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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

A DIACHRONIC ASSESSMENT OF HEALTH AND …ufdcimages.uflib.ufl.edu/UF/E0/04/46/39/00001/PARR_N.pdf · Pre-Latte Period ... The Spanish Colonial Period (1521-1898 CE) ... The First

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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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© 2012 Nicolette Maria Luney Parr

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

encouragement

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ACKNOWLEDGMENTS

First and foremost I would like to thank my committee members, Mike Warren,

John Krigbaum, Dave Daegling, and David Steadman for your tireless 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 way. 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

provided to me by Patrick O’Day, Nicole Vernon, Mike Desilites, 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 good 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 many 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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Barcinas, Boyd Dixon, Jonn Peterson, Anne Stodder, Lawrence Cunningham, Lon

Bulgrin, Michael Pietrusewsky, Bruce 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, Allysha, 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 – here’s to ugly babies!

To Lulu, Kim, Viviana, Amber, and Leila: much of who I am today I owe to having

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, particularly 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 for teaching me

how to travel, eat good food, and to love life – 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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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

Biocultural 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 Environment ............................................................................................... 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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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 Population .................................................................................................. 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 Comparison of dental, craniofacial, and postcranial changes across

time ......................................................................................................... 99

Carious lesions ........................................................................................ 101

Mechanism for Dental Reduction .......................................................................... 102 Probable Mutation Effect ................................................................................ 102 Increasing Population Density Effect .............................................................. 103 Selective Compromise Effect ......................................................................... 104 Masticatory Functional Hypothesis ................................................................. 105

Conclusions .......................................................................................................... 105

5 DEVELOPMENTAL INSTABILITY: ENAMEL HYPOPLASIAS ............................. 110

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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 Materials 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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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 ................................................................... 84

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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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 Descriptive statistics of cross-sectional area by time period and sex ............... 190

A-6 Group comparisons of dental measurements ................................................... 193

A-7 Kolmogorov-Smirnova Test for normality .......................................................... 200

A-8 Levene's Test of Homogeneity of Variance based on the mean ....................... 204

B-1 Two-Way Factorial 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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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 RMax 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 RMax 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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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 for 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 for 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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B-64 Two-Way Factorial ANOVA for RMand M3 BL ................................................. 248

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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 Reconstructed Latte set from Reinman’s Talofofo River Valley Site. .................. 73

2-4 Examples of Pre-Latte Period pottery ................................................................. 74

2-5 Island of Guam depicting Latte Set density and distribution, following Hornbostel’s original 1920s survey ..................................................................... 75

2-6 Examples of Latte Period pottery ........................................................................ 76

5-1 Single 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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LIST OF ABBREVIATIONS

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/m3 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 maxillary second molar

LMax M3 Left maxillary third molar

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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 Right 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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Abstract of Dissertation Presented 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. 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 in size from the Pre-Latte to Latte periods.

Increased reliance on starchy crops would have led to selection for smaller dentition to

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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 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 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, which 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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CHAPTER 1 INTRODUCTION

In these Proes1…these people sail in those seas from Island to Island for several hundred Leagues, the Sun serving them for a compass by day and the Moon and Stars by night. When this comes to be prov’d we Shall be no longer at a loss to know how the Islands lying those Seas came to be people’d.

—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 the last migration into uncharted territory (Spate, 1979) and is a

remarkable feat of ‘maritime discovery, colonization, and adaptation’ (Russell, 1998:

67).

The 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 islands are more likely

to undergo more noticeable phenotypic modification due to environmental changes and

subsequent selective pressures (Houghton, 1991a). Additionally, the diversity of

climate and environment throughout the Pacific Islands provide “the possibility of

investigating on a broad comparative scale the effect of environment on phenotypic

development” (Goodenough, 1957: 154).

1Cook is referring to proas, maritime sailing vessels used in the Pacific.

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 written

about where the Polynesians came from but nobody cares a straw 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, the

paucity of skeletal research in Guam still remains, regardless of evidence that places

the earliest colonization of Remote Oceania in the Marianas Islands (Carson, 2008;

Clarke et al., 2010; Carson, 2010).

Most 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 thorough 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 (Hanson 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, 1992; 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 restricted to a single published volume (vol. 104, 1997)

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 Douglas, 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 Rothschild, 1995) or in small Pacific-

based publications (e.g. Underwood, 1973, 1976; Houghton, 1991b; 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 teeth 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 important 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 associated with subsistence patterns

and health status in prehistoric populations (Armegalos and Van Gerven, 2003).

22

Central to the biocultural analysis of human health and disease is Goodman and

Armelagos’ (1989) model for investigating stress in prehistoric skeletal 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

uses integrative thinking to explain the adaptation process as a ‘mechanism for

responding to environmental stressors’ (McElroy, 1990: 246), while creating testable

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 evolutionary processes and

adaptations in prehistory.

In the 1970s 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 associated 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 in other parts of the world, stems

from the emergence of Binford’s New Archaeology (1962), with its focus on a synthetic

approach to population-based, ecological research (Buikstra, 1997; Buikstra and Beck,

2006). Bioarchaeology can be defined as a contextual study of human skeletal remains

from archaeological contexts, which integrates biology, culture, and environment to

better understand health, disease, and demography in prehistoric human populations

23

(Buikstra; 1977; Larsen, 1997; Buikstra and Beck, 2006). 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 archaeological 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,

1977). Blakely (1977b: 1) states that ‘humans survive not through cultural adaptation

nor biological adaptation, but through biocultural adaptation’ (emphasis his). Recent

studies emphasize a biocultural approach by combining archaeological and osteological

research to answer significant questions about adaptation and the forces 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 Japan (Temple, 2007; Temple and

Larsen, 2007; Oxenham and Matsumura, 2008; Temple, 2010); ecological and

demographic pressure in the Nile Valley (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 twentieth 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

24

development) due to a ‘state of stress’ (Goodman et al., 1988). Selye (1973: 692)

defines stress as ‘the nonspecific response of the body to any demand made upon it’

where alterations of the environment alters the normal and steady state of an organism.

Central to the Selyean concept of stress is the general adaptation syndrome, a defense

mechanism in which homeostatic mechanisms are activated to alleviate a long and

continued stress event (Selye, 1956). Classic laboratory studies showed that increased

exposure to stressors (i.e., noise, cold, and heat) lead to developmental disturbances in

rodents (Siegel and Smookler 1973; Siegel and Doyle 1975a,b; Siegel et al. 1977;

Doyle et al. 1977; Sciulli 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 affects 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).

In Goodman and Armelagos’ (1984) seminal volume Paleopathology at the Origins

of Agriculture, the authors present a model for stress applicable 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).

25

The model 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 stressors will reach the individual or population.

