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NHS Dissertation4 Patterns and Processes in the Dental Evolution of North American Plesiadapiforms and Euprimates from the Late Paleocene and Early Eocene DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University By Naava Hadassah Schottenstein, B.S. Graduate Program in Evolution, Ecology and Organismal Biology The Ohio State University 2020 Dissertation Committee: John P. Hunter, Advisor Mark Hubbe Bryan Carstens Debbie Guatelli-Steinberg Copyright by Naava Hadassah Schottenstein 2020 Abstract This dissertation explores the radiation of Primates during the Paleogene. The first radiation of Primates began with the plesiadapiforms near the Cretaceous-Paleogene boundary and the second radiation marked the introduction and diversification of euprimates at the beginning of the Eocene. Questions surrounding these radiations include their general patterns of evolution, rates of dental evolution, and potential influences of abiotic and biotic drivers. I explore these questions in euprimates by presenting a study of the Tetonius- Pseudotetonius primate lineage. Rates of evolution and the roles of neutral and adaptive processes across this lineage remain unclear. Linking Tetonius and Pseudotetonius are a series of stratigraphic and morphologic intermediates revealing possible functional and developmental reorganization within the dentition. Notable changes involved a reduction of the P3 and the P4 became a robust tall-cusped tooth. I test whether neutral evolution can explain the phenotypic differences in the lineage, and whether P4 lost developmental association with P3 and became integrated with the molars. I calculate the rate of evolutionary differentiation, based on the ratio between inter- and intra-species variation in length and width of the premolars and molars, between lineage segments and the entire lineage. I test for correlations between teeth within lineage segments. Correlations between P3 and the molars diminished, whereas correlations between P4 and the molars increased. I found evidence of varying degrees of stabilizing selection in the lengths and ii widths of most tooth metrics and neutral evolution in P4 width. This suggests a trend towards P4 becoming integrated into the molar field and that rates of evolution and selective pressures vary through time. The following chapters focus on the radiation of the family Paromomyidae during the late Paleocene and early Eocene. Their focus is on two paromomyid genera, Ignacius and Phenacolemur, which provide a series of long-lived lineages that experienced episodes of biotic turnover and dietary shifts. The second chapter explores evolutionary modes and rates that produced the changes in the dentitions of Ignacius and Phenacolemur, and the influence of the biotic turnover events. I test whether directional selection can explain the morphological changes that occurred with a dietary shift and if the changes that occurred with no dietary shift is marked by stasis. A random walk was the dominant model, followed by an Ornstein-Uhlenbeck process and strict stasis. Often, differences in tooth metrics between species are explained by neutral evolution. Dietary shifts did not always produce directional modes, and stasis was not always the best model when no dietary shift occurred. The third chapter focuses on the Ignacius frugivorus – Phenacolemur citatus lineage and assesses whether morphological changes in the dentition can be explained by changing paleotemperatures. I compare the magnitude and timing of dental changes to the paleotemperature curve and found significant positive correlations with temperature over the entire lineage. I found positive correlations between tooth metrics during both warming periods and both positive and negative correlations during the cool period. iii These findings suggest that paleotemperature may have played an important role in the evolution of paromomyids, but exactly how remains unclear. iv Dedication I dedicate this dissertation to my daughter Rose, who’s individuality and sense of self is truly inspiring to me. To my husband Michael, who always shows me patience and kindness and who’s support has helped me get through this stage of my life. To my mother, father, and sister who have been there for me always. Our yearly fishing trips to the beach provided a necessary escape to look forward to. To my Uncle Larry Grimes, who inspired me with his love of science. Visiting him and his lab at Meredith University as a young girl is one of the reasons this dissertation was ever written. To David Goldstein, for being my first academic mentor and a friend during one of the most challenging times of my life. v Acknowledgements I would foremost like to thank my advisor, John Hunter, for his encouragement, ideas, support, and patience throughout my PhD. I can’t imagine ever finding an advisor that would have been better for me. His academic guidance throughout these years has been incredibly helpful and has helped me create a body of work, of which I can be proud. I would also like to thank Mark Hubbe for his support and time spent helping me work through so much of the statistics involved in this dissertation. The insights he provided me throughout the years was immensely helpful. I would like to thank Bryan Carstens and Debbie Guatelli-Steinberg for agreeing to be on my committee. I learned a great deal about teeth from Debbie, and I always appreciated her kindness and encouragement. I am thankful to Bryan for the fun times teaching Evolution and Mammalogy, and for allowing me opportunities to guest lecture and get creative in the Mammology lab. I would like to thank Mary Silcox for providing me with a huge portion of the data needed to make this dissertation possible. I am grateful to her for allowing me to visit her lab in Scarborough and for the hospitality and kindness she showed me during my visits. I would like to thank Kenneth Rose and Thomas Bown for their thorough work and publications on the Tetonius – Pseudotetonius lineage, which inspired a portion of vi this research. I thank Kenneth Rose, Johns Hopkins School of Medicine, Smithsonian National Museum of Natural History, and the University of Michigan Museum of Paleontology for allowing me to measure additional Tetonius – Pseudotetonius specimens housed in their collections. I also thank Will Clyde for his work and help in establishing sediment ages and correlating the sediments of the Clarks Fork and Bighorn Basins, which allowed me to estimate ages of individual Tetonius – Pseudotetonius specimens. vii Vita B.S. Biological Sciences, Wright State University ........................................... 2014 Graduate Teaching Associate, Department of Evolution, Ecology and Organismal Biology, The Ohio State University .................................................. 2014 to present Publications Schottenstein, N. H., Hubbe, M., Hunter, J. P. 2019. Modules and mosaics in the evolution of the Tetonius-Pseudotetonius dentition. Journal of Mammalian Evolution. Evolution doi: 10.1007/s10914-019-09488-3 Fields of Study Major Field: Evolution, Ecology and Organismal Biology viii Table of Contents Abstract .............................................................................................................................. ii Dedication .......................................................................................................................... v Acknowledgements ........................................................................................................... vi Vita .................................................................................................................................. viii List of Tables ................................................................................................................... xii List of Figures .................................................................................................................. xv Chapter 1. Modules and Mosaics in the Evolution of the Tetonius – Pseudotetonius Dentition ............................................................................................................................ 1 1.1 Introduction .................................................................................................................. 1 1.2 Materials and Methods ................................................................................................. 7 1.3 Results ........................................................................................................................ 15 1.4 Discussion .................................................................................................................. 22 1.5 Conclusion ................................................................................................................. 33 1.6 Figures ........................................................................................................................ 36 1.7 Tables ......................................................................................................................... 42 1.8 Literature Cited .......................................................................................................... 47 Chapter 2. Rates and Modes of Dental Evolution in Late Paleocene and Early Eocene Paromomyids ................................................................................................................... 56 2.1 Introduction ...............................................................................................................
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