The Interrelationship of the Premaxilla and the Mandibular Symphysis in Morphometric Terms and on a Wide Comparative Sample of Anthropoid Primates
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THE INTERRELATIONSHIP OF THE PREMAXILLA AND THE MANDIBULAR SYMPHYSIS IN ANTHROPOID PRIMATES Georgia Mortzou November 2003 Dissertation submitted to University College London in partial fulfilment of the requirements for the degree of Doctor of Philosophy ProQuest Number: 10011152 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest. ProQuest 10011152 Published by ProQuest LLC(2016). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code. Microform Edition © ProQuest LLC. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 ABSTRACT The premaxilla and mandibular symphysis have been traditionally studied in isolation, despite their anatomical vicinity and functionally complementary roles. The latter are particularly seen in the accommodation of incisors and the dissipation of masticatory forces. The diversity in the morphology of both premaxilla and mandibular symphysis has been viewed in the past as phylogenetically informative, but recent research has highlighted such a large degree of inherent variability, that its diagnostic validity is being seriously challenged. 50 inter-landmark distances, reflecting the upper and lower anterior masticatory complex, were collected from 26 species of extant anthropoid primates. Morphological correlations in the upper and lower masticatory complex were investigated using Principal Components Analysis. Discriminant Function Analysis (DFA) was employed to determine the extent to which the morphology of this region reflects diet. The contribution of phylogenetic inertia to morphological and dietary correlations was evaluated with Comparative Analysis by Independent Contrasts (CAIC). All analyses were run on both raw and size-adjusted data, except CAIC which was only carried out on size-adjusted data. Intraspecifically, no conclusive correlations were observed between the premaxillary and symphyseal regions. In DFA based on size-adjusted data of the full sample, distinct premaxillary and symphyseal morphologies were found to correspond to diet across phylogenetically diverse taxa. However, when DFA was run on size-adjusted data on each of the large taxonomic units in the sample (hominoids, cercopithecoids, ceboids) interpretation of the results was hampered by the inherent lack of dietary diversity within small taxonomie units and the strong effect of phylogeny. Dietary predictions on fossil taxa were in accordance with published accounts based on other lines of evidence. When the phylogenetic constraints were controlled for, only a small number of correlations between the areas of interest were identified. Small sample sizes, however, rendered the CAIC results inconclusive. Acknowledgments Though completing a thesis feels like (and proves to be) a rather solitary and lonely process, there are always numerous individuals who contribute to the final result, without necessarily sharing the views presented in it. In this respect, I would like to take this opportunity and express my gratitude to Professor Leslie Aiello for generating my interest in hominoid evolution with her truly inspiring lectures, and for -more importantly- providing valuable advice and guidance since the initial stages of this thesis, as well as instilling necessary discipline during the critical last months of its completion. I feel similarly indebted to Professor Peter Andrews for teaching me everything I know about fossil apes (a subject which provided inspiration for the present study), for showing me how to adopt broader perspectives and for always providing immediate and constructive comments on my writings. He and his family have constantly offered moral support, while they also welcomed me in their own home where I found guidance and a more than adequate library and cast collection to work in. I would like to further express my thanks to all the people responsible for the primate collections I worked in, gathering data: Ms Paula Jenkins, the Natural History Museum, London, UK; Mr John Harrison and Mr Malcolm Harman in the Powell-Cotton Museum, Birchington-on-Sea, Kent, UK; Dr Wim van Neer in the Royal Museum of Central Africa, Tervuren, Belgium; Dr Christian Smeenk in the Naturalis Museum, Leiden, The Netherlands; Dr Karin Isler in the Anthropology Department Collection, Zurich University, Zurich, Switzerland; Dr Olav Rohrer-Ertl in the Zoological and Anthropological State Collections of Munich, Germany; and Dr Richard Thorington and Ms Linda Gordon in the Smithsonian Institute, Washington D.C., USA. All these 4 individuals welcomed me in the collections under