Adaptation to a stressor depends on each individual’s host resistance. No biological

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, 1997; Goodman and Martin,

2002). Soft tissue responds more quickly to stress and disease than the skeleton.

Therefore, 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 leprosy, leave diagnostic lesions on the skeleton,

many other pathogens elicit the same response for a number of given stressors

26

(Goodman et al., 1998). For example, periostitis is one of the most 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, such as influenza and other viruses that may result in

decreased health and in some cases death, leave no evidence in the skeleton, which

confounds 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 that stemmed from Goodman and Armelagos’ (1984) volume

regarding population health in the transition to agriculture. The authors identified key

conceptual problems that complicate 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 process 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 example, 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 Benderlioglu, 2006). Nonetheless, since its publications,

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 joint 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

disease need to be evaluated with “other lines of evidence, including subsistence and

settlement, environmental context, cultural context, and population structure.” Thus, an

analysis of the cultural 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-Anderson 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

28

environmental modifications will be accompanied by biological ones. For example,

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 hypoplasias, 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 investigation into biological processes that can lead to biological

change in human 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

“invention, environmental limitation or encouragement, founder influence on small island

cultures, drift through isolation or communication decline, and diffusion/borrowing.”

These temporal 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 resource management litigation.2 This sample

includes both the Pre-Latte (n = 103) and Latte periods (n = 112) and is located on the

west coast of Guam, in the Western Pacific, in Northern Tumon Bay at the Naton Beach

location (Figures 1-2). Four 14C dates were obtained from conus shell bead necklaces

associated with the earliest burials and range from 2790 to 2330 BP (DeFant, 2008).

2The 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.

29

This sample represents the largest Pre-Latte skeletal assemblage to be excavated on

Guam and one of the oldest and largest in the remote Western Pacific. Dates from the

Chelechol ra Orrak cemetery in Palau (also in the Western Pacific) range from 3000 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 Marianas

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 if biological change has occurred

following cultural shifts. The hypotheses are as follows:

Hn1: There is no significant 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, was 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. Additionally, subsistence adaptations are analyzed to

30

determine the impact of agricultural intensification on the oral health of the Chamorro. If

dental reduction, linear enamel hypoplasias, and carious lesion differences are found,

this study will delineate causative factors associated with these changes through further

analysis of the archaeological record. In conclusion, an analysis of dental changes in

the prehistoric Chamorro will elucidate 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 chapters. 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 skeletal remains. The second chapter

details the natural and cultural history 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 Chapter 3, the sample materials are

described with a focus on taphonomic biases and population demography. The fourth,

fifth, and six chapters report the background, data collection methods, including

statistical procedures, results, and discussion of the research. 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 lifestyles of the prehistoric Chamorro.

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)

32

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

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

Hawai’i (Thompson, 1932) (Figure 2-1). The Marianas lie west of the 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 Saipan) 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 forming 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-Adelup 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 Guam is necessary to

elucidate the complexity of human cultural adaptations over time, particularly when

evaluating the archaeological record. This overview provides insight into environmental

factors that may have led to biocultural changes in the Chamorro.

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Biogeographical Divides

In the 17th century, the French voyager, Dumont d’Urville (1832) created a tripartite

classification system for the peoples and islands of the Pacific (Melanesia, Micronesia,

and Polynesia) based on typological physical characteristics of incredibly diverse and

heterogeneous peoples (Green 1991; Kirch, 2000; 2010). d’Urville’s partitions have no

utility in terms of biological, cultural, and historical processes, save for Polynesia which

has proven more homogeneous, both culturally 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 populations 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 biological standpoint, d’Urville’s classification

scheme is geographically significant. Melanesia, ‘the dark islands’, contains the largest

land masses in the Pacific and includes New Guinea, the Solomons, Vanuatu

(previously known as New Hebrides), New Caledonia, and Fiji (d’Urville,1832; Kirch,

2010). Micronesia, ‘the little islands’, is composed of approximately 21,000 islands and

encompasses five main archipelagos: Palau (Belau), the Marianas, the Carolines, the

Marshalls, and Kiribati (formerly the Gilbert Islands) (d’Urville, 1832; Kirch 2010).

Despite the vast number of islands, the totality of Micronesia’s land mass is only ~2,700

km2, with Guam being the largest of the islands (582 km2) (Kirch, 2010). Lastly,

Polynesia, ‘many islands’, encorporates the far eastern Pacific islands, including

35

Hawai’i, Rapa Nui (Easter Island), Tahiti, and New Zealand (d’Urville, 1832; Kirch

2010).

Green (1991) suggests replacing d’Urville’s terms with 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 use of Green’s

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 activity (Bath, 1986; Kurashina and Clayshulte,

1983; Dickinson, 2000; 2003; Carson, 2011). During the mid-Holocene highstand

(~6000 and 4000 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

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 arrival 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, stable 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 associated 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-Latte sites are

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). Additionally,

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, data 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 increased diversity in habitat (Nunn, 1999). Since the end of the HCO

‘cooling, sea-level fall and, in places, a fall in precipitation and loss of biodiversity’

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 from AD 750 to 1300 is known as the ‘AD 1300 Event’,

where rapid cooling temperatures, decline in sea levels, and increased storminess

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

‘most rapid period of climate change to have occurred within the past several millennia’

(Nunn, 2007: 1) and is associated with societal disruption, subsistence change, and

movement to inland habitation 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, associated 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 paleoclimate 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. Likewise, 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 areas 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.

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 predation, 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 introduced

species: the mangrove monitor lizard (Varanus indicus), introduced around 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 extinct.

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 historic 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 remains were found in the prehistoric 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 twenty-four

documented) extant species of birds (herons - two species, swifts, starlings, and crows)

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).

Settlement 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 the 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, including

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 there 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 archaeological, 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 second settlement event began around 3,500 BP with the rapid

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 archaeological 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 three 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 languages, 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. Terrell, 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

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 family (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

Biologically, 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, 1990a; 1990b; Houghton, 1991b; 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 proto-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,

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 the 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, the 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 groups (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 homeland 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

Polynesian 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 Australians 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

44

closer affinity with Southeast 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 pattern 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 mid-Holocene in the Pacific Ocean resulted in the

appearance of attractive coastal environments, which subsequently allowed for rapid

dispersal and initial human colonization in the western parts of Remote Oceania

(Dickinson, 2001; Carson, 2011). Most researchers 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 analysis of pottery, language, and

DNA (Hanson and Butler, 1997; Lum and Cann, 1998; 2000; Callaghan and Fitzpatrick,

2008). Osborne (1958) posited a ‘stepping-stone’ colonization model of Micronesia

through Palau and Yap, however, this theory has not been supported 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

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 and 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 that 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, 1965; 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; Pawley, 2002). The

Chamorro language has a distinctly separate geographic source from the Lapita Proto-

Oceanic group, and historical linguistics suggest direct colonization from Island

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; Clark 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 settlement 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, similar 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

47

Craib,1987; Carson, 2008; Clark et al., 2010). Carson’s radiocarbon findings (2008)

pushes occupation even further back to 3600 to 3420 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 from 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 with

Achugao Incised pottery has been dated more securely to 3690 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 and is

associated with very thin, redware pottery. The slightly younger site, with thicker and

coarser pottery, dates to 3056 to 2842 BP (Carson, 2010). This site illustrates

chronological change in pottery sequence over time with solid radiocarbon dates.

