University of Calgary PRISM: University of Calgary's Digital Repository Graduate Studies The Vault: Electronic Theses and Dissertations 2019-08-12 Using the Price Equation to Quantify Species Selection and Other Macroevolutionary Forces in Cretaceous Molluscs Jordan, Katherine J. Jordan, K. J. (2019). Using the Price Equation to Quantify Species Selection and Other Macroevolutionary Forces in Cretaceous Molluscs (Unpublished master's thesis). University of Calgary, Calgary, AB. http://hdl.handle.net/1880/110722 master thesis University of Calgary graduate students retain copyright ownership and moral rights for their thesis. You may use this material in any way that is permitted by the Copyright Act or through licensing that has been assigned to the document. For uses that are not allowable under copyright legislation or licensing, you are required to seek permission. Downloaded from PRISM: https://prism.ucalgary.ca UNIVERSITY OF CALGARY Using the Price Equation to Quantify Species Selection and Other Macroevolutionary Forces in Cretaceous Molluscs by Katherine J. Jordan A THESIS SUBMITTED TO THE FACULTY OF GRADUATE STUDIES IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE GRADUATE PROGRAM IN BIOLOGICAL SCIENCES CALGARY, ALBERTA AUGUST, 2019 © Katherine J. Jordan 2019 Abstract Species selection and other macroevolutionary forces are challenging processes to study and quantify when using fossil data. Here, I used the Price equation to analyze changes in geographic range sizes prior to and during a mass extinction event to estimate the relative contribution of three macroevolutionary processes (species selection, anagenesis, and immigration). I also tested the hypothesis that larger geographic range size increases a group’s survivability during mass extinctions. I applied a similar method to Rankin et al. (2015) to study marine gastropods and bivalves of the Gulf and Atlantic Coastal Plain (originally studied by Jablonski (1987)) over the last 16 million years of the Cretaceous Period. I found three major changes in mean geographic range size shared by both gastropods and bivalves during the end- Cretaceous: an increase in mean range size during the late Campanian, a decrease in the mid- Maastrichtian, and an increase near the end of the Cretaceous Period (late Maastrichtian). The Price Equation indicates that the late Campanian increase in geographic range size was attributable primarily to immigration, the mid-Maastrichtian decrease was due to different combinations of the three processes (species selection, anagenetic change, and immigration) in gastropods and bivalves, and the late Maastrichtian increase was attributable to species selection. These changes in geographic range size coincide with a marine transgression event, a period of global climate change, and a marine regression event, respectively. A statistically significant correlation between larger geographic range size and increased survivability was found for one time increment (approximately four million years before the KPg boundary). This study shows that the relative contribution of interacting macroevolutionary processes fluctuated over the end- Cretaceous extinction event and suggests that large geographic range size can increase survivability under certain conditions leading up to a mass extinction. ii Keywords: species selection, anagenetic change, immigration, the Price equation, macroevolution, mass extinction. iii Acknowledgements I would like to start with an acknowledgement to those who have helped me in my professional development. A heartfelt thank you is much overdue to my advisor Dr. Jeremy Fox whose patience and kindness have gotten me to this point. I am appreciative of every bit of advice, cooking, and funding I have received as member of his lab. I hope that my hard work and future success will be the biggest demonstration of my thanks, Jeremy. I would also like to thank Dr. Jessica Theodor for being the role model I needed. Her strength and passion are inspiring. I am grateful for all the work she has done to help me become the paleontologist I dreamed I could be. A thank you to Dr. Mindi Summers and Dr. Charles Henderson, as well, for enhancing my invertebrate knowledge to make this thesis possible. To my twin sister, Rebecca: Her wisdom and love have helped me immensely. I am so glad you are my sister. To my undergraduate professor Dr. Brian Penney: It was Dr. Penney who found out about the graduate opportunity here in Calgary. In a way, I hope I have carried on his legacy by attending graduate school in Alberta as he did. Thank you so much, Dr. Penney. My friends here in Canada Selina, Colby, Rachel, and all the rest (from campus and Telus Spark): you all got me through and were more kind than I deserved at