their care and provided useful advice on catalogue information, as well as more practical issues (such as accommodation). For all their help I am grateful. This fieldwork became largely possible due to generous financial support by: the Department of Anthropology (Research Fund), the Graduate School Travelling Fund, the Leakey Foundation Trust, and the Central Research Fund. I would like to further offer my thanks to Dr William Harcourt-Smith for help with the use of the Microscribe digitiser, and Dr Clare Holden for critical help and advice regarding the phylogenetic analysis and the use of the CAIC software. Similar thanks are due to the computer administrator of the Anthropology Department, UCL, Mr Chris Hagisawa. My heartfelt gratitude is due to my family, my parents and my brother, all of whom, though miles away have provided constant support and reminded me the reasons why I started this project in the first place, at times when justification for such actions eluded me. Finally, I would just like to remind Sam Cobb one more time that this project, even if it had been completed, it would have been of a very different form, had he not provided comments, help and moral support throughout the critical stages of data collection, analysis and, especially, writing up. TABLE OF CONTENTS List of Figures 9 List of Tables 12 CHAPTER 1 INTRODUCTION AND LITERATURE REVIEW i6 1.1 Introduction to the topic i6 1.2 Premaxilla i8 1.3 Mandibular symphysis 30 1.4 Incisal morphology 40 1.5 Synthesis 41 1.6 Aims of the study 45 1.7 Structure of the study 46 CHAPTER 2 MATERIALS AND METHODS 50 2.1 Introduction 50 2.2 Extant sample parameters 51 2.3 Fossil sample 54 2.4 Definition of measurements 56 2.5 Clearing of the data and size correction 63 2.6 Measurement error 64 2.7 Principal Components Analysis 65 2.8 Discriminant Function Analysis 66 2.9 Phylogenetic Comparative Analysis 72 CHAPTERS PRINCIPAL COMPONENTS ANALYSIS 87 3.1 Introduction 87 3.2 Results 88 3.2.1 Full Sample - Raw Data 88 3 .2.2 Full Sample - Size Corrected Data 92 3 .2 . 3 Hominoids - Raw Data 95 3.2.4 Hominoids - Size Corrected Data 98 3 .2 . 5 Cercopithecoids - Raw Data 101 3.2.6 Cercopithecoids - Size Corrected Data 103 3.2.7 Ceboids - Raw Data 106 3.2.8 Ceboids - Size Corrected Data 109 3 3 Discussion 111 3 4 Conclusions 117 CHAPTER 4 DISCRIMINANT FUNCTION ANALYSIS 154 4.1 Introduction 154 4-2 Results 155 4 .2.1 Full Sample - Size Corrected Data 155 4 .2.2 Hominoids - Size Corrected Data 159 4.2.3 Cercopithecoids - Size Corrected Data 162 4.2.4 Ceboids - Size Corrected Data 165 4.3 Discussion 167 4.4 Conclusions 173 CHAPTERS PHYLOGENETIC COMPARATIVE ANALYSIS 187 5.1 Introduction 187 5.2 Materials and Methods 187 5.3 Results 192 5.3.1 Analysis of continuous variables (Crunch) 192 5.3.2 Analysis of continuous and categorical variables 194 (Brunch) 5.4 Discussion 196 5.5 Conclusions 200 CHAPTER 6 ANALYSIS OF THE FOSSIL SAMPLE 209 6.1 Introduction 209 6.2 Results 210 6.2.1 Pliopithecus (Epipliopithecus) vindobonensis 210 6.2.2 Afropithecus turkanensis 214 6.2.3 Ankarapithecus meteai 219 6.2.4 Sivapithecus indicus 222 6.2.5 Ouranopithecus macedoniensis 224 6.2.6 Proconsul heseloni 226 6.2.7 Mesopithecus pentelicus 229 6.3 Discussion 231 6.4 Conclusions 241 7 CHAPTER 7 GENERAL DISCUSSION 276 7.1 Introduction 276 7.2 The association of premaxillary and symphyseal 278 morphologies 7.3 Directions for future research 296 7.4 SUMMARY OF CONFUSIONS 299 BIBLIOGRAPHY 301 LIST OF FIGURES CHAPTER 1 1.1 Diagrammatic overview of hominoid subnasal 49 morphology CHAPTER 2 2.1 Measurements employed in this study. 84 Cranium, anterior view 2.2 Measurements employed in this study. 85 Cranium, lateral view 2.3 Measurements employed in this study. 85 Mandible, lingual view of hemisection 2.4 Measurements employed in this study. 86 Palate, inferior view 2.5 Measurements employed in this study. 87 Mandible, superior view CHAPTERS 3.1a PCA: Full Sample (Raw Data). PCi v PC2 130 3 2 a PCA: Full Sample (Size-Corrected Data). PCi v PC2 131 3.2b PCA: Full Sample (Size-Corrected Data). PCi v PC3 132 3 -2C PCA: Full Sample (Size-Corrected Data). PC2 v PC3 133 3.3a PCA: Hominoids (Raw Data). PCi v PC2 134 3.3b PCA: Hominoids (Raw Data). PCi v PC3 135 3.3c PCA: Hominoids (Raw Data). PC2 v PC3 136 3.3d PCA: Hominoids (Raw Data). PCi v PC4 137 3.3e PCA: Hominoids (Raw Data). PC3 v PC4 138 3.4a PCA: Hominoids (Size-Corrected Data). PCi v PC2 139 3.4b PCA: Hominoids (Size-Corrected Data). PCi v PC3 140 3.4c PCA: Hominoids (Size-Corrected Data). PC2 v PC3 141 3.5a PCA: Cercopithecoids (Raw Data). PCi v PC2 142 3.5b PCA: Cercopithecoids (Raw Data). PCi v PC3 143 3.5c PCA: Cercopithecoids (Raw Data). PC2 v PC3 144 3.6a PCA: Cercopithecoids (Size-Corrected Data).