Decorated pottery in the Mariana Islands is most similar to ‘red-slipped, circle- and

punctate-stamped pottery from several sites in the Cagayan Valley on Luzon’ in the

Philippines (Hung et al., 2010: 913). The Neolithic and Iron age site of Nagsabaran,

which is located within the Cagayan 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

Unai 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.

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

other 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 (craniofacial

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 clustered 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 differences 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

49

Near Oceania 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 diversity of its group I sequence. These data are further evidence

against the ‘stepping-stone’ hypothesis of settlement of the Marianas through Palau and

Yap.

Bioarchaeology

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 suggesting 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).

Pietrusewsky’s research (1990a; 1990b; 1994; 1995; 1996; 2000; 2005; 2008)

over the past four decades confirms Howell’s findings in regards to the position of

Micronesia populations in comparison to the rest of Remote Oceania, Near Oceania,

Australia, Island Southeast Asia, mainland Southeast Asia, east Asia, and North Asia.

Multivariate craniometric analyses (27 measurements in 63 cranial 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 D2 shows that the Polynesian and Guam grouping

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) analyzed 22 non-metric cranial traits from the Marianas

populations and compared them with twelve other groups across the Pacific, mainland

Southeast Asia, East Asia, mainland Asia, and North America. Like Pietruwsewsky’s

and Howells’ results, Ishida and Dodo cluster the Chamorro and Polynesian populations

with Southeast Asian and East Asian populations.

One problem with the interpretation of Pietrusewsky’s and Ishida and Dodo’s

findings on Chamorro settlement patterns is that their samples are derived from the

Hornbostel-Thompson Collection1 of skeletal material from the Mariana Islands

originally housed in the Bishop Museum in Hawai’i. These series of skeletons were

collected by Hans Hornbostel and J.C. Thompson in the 1920s 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 undertakings 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

Guam Museum’s collections.

51

Unfortunately, as discussed above, Pre-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 Rainbird (2003: 85) notes, a voyage from the

Philippines to the Marianas ‘would constitute the longest sea-crossing undertaken by

that time in human history’. Thus, the peopling of the Marianas is not only interesting in

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

the Chalan Piao site on the island of Saipan, located approximately 220 km north of

Guam (Figure 2-1). Subsequently, recalibration of Spoehr’s dates yielded a more

recent age dating to 1,700 years later (Cloud et al., 1956). Nonetheless, the majority of

the earliest radiocarbon dates across the Marianas cluster around 3500 to 3000 BP,

therefore substantiating Spoehr’s claim and making Guam the earliest colonized island

of Micronesia and Remote Oceania (Reinman 1977; Kurashina and Clayshulte 1983;

Butler 1995; 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

52

appearance of charcoal particles in the Laguas paleocore, however, this date has not

been supported by archaeological 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.

Spoehr (1957) proposed the chronology of the Marianas dividing the prehistory

into two 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 pillars, 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). Spoehr’s division also distinguished between ceramic types with the Pre-Latte

Marianas Redware and the Latte Period Marianas Plainware.

Moore’s (1983) chronological sequence builds upon Sphoer’s original scheme and

is based on analyses of ceramic sequences from Tarague Beach in Northwestern

Guam. Her classification was later refined into the current standard which has replaced

the term ‘Pre-Latte’ with ‘Unai’, which means beach in Chamorro, and ‘Transitional’ with

‘Huyong’, which translates to ‘going out’ (Moore, 2002) (Table 2-1). The following

53

overview of the chronological sequence is based on a combination of Spoehr (1957)

and Moore’s (2002) divisions.

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 period (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 deposits 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-Anderson 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 northern 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 breakage patterns were not conducted

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, dating 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, lizard, 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 have 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 Marianas during the Holocene. Thus, presence of T. gigas

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 378.37 grams per

cubic meter) in comparison to the Latte layer (187.20 g/m3).

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 common 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 increases to 43% in the Latte period.

Emperorfish (Lethrinidae) and wrasses/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 15N values in 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

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 practices 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 noted 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 among 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 phase (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 exteriors

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 remains, 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 the north

(Leidemann, 1980; Bath, 1986; Dilli et al., 1998; Carson 2010).

The Middle Unai period (3000-2500 BP)

The Middle Unai Phase spans the next 500 years following the Early Unai Phase.

Archaeological assemblages are very similar to the Early Unai Phase, 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 slipped,

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), with 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.

58

Ray (1981) reports another pottery type characteristic 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 impressions, 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 (2500-2400 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, 1983). 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).

59

Quartz inclusions 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 primarily 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 shoreline 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 incurvate 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 purposes, 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 and 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

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

Guam’s prehistory comes from the better preserved Latte period sites (Hunter-Anderson

and Butler, 1991). While ideal settlement 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 (Tomonari-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 function and significance of latte sets

as specialized structures attributed as ceremonial centers, meeting houses, men’s

houses, canoe sheds, and residential structures for the elite (Thompson, 1940;

Thompson, 1945; Spoer, 1957; Reinman, 1977). A survey of cultural 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 previously thought,

specialized to one function. Artifacts associated with cooking, food preparation, tool

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manufacture and maintenance, fishing, and warfare were found homogenously in all the

latte sets analyzed (Table 2-2). Additionally, human burials are uniformly 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 along 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). Archaeological 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

symbols of corporate landholdings and a kin group’s claim to land through linage

(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 and

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

gripping/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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residue 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.

They were likely used to ‘grind, pulverize, and de-husk plant products, such as rice,

cycads, and arrowroot tubers’ (Moore, 2005:100). The presence of scrapers, knives

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, 2005).

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 identifies 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; Crowther 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 years 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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Pollen and phytolyth analysis: 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 and betel palm, in

addition to arrowroot, rice, bananas, canna, and Job’s tears (Hunter-Anderson, 1994;

Pearsall and Collins 2000; Collins and Pearsall, 2001a; Collins and Pearsall 2001b).

Lastly, analysis of charred wood samples from southwestern Guam identified 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, research 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 13C values indicating a significant dependence on terrestrial

foods. The 15N 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 proximity to the ocean.

