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SSStttooonnnyyy BBBrrrooooookkk UUUnnniiivvveeerrrsssiiitttyyy The official electronic file of this thesis or dissertation is maintained by the University Libraries on behalf of The Graduate School at Stony Brook University. ©©© AAAllllll RRRiiiggghhhtttsss RRReeessseeerrrvvveeeddd bbbyyy AAAuuuttthhhooorrr... Morphometric Variation in the Appendicular Skeleton of Recent and Prehistoric Humans A Dissertation Presented by Danielle Frances Royer to The Graduate School in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Anthropology (Physical Anthropology) Stony Brook University August 2009 Copyright by Danielle Frances Royer 2009 Stony Brook University The Graduate School Danielle Frances Royer We, the dissertation committee for the above candidate for the Doctor of Philosophy degree, hereby recommend acceptance of this dissertation. Frederick E. Grine – Dissertation Advisor Professor, Departments of Anatomical Sciences and Anthropology John G. Fleagle – Chairperson of Defense Professor, Department of Anatomical Sciences William L. Jungers Professor and Chair, Department of Anatomical Sciences Osbjorn M. Pearson Department of Anthropology, University of New Mexico This dissertation is accepted by the Graduate School Lawrence Martin Dean of the Graduate School ii Abstract of the Dissertation Morphometric Variation in the Appendicular Skeleton of Recent and Prehistoric Humans by Danielle Frances Royer Doctor of Philosophy In Anthropology (Physical Anthropology) Stony Brook University 2009 The study of phenotypic variation can help elucidate the various influences that have shaped within-population morphometric diversity within Homo sapiens, and provides another line of evidence to complement molecular and craniometric research on human evolution. Moreover, because modern skeletal samples frequently provide a framework for interpreting diversity and taxonomic relationships in the fossil record, it is imperative to test assumptions concerning the magnitude and pattern of past and present-day skeletal variability, since inequalities may contribute to serious biases. The primary aim of this dissertation is to broaden our understanding of phenotypic diversity in the postcranial skeleton of select modern and prehistoric H. sapiens groups. This is accomplished through univariate comparisons of the magnitude of morphometric variability in the major long bones of the appendicular skeleton 1) between recent iii African populations, 2) between recent African populations and prehistoric samples from the terminal Pleistocene and early Holocene, 3) between modern African populations and samples of early H. sapiens from the Middle Stone Age (MSA) and early Upper Paleolithic (EUP), and 4) between MSA and EUP fossil samples. This dissertation demonstrates equal relative variation among recent African populations, although some sex-specific samples exhibit significant differences in the magnitude of appendicular variation. African populations from the terminal Pleistocene and early Holocene are characterized by an equal magnitude and pattern of appendicular variation relative to recent Africans. In contrast, both MSA and EUP fossil samples exhibit an elevated magnitude of variation throughout the appendicular skeleton compared to recent and prehistoric populations, demonstrating that modern diversity substantially underestimates the early diversity of our lineage. However, the MSA sample, which spans over 100,000 years, is not more variable than the EUP sample which spans less than 15,000 years. These results suggest that time averaging alone is insufficient to account for the increase of variation documented in the fossil samples. Furthermore, the reduction of diversity to present-day levels appears to be bracketed between the end of the early Upper Paleolithic and the terminal Pleistocene at approximately 14,000 years BP. This dissertation also tested the accuracy of metric and non-metric methods for classifying sex from fragmentary pelvis remains in modern Africans. The most reliable method, discriminant analysis, is used to diagnose the sex of prehistoric humans, including the first formal diagnosis for Omo I, the earliest H. sapiens fossil currently known. The unambiguous female diagnosis for Omo I suggests that at least some of the greater diversity documented in the past may reflect stronger sexual dimorphism. iv Table of Contents List of Figures…………………………………………………………………....ix List of Tables………………………………………………………………….....xi Acknowledgments……………………………………………………...............xiv Chapter 1 – Introduction………………………………………………………….1 Goals of the Dissertation…………………………………………………..6 Chapter 2 – Data Collection Methods and Skeletal Samples………………….....9 Skeletal Measurements…………………………………………………....9 