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Signatory Page The Pennsylvania State University The Graduate School College of the Liberal Arts PROCONSUL HESELONI FEET FROM RUSINGA ISLAND, KENYA A Thesis in Anthropology by Holly M. Dunsworth © 2006 Holly M. Dunsworth Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy December 2006 The thesis of Holly M. Dunsworth was reviewed and approved* by the following: Alan Walker Evan Pugh Professor of Biological Anthropology and Biology Thesis Adviser Chair of Committee Joan T. Richtsmeier Professor of Biological Anthropology George R. Milner Professor of Professor of Archaeological Anthropology John H. Challis Associate Professor of Kinesiology Nina G. Jablonski Professor of Biological Anthropology Head of the Anthropology Department *Signatures are on file in the Graduate School. ABSTRACT In the mid 1980s, teams from Johns Hopkins University and the National Museums of Kenya colleted a minimum of ten Proconsul heseloni individuals from the Kaswanga Primate Site (KPS), Rusinga Island, Kenya. Five of them have nearly complete right and/or left feet. The preservation and anatomy of the KPS tarsals and metatarsals are described and analyzed here. Bivariate and multivariate analyses of the linear proportions of the feet and hindlimbs of the individuals illuminate arboreal characteristics in the Proconsul foot. Because the KPS individuals died at different stages of development, their foot and hindlimb proportions through ontogeny are investigated. Comparative cross-sectional ontogenetic samples include 222 Pan troglodytes, 65 Gorilla gorilla gorilla, and 115 Macaca mulatta and adult samples include 15 Pan paniscus, 24 Colobus (angolensis, guereza, and badius), 10 Nasalis proboscis, and 12 Hylobates lar. Proconsul aligned with Colobus and Nasalis in overall proportions and with apes in certain metatarsal proportions. Dissimilarities to macaques and similarities to apes in growth patterns suggest Proconsul was not born with large feet like macaques and other non-hominoid primates. Thus, Proconsul may have shared life history traits with Pan and Gorilla like increased altriciality in infant locomotor and positional behaviors. In order to investigate strength properties of the metatarsals, the midshafts of a total of 29 Proconsul metatarsals from six individuals were scanned with high-resolution microCT. Biplanar radiography and the latex cast method were used to obtain cross- sectional properties of the metatarsals from an extant cross-sectional ontogenetic sample of 52 Macaca mulatta (260 metatarsals) and 74 Pan troglodytes (370 metatarsals). Regression plots showed much closer similarity between Proconsul and Macaca than Proconsul and Pan in overall strength properties and cross-sectional shapes when normalized for metatarsal length and for body size. However, the first metatarsal of Proconsul showed greater resistance to bending and torsional forces for its length than Macaca, which is more like Pan. Overall results support previous interpretations on partial and incomplete Proconsul pedal fossils, and on other parts of the postcranial skeleton, that Proconsul was a generalized arboreal quadruped that grasped tree branches with a long, strong hallux. iii TABLE OF CONTENTS List of Tables………………………………………………………………………….....vi List of Figures…………………………………………………………………………..viii Acknowledgments ………………………………………………………………………xii Chapter 1. INTRODUCTION……………………………………………………….…….1 Specific Aims of the Dissertation..……………………………......………………1 Proconsul: State of the Taxon, 2006……………………………………………...2 Proconsul from the Kaswanga Primate Site………………………………………8 Primate Foot Notes: A brief survey of evolutionary research…………………...10 The Proconsul Foot……………………………………………………………....18 Chapter 2. BACKGROUND……………………………………………………………..30 Catarrhine Positional Behavior: A review of the comparative sample………......30 Significance of Ontogenetic Studies……………………………………..............37 Rhesus Macaque Growth and Locomotor Ontogeny……………………….........40 Chimpanzee and Gorilla Growth and Locomotor Ontogeny…………….............41 Chapter 3. NEW PROCONSUL HESELONI TARSALS AND METATARSALS FROM THE KASWANGA PRIMATE SITE..………………….…………………………...44 The Kaswanga Primate Site……………………………………………………...44 Taphonomy………………………………………………………………………47 Age, Sex and Grouping of the Individuals..………………………………….….47 Descriptions of KPS Tarsals and Metatarsals……………………………….…...49 KPS I………………………………………………………………….….51 KPS III…………………………………………………………………...63 KPS IV…………………………………………………………………...76 KPS VI…………………………………………………………………...86 KPS VII…………………………………………………………………..88 KPS VIII…………………………………………………………………96 Comparative Anatomy………………………………………………………….106 Chapter 4. FOOT AND HINDLIMB PROPORTIONS OF PROCONSUL HESELONI ………………………………………………………………………………… 113 Specific Aims……………………………………………………………….