Dental Pathology, Wear and Developmental Defects in South African Hominins
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Dental pathology, wear and developmental defects in South African hominins IAN EDWARD TOWLE A thesis submitted in partial fulfilment of the requirements of Liverpool John Moores University for the degree of Doctor of Philosophy June 2017 Abstract Studying different types of dental pathology, wear, and developmental defects can allow inferences into diet and behaviour in a variety of ways. In this project data on these different variables were collected for South African hominins and compared with extant primates. The species studied include Paranthropus robustus, Australopithecus africanus, A. sediba, early Homo, Homo naledi, baboons, chimpanzees and gorillas. Macroscopic examination of each specimen was performed, with a 10X hand lens used to verify certain pathologies. Variables recorded include antemortem chipping, enamel hypoplasia, caries, occlusal wear, tertiary dentine, abscesses, and periodontal disease. Clear differences in frequencies were found in the different South African hominin species. Homo naledi displays high rates of chipping, especially small fractures above molar wear facets, likely reflecting a diet containing high levels of contaminants. Other noteworthy results include the high levels of pitting enamel hypoplasia in P. robustus molars compared to other species, likely due to a species-specific enamel formation property or developmental disturbance. The low rates of chipping in P. robustus does not fit with this species being a hard food specialist. Instead, the wear best supports a diet of low-quality tough vegetation. Australopithecus africanus likely had a broad diet, with angled molar wear, lack of caries, and high chipping frequencies supporting this conclusion. Seven new carious lesions are described, two from H. naledi and five P. robustus. Other, rarer, pathologies are also highlighted, including abscesses in an early Homo individual, root grooves caused by erosive wear in A. africanus and a case of amelogenesis imperfecta in a female chimpanzee. The main conclusion from this thesis as a whole is the substantial difference in frequencies of the different variables among hominin species, supporting the proposition that their diets differed substantially. ii Acknowledgments First and foremost I want to thank my director of studies Joel Irish for the opportunity to complete this research and for all his help and encouragement during the course of this project. I would also like to thank my second supervisor Isabelle De Groote for her guidance and advice over the last three years. Without both supervisor’s continuous support this thesis would not have been possible. I am indebted to various museum curators and researchers who were tremendously helpful during data collection. In particular, I would like to thank Lee Berger and Bernhard Zipfel from the University of the Witwatersrand, Stephany Potze from Ditsong Museum of South Africa and Inbal Livne from the Powell-Cotton museum. I would also like to thank John Hawks, Lucas Delezene, Jacopo Moggi-Cecchi, Sandra Mathews, and Martin Hausler for their help and hospitality during these trips. I am hugely thankful to Marina Elliott for taking additional photos and descriptions of specimens once I was back in the UK and to Matthew Skinner for access to micro-CT scans. I gratefully acknowledge Liverpool John Moores University for providing funding for this project. This was in the form of a stipend and lab equipment but also travel bursaries which allowed me to attend and present my work at conferences. I am also appreciative of the administrative staff, including Gill Beesley and David Jones, who helped with travel arrangements and purchasing equipment. I would like to thank my colleagues at Liverpool John Moores University, in particular the fantastic group of postgraduates who made this PhD so enjoyable. As well as my main project I have been fortunate enough to undertake side projects, for which I am extremely grateful to Eleanor Dove, Alessio Veneziano and Cal Davenport for these collaborations. I am extremely thankful to my family for all their support and encouragement. Lastly, I am eternally grateful to Clare Davies for her encouragement and support both before and during this project. iii Contents Abstract……….…………………………………………………………………………………………………………………….…………….. ii Acknowledgments…………………….………………………………………………………………………………………..…………….iii Contents page…………………………………………………………………………………………………………………………..………iv List of figures………………………………………………………………………………………..……………………….………………..viii List of tables………………………………………………….………………………………………………………………………………..…X Glossary……………………....…………………………………………………………………………………………………………………..Xi Chapter 1: Introduction .......................................................................................................................... 1 1.1. Structure of thesis ....................................................................................................................... 2 1.2. Justification of research ............................................................................................................... 3 1.2.1. Homo naledi ......................................................................................................................... 3 1.2.2. Australopithecus sediba ....................................................................................................... 3 1.2.3. Paranthropus robustus ......................................................................................................... 4 1.2.4. Australopithecus africanus ................................................................................................... 5 1.2.5. Early Homo ........................................................................................................................... 5 1.2.6. Comparative samples ........................................................................................................... 6 1.3. Project aims ................................................................................................................................. 6 1.3.1. Research questions to be addressed ................................................................................... 7 Chapter 2: Sites and species .................................................................................................................. 8 2.1. The cradle through time .............................................................................................................. 8 2.2. Sites.............................................................................................................................................. 9 2.2.1. Taung .................................................................................................................................... 9 2.2.2. Sterkfontein .......................................................................................................................... 9 2.2.3. Kromdraai ........................................................................................................................... 10 2.2.4. Coopers .............................................................................................................................. 11 2.2.5. Drimolen ............................................................................................................................. 11 2.2.6. Swartkrans .......................................................................................................................... 11 2.2.7. Makapansgat ...................................................................................................................... 12 2.2.8. Malapa ................................................................................................................................ 12 2.2.9. Rising Star ........................................................................................................................... 12 2.3. Site codes ................................................................................................................................... 13 2.4. Species classification ................................................................................................................. 13 2.4.1. Australopithecus africanus ................................................................................................. 14 2.4.2. Paranthropus robustus ....................................................................................................... 15 2.4.3. Early Homo ......................................................................................................................... 16 iv 2.4.4. Homo naledi ....................................................................................................................... 17 2.4.5. Australopithicus sediba ...................................................................................................... 17 2.4.6. Family Tree ......................................................................................................................... 17 2.5. Geology and deposition ............................................................................................................. 18 2.6. Archaeological record and preservation ................................................................................... 19 2.6.1. Taphonomy, excavation and curation ................................................................................ 19 Chapter 3: Dental pathology, developmental defects and wear ......................................................