The Biogeography and Macroevolutionary Trends of Late
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The Biogeography and Macroevolutionary Trends of Late Paleozoic Cephalopods in the North American Midcontinent Sea: Understanding the response of pelagic organisms to changing climate during the Late Paleozoic Ice Age By: Kayla Kolis Submitted to the graduate degree program in Ecology and Evolutionary Biology and the Graduate Faculty of the University of Kansas in partial fulfillment of the requirements for the degree of Master of Science. Chair: Bruce S. Lieberman Member: Christopher K. Beard Member: Kirsten Jensen Date Defended: 02 June 2017 The dissertation committee for Kayla Kolis certifies that this is the approved version of the following dissertation: The Biogeography and Macroevolutionary Trends of Late Paleozoic Cephalopods in the North American Midcontinent Sea: Understanding the response of pelagic organisms to changing climate during the Late Paleozoic Ice Age Chair: Bruce S. Lieberman Member: Christopher K. Beard Date Approved: 02 June 2017 ii Abstract: Geographic range is an important macroevolutionary parameter that is frequently considered in paleontological studies. Species distribution and geographic range size are determined by a variety of biotic and abiotic factors that are well known to affect the differential birth and death of species. Thus, considering how species distributions and geographic range sizes fluctuate over time can provide important insight into evolutionary dynamics across the geologic time scale. In this study, I examine how geographic range size and rates of speciation and extinction changed throughout the Pennsylvanian and Early Permian in the North American Midcontinent Sea in an important pelagic clade, the Cephalopoda, using Geographic Information Systems (GIS). This period is particularly interesting for biogeographic and evolutionary analysis because it is characterized by repetitive glacial cycles, a global transition from an icehouse to greenhouse climate during the Late Paleozoic Ice Age and sluggish macroevolutionary dynamics, i.e. low speciation and extinction rates, that have been repeatedly documented in studies of other marine invertebrate taxa from both Gondwana and Pangea. The analyses presented herein indicate that cephalopod species diversity fluctuated throughout the Pennsylvanian and Early Permian matching the findings of other studies that examined diversity components of the Late Paleozoic marine invertebrate fauna. However, contrary to studies that focused on benthic taxa, my analyses found that: mean geographic range size of cephalopod species did not change significantly through time, despite numerous climate oscillations throughout this period; and further, geographic range size did not correlate with macroevolutionary rates. This result suggests that pelagic organisms may have a different response to climate change than benthic organisms and thus additional consideration of this issue is needed. Finally, these analyses indicate that, in the case of cephalopods, macroevolutionary iii patterns during the Late Paleozoic were more ‘dynamic’ than previously characterized. Thus, the macroevolutionary patterns recovered may depend on the taxon analyzed. iv Author’s Disclaimer: All taxonomic actions in this work are hereby disclaimed for nomenclatural purposes, as recommended in Article 8 of the International Code of Zoological Nomenclature (ICZN, 1999). v Acknowledgements: There are several people that I would like to thank for their guidance and support throughout my academic career. First and foremost, I would like to thank my family. Without their love and support, I would not have been able to accomplish my academic goals or been able to effectively balance being both a student and a mother. I owe my success to them. Next, I would like to thank my advisor, Dr. Bruce Lieberman. His expertise in macroevolutionary theory and biogeography proved essential for providing the framework for my thesis project. His guidance and assistance on many aspects of graduate school life, ranging from; course-work, encouragement to volunteer for outreach activities, conversations on evolutionary theory, and critiques on grant proposals as well as other writing endeavors, proved essential to my success as a student at the University of Kansas. Further, I would like to thank Dr. Chris Beard & Dr. Kirsten Jensen for their willingness to serve as members of my thesis committee. Their perspective and advice proved essential in providing both structure and clarity to my thesis. Further, conversations with Dr. Julien Kimmig and Dr. Lucas Strotz were exceedingly useful for providing structure to my thesis. I would also like to thank my undergraduate advisor and mentor, Dr. Chris Widga, for his support and encouragement throughout my academic career. Finally, I would like to thank several graduate students for their moral support and friendship throughout my graduate school career, including; Theo Kermit, Che-Ling Ho, Emily Arsenault, and Kaila Colyott. Their unique perspectives and expertise in different fields of biology and challenged me to consider my thesis project in innovative ways. vi Table of Contents: Abstract .......................................................................................................................................... iii Author’s Disclaimer ........................................................................................................................ v Acknowledgements ........................................................................................................................ vi Table of Contents .......................................................................................................................... vii List of Figures ............................................................................................................................... xii List of Tables ............................................................................................................................... xiv Chapter 1: The Biogeography and Macroevolutionary Trends of Late Paleozoic Cephalopods in the North American Midcontinent Sea ........................................................................................... 1 Introduction ................................................................................................................................. 1 Background ................................................................................................................................. 4 Geologic Setting of the LPIA in the Midcontinent Region of North America: ...................... 4 Paleoecology of the North American Midcontinent Sea ........................................................ 7 Previous Work ............................................................................................................................ 9 Geographic Range Size and its Association with Various Macroevolutionary Phenomena .. 9 Cephalopods of the North American Midcontinent Sea ....................................................... 12 Materials & Methods ................................................................................................................ 15 Data Collection ..................................................................................................................... 16 Range Reconstruction ........................................................................................................... 18 Analysis of fossil record bias ................................................................................................ 20 Statistical Analyses ............................................................................................................... 22 Results ....................................................................................................................................... 24 Paleoecologic Patterns in Late Paleozoic Cephalopods from the Midcontinent Sea............ 24 Paleobiogeographic Patterns in Late Paleozoic Cephalopods from the Midcontinent Sea .. 28 Analysis of Macroevolutionary Rates in Late Paleozoic Cephalopods from the Midcontinent vii Sea ......................................................................................................................................... 29 The Relationship between Biogeography and Macroevolutionary Rates of Late Paleozoic Cephalopods .......................................................................................................................... 32 Discussion ................................................................................................................................. 33 Conclusion ................................................................................................................................ 38 Bibliography ............................................................................................................................. 39 Chapter 2: Systematic Paleontology of Ammonoidea .................................................................. 51 Description of Ammonoidea ..................................................................................................... 52 Comparison of Ammonoid & Nautiloid Morphology & Behaviors ......................................... 53 Notes on Species Determinations ............................................................................................. 55 Repositories............................................................................................................................... 56 Order: Goniatitida