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Simulating Speciation and Extinction from East And CONTINUOUS OR PULSE? SIMULATING SPECIATION AND EXTINCTION FROM EAST AND SOUTH AFRICAN FAUNA AT PLIO-PLEISTOCENE FOSSIL SITES Thesis Presented in partial fulfillment of the requirements for the degree Master of Arts in the Graduate School of The Ohio State University By Daniel Chad Peart, B.A. Graduate Program in Anthropology The Ohio State University 2015 Thesis Committee: Jeffrey K McKee, PhD, Advisor Mark Hubbe, PhD Debbie Guatelli-Steinberg, PhD Copyright by Daniel Chad Peart 2015 Abstract Fossil fauna at paleoanthropological sites provides evidence for speciation and extinction events throughout the Plio-Pleistocene. Regarding fauna, first and last appearance dates are temporally clustered around time periods that correlate with climatic shift. The Turnover-Pulse Hypothesis asserts climate change as the cause of punctuated speciation and extinction events. Contending that climate is the cause of first and last appearance of species may be spurious due to inherent sampling biases in the fossil record. Species divergence and extinction may be influenced by climate, but the ultimate cause of species turnover is unclear. This research project required compilation of datasets of first and last appearance dates from South Africa and east Africa. These datasets were used to derive rates of species turnover in order to program a model to simulate the fossil record. Development of a species turnover simulation was undertaken utilizing mathematical modeling software (Matlab). Continuous speciation and extinction was simulated over 3.2 million years for South African fauna and 4.4 million years for east African fauna. Simulated continuous speciation and extinction produces peaks of turnover similar to turnover-pulses. Therefore, the fossil record is unable to support the Turnover-Pulse Hypothesis’s reliance on climate change as a causal mechanism for speciation and extinction. Rather, patterns of first and last appearance dates that indicate peaks of species turnover are a product of biased sampling. ii Dedication Dedicated to my grandmother, Nancy Lou Peart who always believed in me, no matter how grand my ambitions. Your support and love helped shape me into the person I am today. You are, and will always be, remembered. iii Acknowledgements I extend my heartfelt gratitude to Dr. Jeffrey McKee, who allowed my continuation of his work on computer simulation of evolutionary processes, as well as, use of his datasets. Thank you for your guidance and continued commitment to my success. I am also thankful for the guidance and support of all faculty members in the Department of Anthropology at The Ohio State University. Your shared knowledge and continuing advice have made me a better scholar. A special thank you to Mark Hubbe for guidance on quantitative methods, logic, and evolutionary principles. Most importantly, thank you to my family. Mom, you have always pushed me to be the best I can be and achieve my dreams. Dad, you are the rock that supports our family and your continuing dedication to the dreams of your children is something I can only hope to one day live up to. Wes, you have always been a free-thinker, and I cannot thank you enough for helping to open my mind to possibilities I had never before considered. My family will always been an important part of my life. I love you all. iv Vita May 2002……………………………………………….…………….Brewer High School December 2006…………………………B.S. Radio-TV-Film, Texas Christian University May 2013…………………………….B.A. Anthropology, University of Texas, Arlington 2013 to present…………………………..Graduate Student, Department of Anthropology, The Ohio State University Fields of Study Major Field: Anthropology Specialization: Paleoanthropology v Table of Contents Abstract…………………………………………………………………………………....ii Dedication………………………………………………………………….…………..…iii Acknowledgements……………………………………………………………………….iv Vita…………………………………………………………………………...……………v List of Tables…………………………………………………………………......……..viii List of Figures…………………………......………………………………………...…....ix Chapter 1. Introduction…………......……………………………………………………..1 Climate and Environment…………………………………………………………3 Fossil Record……………………………………………………………………...7 Evolution and Speciation………………………………………………………….9 Phyletic Gradualism…………………………………………………………….....9 Punctuated Equilibrium………………………………………………………......10 Autocatalysis, Eurytopy, and Stenotopy………………………………………….11 Extinction…………………………………………………………………...........13 Migration………………………………………………………………….......….14 Modeling……………………………………………………………………...….15 Chapter 2. Materials…………………………………………………….………………..17 Chapter 3. Methods………………………………………………………………………20 vi Statistical Analyses……………………………………………………………....25 Chapter 4: Results……………………………………………………………………..