…..118 Materials and Methods………………………………………………………….119 Sample…………………………………………………………………..119 Age Estimation………………………………………………………….122 Body Size Estimation……………………………………………….…..123 Statistical Methods………………………………………………….…..126 Bivariate Results: Ontogenetic Sample………….…………………………..…133 Bivariate Results: Adult Sample………….…………………………………….170 Discussion of Bivariate Results……………………………………………..….186 Multivariate Results……………………………………………………….……191 Discussion of Multivariate Results……………………………………………..207 Chapter 5 – CROSS-SECTIONAL GEOMETRY OF PROCONSUL HESELONI METATARSALS THROUGH ONTOGENY…………...…………………………208 Metatarsal CSG in Primates………..……………………………….……….…211 Ontogeny of Cross-Sectional Geometry………………………………………..213 Specific Aims and Hypotheses…………………………………………………215 Materials and Methods…………………………………………………………216 Sample……………………………………………………………….…216 Age Estimation…………………………………………………………217 Body Size Estimation……………………………………………….….219 Cross-sectional Geometry Methods…………………………………....221 Microcomputed Tomography (microCT) of Fossil Specimens………..229 Statistical Methods…………………………………………………..…231 Results……………………………………………………………………….…232 Discussion……………………………………………………………………...250 Chapter 6 – SUMMARY, CONCLUSIONS & FUTURE RESEARCH………………256 Implications for Proconsul Behavior and Life History……………………….. 257 Problematic Proconsul Individuals…………………………………………… 260 Suggestions for Future Research……………………………...………………..262 Conclusion……………………………………………….…..…………………267 Literature Cited…………………………………………………………………………269 Appendix A: Abbreviations for Measurements…………………………..…..………...287 Appendix B: Statistical Data Tables………………..…………………….…...………..291 Appendix C: Raw Linear Measures for the Modern Comparative Sample………….....321 LIST OF TABLES 2.1. Body mass, locomotion and foot positions of taxa in the comparative collection….31 3.1. Lengths in millimeters of metatarsals of the KPS Individuals……………………...50 3.2. Measurements of intermediate and lateral cuneiforms of the KPS Individuals…….50 4.1 Adult specimens and their locations…………………………………………… ….119 4.2 Sample size by age group for growth sample…………………………………… ...121 4.3 Proconsul individuals in the sample…………….……………………………… …121 4.4 Age groups constructed according to dental eruption stages……………………….123 4.5 Body mass estimates for the sample………………………………………………..124 4.6. ANOVA for Tibia/Femur by age group……………………………………………137 4.7 ANOVA for Tarsus/ Foot by age group…………………………………………....139 4.8 ANOVA for MT3/ Foot by age group…………………………………………… ..141 4.9 ANOVA for MT3/ Calcaneus by age group……………………………………… .143 4.10 ANOVA for PH3/ Foot by age group………………………………….……….....145 4.11 ANOVA for PH3/ Calcaneus by age group……………………………………….148 4.12 ANOVA for Power Arm/ Calcaneal Load Arm by age group…………………….150 4.13 ANOVA for Power Arm/ Pedal Load Arm by age group………………………...152 4.14 ANOVA for Foot/ Femur by age group…………………………………………...155 4.15 ANOVA for MT1/ Foot by age group…………………………………………….157 4.16 ANOVA for MT1/ Calcaneus by age group………………………………………160 4.17 ANOVA for PH1/ Foot by age group……………………………..………………162 4.18 ANOVA for PH1/ Calcaneus by age group……………………….………………164 4.19 ANOVA for PH1/ PH3 by age group……………………………..………………166 4.20 ANOVA for MT1/ MT3 by age group……………………………………………168 4.21 Summary of Proconsul comparative proportions…………………………………190 4.22 Summary of multivariate results…………………………………………………..207 5.1 Number of individuals per age group ……………………………………………...217 5.2 Age groups constructed according to dental eruption stages……………………….218 5.3 Age group assignments for Proconsul heseloni individuals………………………..218 5.4 Body Masses estimates……………………………………………………………..219 5.5 Parallax by institution: ratio of mold diameter/ x-ray diameter…………………….227 5.6 Parallax by institution: percent enlargement………………………………………..227 5.7 Proconsul specimens scanned with microCT for the analysis……………………..230 5.8 Cross-sectional properties referenced and analyzed in this study………………….230 5.9 Error introduced for each method ………………………………………………….250 B.1. Pearson Correlations for metrics of entire extant sample (pooled ages)…………..291 B.2. Regression statistics for log-transformed metrics and indices of the ontogenetic sample of Gorilla gorilla ………………………………………………………………297 B.3 Regression statistics for log-transformed metrics and indices of the ontogenetic sample of Macaca mulatta ……………………………………………………………..299 B.4. Regression statistics for log-transformed metrics and indices of the ontogenetic sample of Pan troglodytes ……………………………………………………………..301 B.5 Regression statistics for Colobus hindlimb metrics versus femoral head breadth (FHB). ……………………………………………………………………………….…303 vi B.6. Regression statistics for Gorilla hindlimb metrics versus femoral head breadth (FHB) …………………………………………………………………………………..304 B.7. Regression statistics
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