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Open Hill Final.Pdf The Pennsylvania State University The Graduate School College of the Liberal Arts EVOLUTIONARY AND DEVELOPMENTAL HISTORY OF TEMPORAL BONE PNEUMATIZATION IN HOMINIDS A Dissertation in Anthropology by Cheryl Ann Hill © 2008 Cheryl Ann Hill Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy December 2008 The dissertation of Cheryl Ann Hill was reviewed and approved* by the following: Joan T. Richtsmeier Professor of Anthropology Dissertation Adviser Chair of Committee Kenneth M. Weiss Evan Pugh Professor of Anthropology and Genetics Alan C. Walker Evan Pugh Professor of Anthropology and Biology Carol V. Gay Professor Emeritus of Biochemistry and Molecular Biology Timothy M. Ryan Research Associate of Anthropology Special Member Nina Jablonski Professor of Anthropology Head of the Department of Anthropology *Signatures are on file in the Graduate School ii ABSTRACT This project investigates ontogenetic and evolutionary changes in temporal bone pneumatization, the process leading to air-filled cavities in adult temporal bones. Important conclusions regarding hominid phylogeny have included temporal bone pneumatization as a phylogenetic, or species-specific, marker. Despite its previous inclusion in evolutionary analyses, a number of questions remain concerning the evolutionary and developmental changes in hominid temporal bone pneumatization, as well as the developmental and functional basis for observed variation in pneumatization. This study includes three sets of analyses aimed at determining whether temporal bone pneumatization is an appropriate and informative marker for evolutionary change. First, ontogenetic changes in temporal bone pneumatization in modern humans were investigated. Second, comparative analysis of adult morphology of temporal bone pneumatization in five extant primate species was completed. Third, evolutionary changes in temporal bone pneumatization in both living and fossil Homininae species were analyzed. For the first and second studies, bone volume fractions, anisotropy, trabecular number, trabecular thickness, surface area and volume were measured in high resolution x-ray computed tomography scans acquired from adult primates and a cross- sectional ontogenetic series of humans. The third study was achieved using medical resolution scans previously acquired from fossil species, extant African apes, and modern humans. Changes in the extent of pneumatization over evolutionary and ontogenetic time are accompanied by changes in the bony material analyzed here. A number of species- specific and age-specific differences in temporal bone pneumatization were noted. The iii species-specific changes did not correspond to known evolutionary or morphological affinities among the species in the study. Analyses of the ontogenetic changes in pneumatization show that temporal bone pneumatization is a dynamic process that is influenced by epigenetic factors. These results suggest that temporal bone pneumatization is not appropriate for inclusion in phylogenetic analyses, since the morphology of temporal bone pneumatization demonstrates patterning, either stochastic or of unknown origin, across species and age groups. iv TABLE OF CONTENTS List of Figures ............................................................................................................. viii List of Tables .................................................................................................................. x . Acknowledgements ........................................................................................................xi Chapter One: INTRODUCTION AND LITERATURE REVIEW I. INTRODUCTION ....................................................................................................... 1 II. BACKGROUND ........................................................................................................ 3 Definitions ................................................................................................................. 3 Pneumatization in H. sapiens temporal bones ............................................................. 5 Disease and pneumatization in H. sapiens .................................................................. 8 Interspecific differences in pneumatization patterns.................................................. 11 Development and evolution of the temporal bone ..................................................... 15 Interspecific differences in development of pneumatized spaces ............................... 18 Functional hypotheses .............................................................................................. 20 Evolutionary changes influencing pneumatization of the craniofacial complex ......... 23 III. PREVIOUS METHODS FOR ANALYZING PNEUMATIZATION....................... 24 Gross anatomy, histology and casts .......................................................................... 24 Imaging .................................................................................................................... 25 Quantification .......................................................................................................... 29 IV. CONCLUSIONS .................................................................................................... 30 Chapter Two: METHODS AND PRELIMINARY WORK I. INTRODUCTION ..................................................................................................... 32 II. METHODOLOGY FOR CURRENT STUDY .......................................................... 32 Scan acquisition protocol ......................................................................................... 33 Image processing...................................................................................................... 35 Quant3D analyses .................................................................................................... 39 Surface area and volume estimation ......................................................................... 42 III. POTENTIAL LIMITATIONS AND THEIR RESOLUTION................................... 43 Impact of image resolution ....................................................................................... 43 Precision and repeatability........................................................................................ 48 IV. CONCLUSIONS .................................................................................................... 50 Chapter Three: ONTOGENETIC CHANGE IN TEMPORAL BONE PNEUMATIZATIONINH. SAPIENS I. INTRODUCTION ..................................................................................................... 51 II. BACKGROUND ...................................................................................................... 51 III. RESEARCH DESIGN ............................................................................................. 54 SAMPLE .................................................................................................................... 54 Scanning and image processing ................................................................................ 55 v Data collection and statistical analyses ..................................................................... 57 IV. RESULTS ............................................................................................................... 58 Qualitative observations ........................................................................................... 58 Sexual dimorphism in adult specimens ..................................................................... 58 Age-specific regional variation in bony organization and epithelial expansion .......... 60 Age-related changes in bony organization in each region.......................................... 62 Principal components analysis of measures of bone .................................................. 69 Age-related chagnes in epithelial expansion ............................................................. 71 V. DISCUSSION .......................................................................................................... 76 Morphology and variation ........................................................................................ 76 Changes in epithelial expansion during ontogeny ..................................................... 78 Timing of ontogenetic change .................................................................................. 79 Implications for phylogenetic analyses ..................................................................... 80 Implications for the epithelial hypothesis.................................................................. 81 VI. CONCLUSIONS ..................................................................................................... 82 Chapter Four: DIFFERENCES IN TEMPORAL BONE PNEUMATIZATION IN EXTANT PRIMATES I. INTRODUCTION ..................................................................................................... 83 II. BACKGROUND ...................................................................................................... 84 III. RESEARCH DESIGN ............................................................................................. 87 Sample ..................................................................................................................... 87 Scanning and image processing ................................................................................ 88 Data collection and statistical
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