64

Quinn’s analysis (1990) of two individuals from different locations in Saipan

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 15N value)

and 40% (based on mean 13C value) of the diet in Saipan, while these values 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’s (1997) data for Saipan reveal large discrepancies

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 higher estimate of 60%. Nonetheless, the very high apatite carbonate

13C values suggest a large component of the Saipan diet came from C4 or marine

plants with very low protein component, such as sugar cane and seaweeds. Small

amounts of marine foods, likely 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 15N values suggesting exploitation of pelagic fish,

while others have negative 13C and 15N values indicative of a diet dominated by

terrestrial foods, suggesting differential access to resources between individuals.

65

Likewise, Ambrose and colleagues (1997) found that marine resources contributed

between 21% and 40% to the diet in Rota. High 15N values suggest a significant

contribution of pelagic fish to the diet in some individuals, while other individuals have

low 13C 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 15N and 13C values are similar to what is

seen in Rota, however, the 15N values in Guam are slightly lower than those from Rota.

This suggests that the diet was comprised primarily of terrestrial C3 resources, such as

rice, root crops, and vegetables, and some marine protein, but little reliance on

seaweeds or C4 plants (Ambrose et al., 1997).

Shellfish consumption: A transition from bivalve to gastropod consumption

occurs in the Latte period (Amesbury et al., 1996; Amesbury, 2007). Leidemann’s study

(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 concomitant 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 marks the beginning of the post-contact era as well as the first contact between

66

Europeans and any indigenous Micronesian populations (Alkire, 1977; Lévesque,

1995). The post-contact area is commonly divided into the Spanish Colonial, First

American, Japanese 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, culture, 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 or 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 missionaries 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

(Lévesque, 1995).

Juan Pobre, a Franciscan brother who lived 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. Taro, 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

20,000 but by the end of that century diminished to approximately 1,600 individuals due

67

to widespread epidemic disease and rebellion after European contact (Russell and

Fleming, 1990; Tomonari-Tuggle et al., 2005). The Spanish instituted the Reducción

which consolidated 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

importance 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 farming and raising livestock (Russell, 1988). The village of “Tumhan” is illustrated

on the 17th century map from the Jesuit Charles Le Gobien (1700), yet, according to

Kurishina et al. (1987; 1988), little activity is known to have occurred in the area during

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

(Burtchard 1991).

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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 was

not firmly set until the 1900s 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 7th attack on Pearl Harbor, Hawai’i, the Japanese attacked Guam. Two days

of bombing by Japanese air forces culminated with a three-pronged ground attack at

Tumon Bay, Agaña Bay, and Merizo, and the forced surrender of Guam to the

Japanese by the American governor (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 Chamorro reverted back to traditional practices

69

of fishing and agriculture, particularly in more remote areas (Russel et al., 1993;

Rogers, 1995).

With the intensification of the war and Japan’s decreasing control in the Pacific

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 was succeeded to the

Americans on July 27th, 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 Japanese 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, control 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

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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Table 2-1. Archaeological and historical chronological sequences of the Marianas 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 setsa

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

Figure 2-3. Reconstructed Latte set from Reinman’s Talofofo River Valley Site (Photo

by author)

74

Figure 2-4. Examples of Pre-Latte Period pottery. Photo taken at Lina’la’ Chamorro

Cultural Park Museum (Photo by author)

75

Figure 2-5. Island of Guam depicting Latte Set density and distribution, following

Hornbostel’s original 1920s survey (Map courtesy of Rad Smith/GANDA)

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Figure 2-6. Examples of Latte Period pottery. Photo taken at Lina’la’ Chamorro

Cultural Park Museum (Photo by author)

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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

determination of the burials’ age and cultural affiliation was based on the stratigraphic

location, associated artifacts, and radiocarbon dates (DeFant, 2008). Dating to roughly

2,500 BP, the Pre-Latte group represents 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 remains in the Latte period between AD 1000 to 1521.

A comprehensive osteological analysis was conducted by Ms. Cherie Walth, of

SWCA. All demographic information, as well as information on cranial and postcranial

metrics and non-dental pathological conditions, 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 unpublished manuscript to the Guam

Historic Preservation Office.

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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 (Dickinson, 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, changing 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 encountered (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 of Guam’s earliest prehistoric peoples.

Disturbances due to human activity also affect bone preservation, as

archaeological 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 that the Latte believed that their ancestors’ skeletal remains

possessed power, which could be called upon by descendants for supernatural

purposes (Thompson, 1940; Driver, 1983). Thus, skeletal remains were buried beneath

the Latte sets, as a means for the familial social unit to demonstrate their loyalty and

79

identify with a particular lineage as well as to show strength through close proximity to

their ancestors (Thompson, 1945; Graves, 1986).

Further, Latte 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

Franciscan priest, Fray Juan Pobre, reported: ‘the one thing for which they have high

regard are the skulls of the ancestors, especially those of their parents and

grandparents’ (Driver, 1983: 214), 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 1930s the Tumon coastline was altered

during the occupation of the Japanese who built commercial establishments and

extensive 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

recent 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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Sample Population

A total 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 undetermined 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

composed of adult individuals (n = 179). Only juveniles with permanent 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 (n = 4). Thirteen individuals were placed into the ‘adult’ category without any

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 determined 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

determined 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; 28.6%),

81

and lastly old-adults (n = 4; 3.6%). Eight individuals are placed in the ‘adult’ category

due to lack of other skeletal indicators to further delineate age (Table 3-6).

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Table 3-1. Radiocarbon dates from Naton Beach site, Tumon, Guam.a

Sample No. Provenience Material

Measured radiocarbon age

Conventional radiocarbon age

Calibrated 2σ age range

Beta- 238482 Feature 2 Soil 1700±40 BP 1680±40 BP AD 250–430

Beta- 238483 Burial 173b Conus shell beads 2490±40 BP 2940±40 BP 770–400 BC

Beta- 238484 Burial 156b Conus shell beads 2330±40 BP 2790±40 BP 590–330 BC

Beta- 238485 Burial 273b Conus shell beads 2490±40 BP 2860±40 BP 720–360 BC

Beta- 238486 Burial 286b Conus shell beads 2640±40 BP 2970±40 BP 790–490 BC a Table recreated from Defant (2008:150). b All burials are from the Pre-Latte assemblage.

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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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.

