excavations at Chalan Piao, Saipan, Micronesia. Indo P acific Prehistory Association Bulletin 15:53 69. Anderson AJ. 1991. The chronology of colonization in New Zealand. Antiquity. 65:767 795. Anderson AJ. 1996. Adaptive voyaging and subsistence strategies in the early settlement of East Polynesia. In T Akazaw a and EJE Szathmary, eds.: Prehistoric Mongoloid dispersals. Oxford: Oxford University Press. p 359 373.

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279 BIOGRAPHICAL SKETCH Nicolette Maria Luney Parr graduated with highest honors from the University of Florida in 20 eligion. Ms. Indianapolis, were she completed a Master of Science degree in h uman b iology. While at the University of Indianapolis, Ms. Parr served as the Laboratory Coordinator for the Archaeology and Forensics Laboratory, performed recovery of human remains from cri me scenes, and assisted with forensic casework and served as a teaching assistant for anatomy, skeletal biology, and dental anthropology courses. After an internship at the University of Pretoria in South Africa, Ms. Parr completed her m determining ancestry from the mandible. Ms. Parr entered the a nthropology Ph.D. program at the University of Florida in 2005, where she served as a graduate analyst at the C.A. Pound Human Identification Laboratory, was a teaching assistant for an introduc tory biological anthropology course, and served as in instructor for an introductory course on forensic anthropology and a laboratory based course in human osteology. She co authored a book entitled Bare Bones: An Introduction to Forensic Anthropology, wh ich is currently in its second edition. After completing coursework, Ms. Parr worked part time as the physical anthropologist for Garcia and Associates in Guam. Her work as an osteologist in Guam led to the culmination of this project. Ms. Parr received her Ph.D. from the University of Florida in August 2012. Upon graduation, M s. Parr was hired by the Joint Pow/MIA Accounting Command Central Identification Laboratory in Honolulu, Hawaii, where she works as a forensic anthropologist.