…26 Tests between Observed and Simulated Data………………………………........27 Chapter 5: Discussion………………………………………………………………….…30 Limitations of Simulated Models………………………………………………...31 Chapter 6: Conclusion……………………………………………………………………33 References………………………………………………………………………………..34 Appendix A: Additional Tables………………………………………………………….42 vii List of Tables Table 1. Species Frequencies from Previous Publications……………......………………19 Table 2. P-Values for Kruskal-Wallis tests performed between observed and simulated data at 100,000 year intervals………………………………………………………………….28 Table 3. East African Fauna – First and Last Appearance Dates…………………………45 Table 4. South African Fauna – First and Last Appearance Dates………………………49 viii List of Figures Figure 1. Benthic d18O Enrichment through the Pliocene and Pleistocene………......……6 Figure 2. Fossil Record First and Last Appearance Dates – East Africa………………….21 Figure 3. Fossil Record First and Last Appearance Dates – South Africa………………...22 ix Chapter 1: Introduction Climate is constantly changing. Throughout the Pliocene and Pleistocene, global climate cycled through a spectrum from warm to cold and wet to dry (deMenocal, 1995). Cyclical climatic change impacts evolution and divergence of species since changes in long-term weather patterns are a major component of physical environment. Changes in a lineage’s physical surroundings may influence speciation and extinction. Species turnover has been interpreted from the fossil record as some fauna go extinct and other species appear. Through turnover of species, by means of speciation, extinction, and migration, different species will appear in the fossil record through time (Vrba, 1985). Change in physical environment is frequently cited as the ultimate cause of species turnover (Vrba, 1974, 1985, 2005). First appearances and last appearances of species in the fossil record are commonly interpreted as evidence in support of this turnover based on climatic shift; however, fossils are differentially preserved, and provide only a snapshot of current species temporally and spatially. Conclusions about species turnover based on the fossil record reflect differential preservation that may be accounted for by chance rather than by climatically driven punctuated events. The fossil record should be intensely scrutinized when attempting to answer research questions, especially those about evolutionary processes. Although inferences can be made from the fossil record, causes for past events are difficult to decipher. Vrba (1985) 1 proposed the Turnover-Pulse Hypothesis to explain speciation, extinction, and migration events as an outcome of climate change. While species turnover occurs continuously at a low rate, Vrba noted punctuated periods of increased turnover. These periods correlate with short spans of intense global climate shift. Through the Turnover-Pulse Hypothesis, Vrba contends that periods of increased turnover are responsible for a majority of speciation and extinction while species remain stable without climatic forcing. This thesis addresses causal mechanisms applied to first and last appearance dates in the fossil record. There are multiple mechanisms that could be responsible for patterns of faunal turnover. The goal of this inquiry is to investigate the potential of the fossil record as an interpretive proxy for climate induced species turnover. Was variance between rates of turnover in the fossil record indicative of climate shift, or are changes in frequency of first and last appearance dates of species due to chance? In order to test the viability of the fossil record as an indicator of species turnover, a model of speciation and extinction was developed. Models have been used to simulate complexity of species radiations, extinctions, and phylogenetic diversification (Raup et al., 1973; Hey, 1992; Rabosky and Lovette, 2008). Using rates of turnover observed in the fossil record, models simulating continuous and punctuated turnover were developed and compared to fossil frequencies of first and last appearances. If the fossil record was not significantly different than simulated continuous turnover, then apparent pulses of speciation and extinction that correlate with climatic shifts may have been the product of random chance. 2 Besides the turnover-pulse hypothesis, other scholars have cited problems with fossil record biases in drawing conclusions about catastrophic extinction of species (Signor and Lipps, 1982), and biodiversity changes through time (Smith, 2001). Understanding causal connections between physical environment and species turnover will bolster understanding of how researchers can use the fossil record as a proxy for past environmental processes. Climate and Environment Environment and climate may be conflated in evolutionary discourse. Environment encompasses all aspects of an organism’s physical
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