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

prehistoric 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, 1989; y’Edynak,

1989). Others 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 Valley 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; Brace 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

86

processing techniques, may allow for an accumulation of mutations, which lead to

decreased tooth size. Calcagno (1986, 1989) introduced the Selective Compromise

Effect (SCE), suggesting that decreased surface area and tooth complexity results in

fewer carious lesions, thus increasing an individual’s fitness. As 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

Density 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 functional 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 changes in

prehistoric society. The Pre-Latte (3500 BP – 1000 CE) and Latte (1000 – 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 populations, 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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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 intensification 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 Period (Butler, 1988; Ambrose et al., 1997;

Pietrusewsky et al., 1997). Material artifacts associated with 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 Melanesian dentition and the smaller

Polynesia dentition (Brace et al., 1981; Brace et al., 1990; 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 earlier 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 short 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 significant differences between parents and offspring dental sizes due to

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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:

Ho: 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 in 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

begin to accumulate (Brace, 1964; 1967; 1978; Brace and Mahler, 1971; Wolpoff, 1971;

Wolpoff, 1975; McKee, 1984). Following Sewell Wright’s (1931; 1964) theory of

mutation pressure, which suggests that mutations lead to a reduction in structures,

Brace and colleagues 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 period to the next (Christensen, 1998; Pinhasi et al., 2008).

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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. Smaller teeth are more resistant to dental caries

and crowding but are less resistant 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 forces 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 show 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

accordance with Wolff’s Law of Transformation (Wolff, 1892), that the mandible and

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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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 functional 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 body 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 (Machiarelli 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 indicators) (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 measurements 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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mesial and distal portion of the tooth, as expected in proper anatomical position. The

buccolingual 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 closest 0.01 mm and were not taken on teeth with moderate 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 the Kolmogorov-

Smirnov test and normal Q-Q plots on each dental measurement. Levene’s test of

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 period 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 the interaction between those

two variables.

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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 level 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 analyzed 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 total 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

population.

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 dental

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 mm) (see

Appendix A, Table A-3). However, in terms of overall cross-sectional area, maxillary

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first molar is the largest tooth in both males (AVG CX1 = 147.5 mm1) and females (AVG

CX1 = 138.2 mm2) (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

males’ and females’ mesiodistal diameter of 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 mm2. However, the maxillary first molar is

the largest tooth in females with a cross-sectional area of 128.1 mm2 (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 rates 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 the left and right sides.

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right maxillary molar (14%), right fourth premolar (11%), left mandibular second molar

(10%) and, left and right mandibular first incisors (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 Levene’s test and all but three 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 level (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

statistically 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 mandibular 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 slightly higher rate.

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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 come 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 buccolingual 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 between 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 time 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 between Time and Sex

Only six measurements have a significant interaction between time period and

sex: left maxillary lateral incisor BL, left mandibular fourth premolar MD, right

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mandibular canine BL, right mandibular second molar MD, right mandibular fourth

premolar BL, and right mandibular 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 measurements 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 slightly 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. These

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; Trembly,

1999; Pietrusewsky 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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range. The largest tooth summary comes from the Fujita sample, which were also

recovered 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

Fujita 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 San 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 measurements 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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and CX, show a slight increase in size from 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 biological significance or may simply

attest to the evolutionary importance of large canine size and it’s genetic stability. The

largest amount of dental reduction occurred in the lower third molars followed by the

upper second molars, upper right fourth premolar, 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 testing, 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, there 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 majority 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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the incisors and canines. Thus, over time the premolars and molars are reducing at a

significant 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 are 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 reduce 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 data2, 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 time2

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 Complex, which is currently being compiled for submission to the Guam Historic Preservation Office.

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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, pers. 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, males

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-

Latte: 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 very 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, pers. comm.).

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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 and 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.

Nonetheless, Walth’s stature data (pers. comm.)2 show very little difference in stature

between the time periods. The Pre-Latte male mean stature is 5’8” while the female

stature averages at 5’3”. The average Latte male height is 5’7” and the female mean

height is 5’2”. Overall, the Latte 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 stature 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 addressed 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. Looking 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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this drastic decrease in infectious disease may have more to do with a the introduced

cultural practice 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, these 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 density, 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-Functional 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 colleagues’ theory of PME (Brace, 1963; Brace and Mahler, 1971;

Brace and Hinton, 1981; Brace, 1987) as an explanation dental reduction is based on

Wright’s (1931; 1964) theory of mutation pressure and 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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changes in the exploitation of certain foods and also an increased amount of food

processing associated with the Latte period 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 be 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 reduces environmental stress due to competition for

nutritional resources (Machiarelli and Bondioli, 1986). All 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 periods 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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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 carious 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 the 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 practice of betel-nut chewing which is known to have

cariostatic properties.

The 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 SCE is a likely model to explain the dental reduction observed in the

prehistoric Chamorro.

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Masticatory Functional Hypothesis

Following Wolff’s Law (1892), the MFH posits that dental reduction occurs as a

result of decreased functional demands placed on the 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 with 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 affected 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

periods. 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 over 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 significant 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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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, greatly 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 tough 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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Table 4-1. Tooth summary data

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 populations 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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Table 4-3. Tooth summaries of Pacific and circum-Pacific samples

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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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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CHAPTER 5 DEVELOPMENTAL INSTABILITY: ENAMEL HYPOPLASIAS

Hans Selye (1973: 692) defines stress as ‘the nonspecific response of the body to

any demand made upon it’ where alterations in the environment change the normal and

steady state of an organism. During periods of environmental stress, an organism

undergoes 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 and 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 enamel 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 populations 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 deficiencies 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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1970; Goodman and Armelagos, 1985). These defects are a permanent record of

developmental disturbances that occurred during 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 disruption 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 enamel 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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et al., 2002; Steckel and Rose, 2002; Hillson, 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, 2003; 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 advances

in human culture that led to the rise of civilizations (Braidwood, 1960; Cohen, 1989).

However, research presented in Cohen and Armelagos’s (1984) edited volume

‘Paleopathology at the Rise of Agriculture’ demonstrated that overall rates of disease 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) coupled with

decrease in stature, and life expectancies were reported as evidence for declining

health (Cohen and Armelagos, 1984). While Cohen and Armelagos’ book (1984)

changed the paradigm of health and agriculture, it was faulty in that it was skewed

towards 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-1980s, 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 conducted on nearly every continent: North America (Larsen et al., 2002),

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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 and 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 undergoing 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

(Hodges, 1987; Neves and Wesolowski, 2002; Pietrusewsky and Douglas, 2002; Eshed

et al., 2004; Douglas and Pietrusewsky, 2007). In some cases, paleodemographic

studies have shown 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 series 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 decrease 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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BP – 1800 BP) has a decrease in carious lesions while LEH remain 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 into 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 (3500 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 Pre-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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Guam by its earliest inhabitants 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-Latte/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 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 tuber. 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; Dixon 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 C3 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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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-Anderson, 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. 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 population, at its

largest, ranged between 30,000 to 40,000.

The shift to agricultural intensification coupled with newly adapted food processing

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 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 over

a year and a half period, will be considered to assist in the construction a better health

profile of the Chamorro population.

