The Utility of Cladistic Analysis of Nonmetric Skeletal Traits for Biodistance Analysis by James Christopher Reed B.A., Mississi
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The Utility of Cladistic Analysis of Nonmetric Skeletal Traits for Biodistance Analysis by James Christopher Reed B.A., Mississippi State University, 1994 M.A., University of South Carolina, 1998 Submitted to the Graduate Faculty of Arts and Sciences in partial fulfillment of the requirements for the degree of Doctor of Philosophy University of Pittsburgh 2006 UNIVERSITY OF PITTSBURGH Faculty of Arts and Sciences This dissertation was presented by James Christopher Reed It was defended on April 17, 2006 and approved by Dr. Michael I. Siegel, Professor, Department of Anthropology Dr. Mark P. Mooney, Professor, Department of Oral Medicine and Pathology Dr. Marc P. Bermann, Associate Professor, Department of Anthropology Dr. Jeffrey H. Schwartz, Professor, Department of Anthropology Dissertation Director ii Copyright © by James Christopher Reed 2006 iii The Utility of Cladistic Analysis of Nonmetric Skeletal Traits for Biodistance Analysis James Christopher Reed, PhD University of Pittsburgh, 2006 A significant focus of bioarchaeology is biodistance analysis, which seeks to determine the biological affinities of human groups and to support arguments about prehistoric and historic cultural topics, such as migration, marriage, and residential patterns. Although genetic comparisons are becoming more common, metric and nonmetric skeletal traits remain the primary source of information on human populations. Biodistance analysis is grounded theoretically and methodologically in phenetics, which is an approach developed by systematists to group organisms on the basis of overall similarity. However, while phenetics was adopted by physical anthropologists and bioarchaeologists as the foundation of biodistance analysis, systematists have long since moved away from phenetic approaches for determining relatedness to hypothetico-deductively based cladistic analyses. It is time for physical anthropologists and bioarchaeologists engaged in biodistance analysis do so as well. It is perhaps an irony that biodistance analysis, which seeks to delineate the biological relationships of group, begins by defining the groups (samples) on the basis of archaeological, cultural, or linguistic information prior to any morphological/biological comparison. However, the delineation and comparison of groups should be based from the beginning on the biology (morphology) of individuals and then of groups and, more specifically, on unique biological features, not cultural or linguistic criteria. A cladistic analysis can provide a biologically based delineation of groups. In this study I investigate whether unique, nonmetric characters can be delineated for small groups such as those traditionally the focus of biodistance analysis and, thus, iv whether cladistic analysis is an appropriate substitute for the phenetic approach in biodistance analysis. Four samples of skeletal material were examined. One, the Spitalfields collection, consists of burials of individuals whose familial relationships are well documented. The other samples are undocumented and compared to the Spitalfields sample in an attempt to delineate unique characters that might define groups. The result was that no unique characters could be delineated, which means that cladistic analysis, while perhaps applicable to study of higher-level groups within the species, fails at the population level. Consequently, while unsatisfactory, biodistance analysis must continue to rely on abiological criteria for defining populations. v TABLE OF CONTENTS TABLE OF CONTENTS.................................................................................................VI LIST OF TABLES ..........................................................................................................XI LIST OF FIGURES........................................................................................................XII PREFACE ................................................................................................................... XVI 1.0 THESIS AND RESEARCH ORIENTATION..................................................... 1 1.1 INTRODUCTION ...................................................................................... 1 1.2 THEORETICAL ORIENTATION............................................................... 4 1.3 HYPOTHESES FOR THIS PROJECT...................................................... 6 1.4 CHAPTER SYNOPSES............................................................................ 7 2.0 PHYSICAL ANTHROPOLOGY, TYPOLOGICAL ANALYSIS, AND BIODISTANCE ANALYSIS ........................................................................... 11 2.1 INTRODUCTION .................................................................................... 11 2.2 RACE, TYPOLOGY AND EARLY PHYSICAL ANTHROPOLOGY ....... 12 2.2.1 Race and Forensic Anthropology ................................................. 18 2.2.2 Prehistoric groups.......................................................................... 19 2.3 TRADITIONAL ANALYSIS OF NONMETRIC SKELETAL TRAITS ...... 22 2.3.1 Animal models and the application of nonmetric trait analysis .23 2.3.2 Current standards of analysis ....................................................... 25 2.3.3 Heritability of nonmetric traits....................................................... 27 2.4 POPULATION GENETICS..................................................................... 28 2.5 HUMAN VARIATION, BIOARCHAEOLOGY, AND BIODISTANCE ANALSYIS............................................................................................... 30 3.0 BIODISTANCE, GENETICS, AND PHENETICS ........................................... 33 3.1 INTRODUCTION .................................................................................... 33 vi 3.2 CONCEPTS OF GENETICS AND THE ANALYSIS OF TRAITS........... 34 3.2.1 Thresholds and epigenetics .......................................................... 34 3.2.2 Phylogeny and Developmental genetics ...................................... 36 3.2.3 Mitochondrial DNA and the molecular clock................................ 37 3.3 BIODISTANCE AND PHENETICS......................................................... 40 3.3.1 Phenetics/numerical taxonomy..................................................... 40 3.3.2 The statistics of biodistance ......................................................... 45 3.3.3 Phenetics and physical anthropology .......................................... 51 3.4 PHENETIC ANALYSIS AND ITS USEFULNESS IN ANTHROPOLOGICAL RESEARCH ....................................................... 53 3.4.1 Logical problems of phenetic analysis......................................... 54 3.4.2 The assumption of similarity ......................................................... 56 3.5 AN ALTERNATIVE APPROACH ........................................................... 57 4.0 CLADISTICS AND CHARACTER CHOICE .................................................. 59 4.1 INTRODUCTION .................................................................................... 59 4.2 CLADISTICS .......................................................................................... 61 4.2.1 Cladistic methods of phylogenetic reconstruction ..................... 63 4.2.2 Homology and Synapomorphy...................................................... 66 4.2.3 Cladistic hypothesis formulation .................................................. 69 4.2.4 Problems of identification of biological human groups through morphological analysis.................................................................. 71 4.2.5 Cladistics and Biodistance analysis ............................................. 73 4.2.6 Outgroup comparison .................................................................... 74 4.3 CHARACTER CHOICE .......................................................................... 76 5.0 METHODS AND SAMPLES .......................................................................... 79 5.1 INTRODUCTION .................................................................................... 79 5.1.1 Trait choice and cladistics............................................................. 80 5.2 SUMMARY OF METHODS .................................................................... 80 5.3 SAMPLES .............................................................................................. 85 5.3.1 Sample Choice................................................................................ 85 5.3.2 Spitalfields ...................................................................................... 88 vii 5.3.3 Terry Collection .............................................................................. 90 5.3.4 Prehistoric North American Skeletal Sample 1 – The Campbell Farm Site ......................................................................................... 91 5.3.5 Prehistoric North American Skeletal Sample 2 – The Perry Site 92 6.0 RESULTS ...................................................................................................... 93 6.1 INTRODUCTION .................................................................................... 93 6.2 METHOD OF EXAMINATION ................................................................ 95 6.3 CRANIAL ANALYSIS ............................................................................ 96 6.3.1 Pterygobasal region ....................................................................... 97 6.3.1.1 Lateral pterygoid plate/ptergospinous bridging.................. 97 6.3.1.2 Foramen ovale and foramen