times. I appreciate you more than you will ever know. I would also like to extend my gratitude to my mother, Vicki, my partner, Aaron, my dearest friends, Caitlyn, Jess, and Matt. I am the best person I can be because of you all. Thank you for all your support in the last two years. I hope to make you all proud. iv Dedication This thesis is dedicated to my grandparents, Leroy and Dorothy Weed of Stonington, ME. Thank you for giving me a love of the ocean that has stayed with me and brought me, ironically, to the prairies of Alberta. v Table of Contents Abstract .............................................................................................................................. ii Acknowledgements .......................................................................................................... iv Dedication ...........................................................................................................................v Table of Contents ............................................................................................................. vi List of Tables and Equations ........................................................................................ viii List of Figures and Illustrations ..................................................................................... ix Epigraph .............................................................................................................................x CHAPTER ONE: INTRODUCTION ..............................................................................1 1.1: Macroevolution and the Price Equation………………………………………1 1.2: The Cretaceous Mass Extinction and Cretaceous Molluscs…………………10 CHAPTER TWO: METHODS ......................................................................................13 2.1: The Price Equation…………………………………………………………. 13 2.2: Dataset Selection and Preparation for Analysis……………………………. 16 CHAPTER THREE: RESULTS ....................................................................................22 3.1: The Price Equation Results………………………………………………….22 3.2: Statistical Analysis of Datasets……………………………………………. 26 3.3: Further Analysis of Datasets………………………………………………...27 CHAPTER FOUR: DISCUSSION .................................................................................29 4.1: Macroevolutionary Change Within Time Increment T2-T3………………...29 4.2: Macroevolutionary Change Within Time Increment T6-T7………………...30 4.3: Macroevolutionary Change Within Time Increment T7-T8………………...32 4.4: Periods of Macroevolutionary Stasis in the Data……………………………33 4.5: Differential Survivorship Among Cretaceous Molluscs…………………….35 4.6: Limitations of the Study……………………………………………….….... 38 CHAPTER FIVE: CONCLUSION ................................................................................41 REFERENCES .................................................................................................................43 APPENDIX 1: Table of gastropod ancestor-descendant pairs of the Gulf and Atlantic Coastal Plain. Species names and ranges from Hunt et al. (2005) Appendix B………………….51 APPENDIX 2: Table of bivalve ancestor-descendant pairs of the Gulf and Atlantic Coastal Plain. Species names and ranges from Hunt et al. (2005) Appendix B………………….57 APPENDIX 3: Species used in PAST biostratigraphy analysis. Species were used based on their occurrence and presence in the formations used in the study. Data collected from the Paleobiology Database (PBDB)………………………………………………………….62 APPENDIX 4: Results of the PAST unitary association analysis. Maximal cliques presented from the unitary associated output……………………………………………………….66 vi APPENDIX 5: Gastropod dataset arranged in time bin for Price equation analysis…….67 APPENDIX 6: Bivalve dataset arranged in time bin for Price equation analysis……….77 APPENDIX 7: Gastropod Price equation analysis by time increment. The title “n/a” is indicative of those descendants without ancestors (i.e. immigrants) ……………………………….84 APPENDIX 8: Bivalve Price equation analysis by time increment. The title “n/a” is indicative of those descendants without ancestors (i.e. immigrants) ……………………………….109 vii List of Tables and Equations Table 1: Results of Permutation Test for Each Time Increment. P values are reported. Null hypothesis: geographic range size and number of descendants left per time bin are independent of each other……………………………………………………………………………………25 Equation 1: Modified from Rankin et al. (2015) ………………………………………….... 13 Equation 2: The extended Price equation modified from Rankin et al. (2015), eq. 2.1………14 viii List of Figures and Illustrations Figure 1: Total mean geographic range size change (blue line) gives the Price Equation answer for the gastropod analysis. In comparison, the three separate