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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). The determination of the burials’ age and cultural

affiliation was based on the stratigraphic 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

categorized as a ‘Type 4’ defect in the Developmental Defects of Enamel Index (FDI,

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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Walth1. 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.

LEH were scored macroscopically by both individual and tooth count methods, in

all available and unmodified teeth, as suggested by Lukacs (1989). LEH were not

scored 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 = absence 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 population 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 obliterate 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 likely 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 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 Preservation Office.

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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 surface, 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 incisors 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

frequencies of linear enamel hypoplasias 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 the mandibular canine (n = 27; 17%) and the maxillary central

incisor (n = 21; 14%).

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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 sample 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 differences between subadults and adults were 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 specific 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 females (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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Sex differences (with and without indeterminate sex category inclusion) 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 males 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

Linear enamel 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

terms 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 difference,

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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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 I1: p

= 0.008; right I1: p = 0.031; left C1: p = 0.012; right C1: 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 (Goodman 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 data 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

findings suggest 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 physiological stress than the Pre-Latte. The raw

frequency data 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 collected 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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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 (Slaus, 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 noted 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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Pietrusewsky 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 female Latte populations 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 physiological stress between the

time periods may indicate a cultural change in women’s access to resources over time.

Studies analyzing division of labor in hunter-gathering and farming populations have

shown significant dietary differences between the sexes (Hill and Hurtado, 1989; Walker

and Hewlett, 1990). Pre-Latte women 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 the 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 higher 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 combines both Pre-Latte and Latte

groups. The low frequency of the current study’s findings may be exacerbated by the

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inclusion of all observable teeth in the dental arcade, including third molars.

Additionally, the Pre-Latte cultural practice of labial abrasion (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 external 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

periostitis 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 population

(Walth, pers. comm.). Non-specific periostitis is the most common infectious disease

occurring in primarily the Latte sample and seven Latte individuals displayed periosteal

inflammation and infection indicative 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, just 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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Tinan, housed at the Bishop Museum in Honoulu, reported very high rates of

treponemal infection, 77%, (Suzuki, 1986). However, these remains were collected in

the early 1900s, 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 counterparts.

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 practice 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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positive correlation between an increase of hypoplastic defects with a shift in

subsistence economies and agricultural intensification. This trend is best illustrated in

the Dickinson Mounds burial complex of Illinois where Goodman and colleagues (1980;

1984) documented 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 well 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, 1992); 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

1350) to the Little Ice Age (AD 1350 to AD 1900), also known as the ‘AD 1300 Event’,

where rapid cooling temperatures, decline in sea levels, and increased storminess lead

to greater climatic variability (Bridgman, 1983; Hanson, 1991; Nunn and Britton, 2001;

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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

droughts (Nunn, 2007). Nunn (2007:1) describes the AD 1300 Event as the ‘most rapid

period of climate change to have occurred within the past several millennia’ and thus,

would have been associated with societal disruption, subsistence change, and

movement 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/lagoon 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). Additionally, 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 agricultural 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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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 to 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 reliance 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-1900s China (Zhou and Corruccini, 1998), have also shown a correlation between

seasonal resource depletion and systemic stress. Specifically, Temple’s analysis

(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 instability, 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 frequency 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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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 reason 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, while 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 to 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 fish 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 with less frequency of LEH and infectious disease. The increased

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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 and 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 climatic 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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Table 5-1. Percentage of teeth with one or more linear enamel hypoplasiaa

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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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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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, 1999

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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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 using 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 Table 5-8. Pre-Latte and Latte differences 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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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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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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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

Maxillary38 38 38 38 38 38Mandibular38 38 38 38 38 38 38

LE

H F

req

ue

ncy

Tooth Type

Pre-Latte

Latte

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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 more 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 populations (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 agriculture 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 population 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 carious lesions between the Pre-

Latte and Latte time periods.

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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) describes dental caries as ‘a disease process characterized by the focal

demineralization of dental hard tissues by organic acids produced by bacterial

fermentation of dietary carbohydrates, especially sugars.’ The pH balance of plaque is

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 carious 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

(Newbrun, 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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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;

Bernal et al., 2007). Cohen and Armelagos’ (1984) benchmark book, Paleopathology at

the Origins of Agriculture, validated Turner’s claim with the consensus that carious

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

Larsen, 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 Tayles,

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 evidence 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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2007; Domett and Tayles, 2007). This geographical trend has been attributed to the

transition to rice-based agriculture, which was not analyzed in Cohen and Armelagos’

(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

(Temple 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, Tayles and colleagues (2009:163) suggest that the variability in rice cariogenicity

is the result of ‘structure, the method of processing, and the extent to which [it is]

gelatinised during cooking.’ Thus, the decreased frequency of carious lesions in some

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 agriculture. 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 population 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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et al., 1991; McGovern and Wilson, 1996). Horticultural practices were likely brought to

Guam by its earliest inhabitants 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-Latte/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 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 tuber. 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; Dixon 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 C3 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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preferential exploitation of reef and/or lagoon fish over pelagic resources (Hanson,

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-Anderson, 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. 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 population, 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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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. The determination of the burials’ age and cultural affiliation was based

on the stratigraphic location, associated artifacts, and radiocarbon dates (DeFant,

2008). Dating to roughly 2,500 BP, the Naton Beach 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 remains 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 individual 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 total of 3,666 teeth were analyzed: 1,930

from the Pre-Latte and 1,736 from the Latte (Table 6-2). Females1 are represented

1 Demographic data, as well as postcranial pathological conditions, 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 Preservation Office.

146

more often 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 bioarchaeological 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, 1996). 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 differences 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), especially 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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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). Again, 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 small. The Pre-Latte population shows an increase of

carious lesion frequencies from 40% (n = 2) in juveniles to 78.1% (n = 50) in young

adults, which then decreased to 73.9% (n = 17) in middle-aged adults. 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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display carious lesions. Similarly, in the Latte period, 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 infected teeth. A

Pearson Chi-Square test found no significant 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).

Time 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 tooth 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 significant 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., 2010; Meng et

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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 anterior dentition typically occurs at the cemento-

enamel junction, and at times, the root surface was also involved (Figure 1). Similar

findings of high rates of carious lesions in the anterior 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-period Kangjia of Northern 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.

No 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 sites 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. In the few individuals where

maxillary or mandibular reconstruction was possible, dental crowding was evident in the

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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 period. 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 two 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 nature 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 dentition 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 studies 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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sexes were eating foods with similar levels of cariogenicity. 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 carious 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 differential 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 females (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 carious 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 possibility of

temporal changes in health and disease between the periods with 14.3% of the Pre-

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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 colleagues (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-Latte

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 sample

(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 disparity 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; Ubelaker and Newson, 2002; Pechenkina et al.,

2002; Krigbaum, 2007; Temple, 2007; Temple and Larsen, 2007). In the current study,

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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; however, other causative factors will also be

explored. Turner (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 unexpectedly 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 of 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 may have had participated more in incipient agricultural

activities than has been observed through the archaeological record.