Measurement Error and Reliability………………………………............10 Data Transformation and Analysis: An Overview…………………….....14 Skeletal Samples…………………………………………………………15 Reference Samples: Rationale for Choice……………………….16 Recent Africans: Zulu……………………………………………18 Recent Africans: Kikuyu…………………………………………18 Recent Africans: Nilotic Ugandans………………………….......19 Archaeological Africans: Khoe-San……………………………..20 Archaeological Africans: Sudanese……………………………...22 Archaeological Africans: Taforalt……………………………….23 Early Homo sapiens Fossils……………………………………...24 Middle Stone Age Sample……………………………………….26 Early Upper Paleolithic Sample………………………………….37 Figures……………………………………………………………………46 Tables…………………………………………………………………….47 Chapter 3 – A Test of Methods for Determining Sex Using Fragmentary Os Coxae………………………………………………………...……………….61 Introduction………………………………………………………………61 Goals……………………………………………………………..61 Determining Sex from Skeletal Remains………………………...62 Skeletal Samples………………………………………………………....67 Method: Testing the Accuracy of the Visual Method…………………...68 Preauricular Surface Character…………………………………..69 Composite Arch Character…………………………………….…71 Greater Sciatic Notch Character…………………………………72 Sacroiliac Complex: Final Diagnosis…………………….………74 Results: Accuracy of the Visual Method………………………………...74 Discussion: Utility of the Visual Method……………….…………….....75 Method: Testing the Accuracy of Discriminant Function Analysis…..…77 Assumptions……….……………………………………….........79 Os Coxae Variables: Form versus Shape………..………….........80 v Stepwise Procedure…………………………………....................81 Determining Accuracy…………………………………………...82 Results: Accuracy of Discriminant Function Analysis…………………..83 Accuracy of Form DFA…………………….…………………....83 Accuracy of Shape DFA…………………………………………86 Discussion: Utility of Discriminant Function Analysis …........................91 Conclusions: Comparison of the Methods……………………………….94 Figures……………………………………………………………………96 Tables…………………………………………………………………...106 Chapter 4 – Classifying Sex in Prehistoric Humans Using Fragmentary Os Coxae………………………………………………………………………..117 Introduction……………………………………………………………..117 Skeletal Samples………………………………………………………..117 Methods………………………………………………………………....118 Results……………………….………………………………………….122 Classification of Sex: Archaeological Samples………………...122 Classification of Sex: Early H. sapiens Fossils………………...123 Discussion: Utility of the Methods……………………………………..125 Archaeological Samples………………………………………...125 Early H. sapiens Fossils………………………………………...129 Omo I…………………………………………………………...131 Qafzeh 9………………………………………………………...132 Skhūl 4………………………………………………………….133 Grotte des Enfants 4…………………………………………….133 Grotte des Enfants 5…………………………………………….134 Nazlet Khater 2…………………………………………………135 Paviland 1……………………………………………………….135 Skhūl 5………………………………………………………….136 Barma Grande 2………………………………………………...137 Cro-Magnon 1…………………………………………………..138 Cro-Magnon 4315………………………………………………139 Grotte du Cavillon 1…………………………………………….139 Mladeč 21……………………………………………………….140 Conclusions……………………………………………………………..141 Figures…………………………………………………………………..143 Tables…………………………………………………………………...158 Chapter 5 – Morphometric Variation in the Appendicular Skeleton: Testing the Assumption of Equality Among Recent Sub-Saharan Africans………………..173 Introduction……………………………………………………………..173 Goals……………………………………………………………173 Assumption of Equality of Variation in Modern Humans……...174 vi Sources of Variation in Modern Humans………………………176 Skeletal Samples………………………………………………………..179 Methods: Equality of Relative Variation……………………………….180 Hypotheses……………………………………………………………...186 Results…………………………………………………………………..190 Equality of Variation Between Females………………………..190 Equality of Variation Between Males…………………………..193 Equality of Variation Between the Sexes………………………194 Equality of Variation Between Populations…………………….197 Discussion: The Assumption of Equality………………………………199 Conclusions……………………………………………………………..207 Figures…………………………………………………………………..210 Tables…………………………………………………………………...239 Chapter 6 – Morphometric Variation in the Appendicular Skeleton of Prehistoric Humans……………………………………………………………..270 Introduction……………………………………………………………..270 Goals……………………………………………………………270 Variation in Early H. sapiens…………………………………...271 Measurements…………………………………………………………..278 Skeletal Samples………………………………………………………..278 Methods: Testing for Differences in Relative Variation………………..280 Hypotheses……………………………………………………………...280 Results…………………………………………………………………..284 Equality of Variation Between the Zulu and Archaeological Samples…………………………………………………………285 Excess Variation in MSA and EUP Fossils Compared to Modern Humans………………………………………………...287 Excess Variation in MSA Compared to EUP H. sapiens………293 Discussion………………………………………………………………293 Variability in the Terminal Pleistocene and Holocene ………...293 Variability