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Evolutionary Dynamics of Speciation and Extinction

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Evolutionary Dynamics of Speciation and Extinction Scholars' Mine Doctoral Dissertations Student Theses and Dissertations Fall 2015 Evolutionary dynamics of speciation and extinction Dawn Michelle King Follow this and additional works at: https://scholarsmine.mst.edu/doctoral_dissertations Part of the Biophysics Commons Department: Physics Recommended Citation King, Dawn Michelle, "Evolutionary dynamics of speciation and extinction" (2015). Doctoral Dissertations. 2464. https://scholarsmine.mst.edu/doctoral_dissertations/2464 This thesis is brought to you by Scholars' Mine, a service of the Missouri S&T Library and Learning Resources. This work is protected by U. S. Copyright Law. Unauthorized use including reproduction for redistribution requires the permission of the copyright holder. For more information, please contact [email protected]. 1 EVOLUTIONARY DYNAMICS OF SPECIATION AND EXTINCTION by DAWN MICHELLE KING A DISSERTATION Presented to the Faculty of the Graduate Faculty of the MISSOURI UNIVERSITY OF SCIENCE AND TECHNOLOGY and UNIVERSITY OF MISSOURI AT SAINT LOUIS In Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHILOSOPHY in PHYSICS 2015 Approved by: Sonya Bahar, Advisor Ricardo Flores Nevena Marić Paul Parris Thomas Vojta 1 iii ABSTRACT Presented here is an interdisciplinary study that draws connections between the fields of physics, mathematics, and evolutionary biology. Importantly, as we move through the Anthropocene Epoch, where human-driven climate change threatens biodiversity, understanding how an evolving population responds to extinction stress could be key to saving endangered ecosystems. With a neutral, agent-based model that incorporates the main principles of Darwinian evolution, such as heritability, variability, and competition, the dynamics of speciation and extinction is investigated. The simulated organisms evolve according to the reaction-diffusion rules of the 2D directed percolation universality class. Offspring are generated according to one of three reproduction schemes. Mate choice dictates offspring placement, and it defines a species based on reproductive isolation (known as the biological species concept), while a globally enforced death process ensues within each generation. This system is shown to exhibit nonequilibrium, continuous phase transitions as a function of the individual death probability. The dynamical rules that enable phase transition and clustering behavior to transpire behavior is discussed, and a connection is drawn to another type of phase transition that arises by mate choice alone. Coalescent theory is then used to explore common descent in evolved phylogenetic tree structures at both the individual and cluster level. Finally, an extinction scenario is implemented where, after reaching a steady-state, a large population percentage is killed. Historical contingency is shown to play a major role in recovery from mass extinction at criticality. iv ACKNOWLEDGEMENTS I would first like to express my gratitude to Professor Sonya Bahar. Because of her, I was able to work on this amazing project, and graduate school has been an absolutely wonderful experience. She has helped me grow by showing me how to address all of life’s problems, both personal and professional, scientifically. I am forever changed. Thank you, Sonya, for being so wonderful, academically curious, brilliant, and strong-willed (because I now understand just how hard your job really is). I look forward to many more years of collaboration and friendship. I owe many thanks to my former lab partner and friend Dr. Adam Scott, who I worked with on various aspects of this project for several years. This work is an extension of Adam’s dissertation, and thus, working with him provided the foundation for this work to happen. Adam is an excellent teacher; he taught me how to build and program computers. Thank you to Tera Glaze and Shane Meyer who provided valuable input on the content and helped to edit, and to Nevena Marić for explaining difficult mathematics and being on my dissertation committee. Also, I would like to thank my other committee members, Ricardo Flores, Paul Parris, and Thomas Vojta for their input and guidance at various stages of this work. Stephen Ordway, my soon-to-be-husband and my best friend, has provided help through this process that has been invaluable, from editing, to listening to me talk through ideas, to putting up with the ‘all hours of the night’ writing binges, to taking Blake to school every morning. Last but not least, I would not even be at this point if it was not for my mother, Ginger Granado, and my son, Blake King. Even though I did not go college out of high school, it never left me that my mother practically pleaded with me to go. She always made education the most important priority for me growing up. Finally, I would like to thank the funding sources for this work, The Missouri Research Board and the James S. McDonnell Foundation. v TABLE OF CONTENTS Page ABSTRACT……………………………………………………….…….…………………….………………………....….…iii ACKNOWLEDGEMENTS…………………………………….………………………………………………..…..….…iv LIST OF ILLUSTRATIONS………………………………….…….……………………………………………….…….viii LIST OF TABLES………………………….………………………….………………………….…………………………….x GLOSSARY………………………….…….……………………………………………………….……………………..……xi SECTION 1. INTRODUCTION…..……………………………………………………………………………………………..1 1.1 BIOLOGICAL BACKGROUND .............................................. ………………………...….1 1.1.1 Darwinian Evolution. .............................................................................. 2 1.1.2 Neutral Evolution. .................................................................................. 2 1.1.2.1 Genetic drift. ............................................................................... 3 1.1.2.2 Ecological drift. ........................................................................... 3 1.1.3 Singular Evolutionary Viewpoints. .......................................................... 8 1.1.3.1 Gene’s eye view. ........................................................................ 8 1.1.3.2 Individual level. .......................................................................... 9 1.1.3.3 Group selection. ....................................................................... 10 1.1.4 The Synthesis of Hierarchical Thinking: Multi-level Evolution. ............ 13 1.1.5 Phyletic Gradualism v. Punctualism. ..................................................... 20 1.1.6 The Hypothesis of Historical Contingency. ........................................... 28 1.2 PHYSICAL BACKGROUND ................................................................................ 31 1.2.1 Phase Transitions. ................................................................................ 31 1.2.1.1 Classification. .......................................................................... 32 1.2.1.2 Universality. ............................................................................ 33 1.2.2 Ordinary, Continuous, and Directed Percolation. ............................... 33 1.3 DISSERTATION SUMMARY ............................................................................. 38 1.3.1 Chapter Two. ....................................................................................... 38 vi 1.3.2 Chapter Three. .................................................................................... 38 1.3.3 Chapter Four. ...................................................................................... 38 1.3.4 Chapter Five. ....................................................................................... 38 2. PHASE TRANSITIONS IN A NEUTRAL EVOLUTION MODEL ...................................... 39 2.1 INTRODUCTION ................................................................................................ 39 2.2 THE MODEL ...................................................................................................... 44 2.2.1 Birth Process. ......................................................................................... 44 2.2.1.1 Reproduction schemes .............................................................. 44 2.2.1.2 Offspring dispersion. ................................................................. 44 2.2.2 Death Processes. .................................................................................... 45 2.2.3 Clustering. .............................................................................................. 46 2.3 ANALYTIC METHODS ....................................................................................... 47 2.3.1 Continuum Percolation. ......................................................................... 47 2.3.2 Nearest Neighbor Index, R. .................................................................... 48 2.4 RESULTS………………………………………………………………………………………………………49 2.5 DISCUSSION……………… .................................................................................... 56 3. MULTI-LEVEL ANALYSIS OF PYHLOGENETIC TREE STRUCTURES ............................ 61 3.1 INTRODUCTION ................................................................................................ 61 3.1.1 Phylogenetic Trees.................................................................................. 61 3.1.2 Coalescent Theory. ................................................................................. 64 3.1.2.1 Wright-Fisher and Moran models. ............................................ 64 3.1.2.2 Times to most recent common ancestor .................................. 68 3.1.2.3 Other coalescent models. .........................................................
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