An alternative explanation for the high rates of carious 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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analyses (Ambrose et 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 archaeological 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).

Latte: 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

period (640 ± 50 BP; calibrated date range, 2σ: AD 1284-1425) (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. The 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 (Krigbaum, 2007; Temple and Larsen, 2007), while

others have shown the opposite (Oxenham et al., 2000; Tayles et al., 2000;

Pietrusewsky and Douglas, 2002; Oxenham et al., 2006; Douglas and Pietrusewsky,

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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 Butler, 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 practices 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 significantly to the prehistoric

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diet. Even if this is the case and rice was not 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 sugarcane, 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 carious 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 carious

lesions 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

established 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 (CaCO3) (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 effects 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, 1984; Moller et al., 1977; Chatrchaiwiwatana, 2006). Hanson

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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 chemical 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 displaying betel-nut staining. The

majority of the adult Latte population displays betel-nut staining (64%), while no

juveniles under the age of 18 were found with staining (Parr, 2012). When 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 lesions, 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 dental 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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associated with intensification of agricultural practices, resulting 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 in 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 elevated 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 horticultural one (Turner, 1979). This finding may be

suggestive increased 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 carbohydrate-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 relationship 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 cultural

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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infection and enamel destruction through its physical and chemical properties, despite

the highly cariogenic diet of the agricultural population.

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Table 6-1. Individual occurrence carious lesion frequencies

Pre Latte Latte

n/Na

% n/Na %

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/Na % n/Na %

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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Table 6-4. Frequency of carious lesions by tooth class

Pre Latte Latte

Tootha n/Nb % n/Nb %

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/Na % n/Na %

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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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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Table 6-7. Carious lesion frequencies of the prehistoric Chamorro populations across 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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Figure 6-1. Pre-Latte carious lesions in the anterior dentition (Photo by author).

Figure 6-2. Pre-Latte dental crowding (Photo by author).

Figure 6-3. Betel-nut with piper leaf and slacked lime (Photo by author).

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CHAPTER 7 SUMMARY

This research is a diachronic investigation of the evolutionary 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 prehistoric 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 Beach

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 supplemented 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 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). Thus, it was

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predicted that cultural and environmental shifts are likely to be accompanied 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% decrease 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 conjunction 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, which 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

best support a combination of Calcagno’s (1986; 1989) Selective Compromise Effect

and Carlson and Van Gerven’s (1977) Masticatory Functional Hypothesis.

167

There were no significant differences in the rates of juvenile and adult hypoplasias;

however, 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 demonstrated 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 Pre-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 period, 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 number 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

168

staple 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 despite 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 broad

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 Lapita expansion that began around 3200

BP and led to the colonization of Polynesia (Kirch, 2010). However, settlement of the

Marianas Islands has been securely dated to 3600 to 3420 BP (Carson, 2008; Clarke et

al., 2010), approximately 400 years before the Lapita peoples ventured 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 (Pietrusewsky,

1990a; 1990b; 1994; 2006; Ishida and Dodo, 1997). The 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.

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 prehistoric 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. Archaeological 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 betel-nut chewing. This temporal assessment of the dentition identifies cultural

and environmental processes that led to biological change in the prehistoric Chamorro.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

193

Table A-6. Group comparisons of dental measurements

Measurement Time Period N Mean Std. Dev.

Std. Error Mean Differencea

Percent of Changeb

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

194

Table A-6. Continued

Measurement Time Period N Mean Std. Dev.

Std. Error Mean Differencea

Percent of Changeb

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

195

Table A-6. Continued

Measurement Time Period N Mean Std. Dev.

Std. Error Mean Differencea

Percent of Changeb

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

196

Table A-6. Continued

Measurement Time Period N Mean Std. Dev.

Std. Error Mean Differencea

Percent of Changeb

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

197

Table A-6. Continued

Measurement Time Period N Mean Std. Dev.

Std. Error Mean Differencea

Percent of Changeb

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

198

Table A-6.. Continued

Measurement Time Period N Mean Std. Dev.

Std. Error Mean Differencea

Percent of Changeb

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

199

Table A-6. Continued

Measurement Time Period N Mean Std. Dev.

Std. Error Mean Differencea

Percent of Changeb

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

200

Table A-7. Kolmogorov-Smirnova Test for normality

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*

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*

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*

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.

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

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

206

APPENDIX B ANALYSES OF VARIANCE TESTS FOR DENTAL MEASUREMENTS

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 3.596a 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 1.209a 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)

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 1.651a 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)

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 0.00007a

Intercept 3285.352 1 3285.352 27927.215 0.00

TimePeriod 0.494 1 0.494 4.202 0.04b

SexCombined 1.848 1 1.848 15.709 0.00b

TimePeriod * SexCombined 0.573 1 0.573 4.875 0.03b

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

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 3.989a 3 1.33 8.015 0.00011a

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 0.00b 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

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 3.367a 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)

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 6.572a 3 2.191 9.997 0.00001a

Intercept 5168.768 1 5168.768 23586.009 0.00

TimePeriod 4.613 1 4.613 21.049 0.00b

SexCombined 1.462 1 1.462 6.673 0.01b

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

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 6.839a 3 2.28 6.899 0.00033a

Intercept 8777.969 1 8777.969 26566.965 0.00

TimePeriod 4.814 1 4.814 14.571 0.00b

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

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 3.817a 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)

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 7.387a 3 2.462 6.679 0.00040a

Intercept 9134.581 1 9134.581 24776.573 0.00

TimePeriod 3.061 1 3.061 8.303 0.00b

SexCombined 2.669 1 2.669 7.239 0.01b

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

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 6.835a 3 2.278 8.657 0.00005a

Intercept 9960.838 1 9960.838 37847.924 0.00

TimePeriod 3.734 1 3.734 14.187 0.00b

SexCombined 2.77 1 2.77 10.524 0.00b

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

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 9.023a 3 3.008 10.362 0.00001a

Intercept 12272.892 1 12272.892 42282.653 0.00

TimePeriod 4.685 1 4.685 16.141 0.00b

SexCombined 3.665 1 3.665 12.626 0.00b

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 13.957a 3 4.652 15.346 0.00000a

Intercept 7942.252 1 7942.252 26198.798 0.00

TimePeriod 7.42 1 7.42 24.475 0.00b

SexCombined 2.185 1 2.185 7.208 0.01b

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

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 22.942a 3 7.647 21.584 0.0000a

Intercept 11879.491 1 11879.491 33527.741 0.00

TimePeriod 14.303 1 14.303 40.368 0.00b

SexCombined 4.26 1 4.26 12.023 0.00b

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 9.775a 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)

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 6.851a 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)

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 1.381a 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 3.021a 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)

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 2.116a 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)

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 3.435a 3 1.145 8.345 0.00007a

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 0.00b

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

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 3.010a 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 3.284a 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)

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 6.590a 3 2.197 8.209 0.00007a

Intercept 9185.092 1 9185.092 34324.858 0.00

TimePeriod 4.824 1 4.824 18.026 0.00b

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 5.308a 3 1.769 9.707 0.00001a

Intercept 4896.429 1 4896.429 26863.383 0.00

TimePeriod 3.521 1 3.521 19.32 0.00b

SexCombined 1.696 1 1.696 9.303 0.00b

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

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 13.591a 3 4.53 17.023 0.00000a

Intercept 8849.841 1 8849.841 33254.095 0.00

TimePeriod 7.204 1 7.204 27.07 0.00b

SexCombined 4.575 1 4.575 17.19 0.00b

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 6.817a 3 2.272 9.041 0.00003a

Intercept 11153.872 1 11153.872 44379.428 0.00

TimePeriod 6.286 1 6.286 25.011 0.00b

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

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 10.285a 3 3.428 13.18 0.00000a

Intercept 13092.688 1 13092.688 50334.755 0.00

TimePeriod 5.236 1 5.236 20.132 0.00b

SexCombined 4.575 1 4.575 17.59 0.00b

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 16.801a 3 5.6 18.605 0.0000a

Intercept 9320.397 1 9320.397 30962.461 0.00

TimePeriod 11.614 1 11.614 38.581 0.00b

SexCombined 2.76 1 2.76 9.168 0.00c

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

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 19.825a 3 6.608 17.436 0.00000a

Intercept 12923.433 1 12923.433 34099.631 0.00

TimePeriod 13.13 1 13.13 34.645 0.00b

SexCombined 4.096 1 4.096 10.807 0.00c

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 3.030a 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)

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 4.869a 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 3.498a 3 1.166 8.023 0.00014a

Intercept 1881.394 1 1881.394 12944.752 0.00

TimePeriod 3.145 1 3.145 21.636 0.00b

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

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 1.021a 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 2.137a 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)

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 7.250a 3 2.417 9.571 0.00002a

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 0.00b

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

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 8.290a 3 2.763 13.657 0.00000a

Intercept 5498.925 1 5498.925 27176.031 0.00

TimePeriod 1.806 1 1.806 8.926 0.00b

SexCombined 6.223 1 6.223 30.752 0.00b

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

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 7.152a 3 2.384 14.168 0.00000a

Intercept 5576.195 1 5576.195 33138.75 0.00

TimePeriod 3.141 1 3.141 18.665 0.00b

SexCombined 3.193 1 3.193 18.977 0.00c

TimePeriod * SexCombined 0.978 1 0.978 5.81 0.02b

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 3.386a 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)

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 5.820a 3 1.94 10.709 0.00000a

Intercept 5933.765 1 5933.765 32751.291 0.00

TimePeriod 1.424 1 1.424 7.861 0.01b

SexCombined 3.369 1 3.369 18.594 0.00b

TimePeriod * SexCombined 0.891 1 0.891 4.917 0.03c

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

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 6.441a 3 2.147 8.575 0.00004a

Intercept 7381.609 1 7381.609 29484.726 0.00

TimePeriod 2.376 1 2.376 9.489 0.00b

SexCombined 3.874 1 3.874 15.473 0.00b

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

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 4.344a 3 1.448 6.442 0.00061a

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 0.00b

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 6.261a 3 2.087 11.262 0.00000a

Intercept 10230.494 1 10230.494 55205.338 0.00 TimePeriod 1.255 1 1.255 6.774 0.01b

SexCombined 4.932 1 4.932 26.615 0.00b

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

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 5.804a 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 16.986a 3 5.662 13.04 0.00000a

Intercept 9889.247 1 9889.247 22776.433 0.00

TimePeriod 9.171 1 9.171 21.123 0.00b

SexCombined 6.205 1 6.205 14.291 0.00b

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

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 14.361a 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)

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 10.903a 3 3.634 10.974 0.00001a

Intercept 5020.924 1 5020.924 15161.674 0.00

TimePeriod 7.384 1 7.384 22.297 0.00b

SexCombined 2.949 1 2.949 8.905 0.00b

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)

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)

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 0.00000a 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 0.00b 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

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 0.00008a 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 0.00b TimePeriod * SexCombined

1.459 1 1.459 7.079 0.01b

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

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 0.00001a Intercept 5266.893 1 5266.893 25532.716 0.00 TimePeriod 2.49 1 2.49 12.073 0.00b SexCombined 2.508 1 2.508 12.159 0.00b 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)

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 0.00000a Intercept 5637.778 1 5637.778 35835.681 0.00 TimePeriod 2.59 1 2.59 16.462 0.00b SexCombined 4.774 1 4.774 30.344 0.00b 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

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 0.00018a Intercept 10197.624 1 10197.624 31229.701 0.00 TimePeriod 3.274 1 3.274 10.028 0.00b SexCombined 5.262 2 2.631 8.058 0.00b 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

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 0.00000a Intercept 20146.669 1 20146.669 65924.701 0.00 TimePeriod 3.391 1 3.391 11.095 0.00b SexCombined 5.407 2 2.704 8.847 0.00b 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

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 0.00000a Intercept 11000.381 1 11000.381 58862.344 0.00 TimePeriod 2.322 1 2.322 12.427 0.00b SexCombined 6.328 1 6.328 33.859 0.00b 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)

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 0.00000a Intercept 9750.763 1 9750.763 38183.968 0.00 TimePeriod 6.657 1 6.657 26.068 0.00b SexCombined 3.592 1 3.592 14.067 0.00b 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

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 0.00000a Intercept 5082.833 1 5082.833 13456.099 0.00 TimePeriod 16.495 1 16.495 43.668 0.00b SexCombined 2.826 1 2.826 7.48 0.01b 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)

249

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BIOGRAPHICAL SKETCH

Nicolette Maria Luney Parr graduated with highest honors from the University of

Florida in 2002 with a bachelor’s degree in anthropology and minor in religion. Ms.

Parr’s graduate studies began at the University of Indianapolis, were she completed a

Master of Science degree in human biology. 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 crime 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 master’s thesis on determining ancestry from the

mandible.

Ms. Parr entered the anthropology 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 introductory 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, which 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, Ms. Parr was hired by the Joint Pow/MIA

Accounting Command-Central Identification Laboratory in Honolulu, Hawaii, where she

works as a forensic anthropologist.