Predation, resistance, and escalation in sessile crinoids by Valerie J. Syverson A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy (Geology) in the University of Michigan 2014 Doctoral Committee: Professor Tomasz K. Baumiller, Chair Professor Daniel C. Fisher Research Scientist Janice L. Pappas Professor Emeritus Gerald R. Smith Research Scientist Miriam L. Zelditch © Valerie J. Syverson, 2014 Dedication To Mark. “We shall swim out to that brooding reef in the sea and dive down through black abysses to Cyclopean and many-columned Y'ha-nthlei, and in that lair of the Deep Ones we shall dwell amidst wonder and glory for ever.” ii Acknowledgments I wish to thank my advisor and committee chair, Tom Baumiller, for his guidance in helping me to complete this work and develop a mature scientific perspective and for giving me the academic freedom to explore several fruitless ideas along the way. Many thanks are also due to my past and present labmates Alex Janevski and Kris Purens for their friendship, moral support, frequent and productive arguments, and shared efforts to understand the world. And to Meg Veitch, here’s hoping we have a chance to work together hereafter. My committee members Miriam Zelditch, Janice Pappas, Jerry Smith, and Dan Fisher have provided much useful feedback on how to improve both the research herein and my writing about it. Daniel Miller has been both a great supervisor and mentor and an inspiration to good scholarship. And to the other paleontology grad students and the rest of the department faculty, thank you for many interesting discussions and much enjoyable socializing over the last five years. Many thanks to the other participants and instructors in the 2011 Paleobiology Database Intensive Workshop in Analytical Methods at Macquarie University, Sydney, Australia. The things I learned there, both concrete and intangible, would have been hard to find elsewhere. And belated thanks also go to Don Prothero and Joe Kirschvink, my undergraduate advisors, for giving me a jump start into paleontological research. Each of the chapters benefited from particular help from my collaborators and reviewers. Thanks to Karl Stanley for enabling data collection, and to Charles Messing for identification and editorial comments, in chapter II. J. M. Koniecki kindly provided access to his private collection of Paleozoic crinoid specimens for Chapter III. Anna Reed and Forest Gahn collected the data used in Chapter IV. Dr. Carlton Brett was of great help in providing guidance on Chapter V. Chapters II and III also benefited from helpful comments from the editors of, respectively, the Bulletin of Marine Science and Paleobiology, and thoughful reviews from Dave Meyer, Tatsuo Oji, and several anonymous reviewers. I would also like to thank the various foundations and programs that provided the funding for my graduate studies and the research projects presented herein. These include a Shell Graduate Fellowship; NSF grant National Science Foundation (DEB 1036393, 1036219; EAR iii 0824793); the National Geographic Society (NGS 8505-08; CRE Research Grant 9283-13); and a Rackham Dissertation Fellowship. And thanks, last and most, to my friends and family. To all my friends here in Ann Arbor, especially Emile Moacdieh and Abby Purens: I’m so glad I got to meet you, and that we’ve had the chance to do this together. To my dear friends from Caltech, many of whom are also embarking on their academic careers at this very moment: let’s go forth and conquer! To my parents, Rebecca Stickler and Bryan Syverson, thank you for seeding my mind with books on paleontology, convincing me I could do anything I wanted to do, and then supporting me through college so that I could have a chance to prove them right. To my brother Joe Syverson, thanks for making me want to be an awesome big sister. To Phyllis and Patrick Poon, thank you for all your support and for making Ann Arbor feel like home. And to my wonderful fiancé Daniel Poon, who has been a rock of patience and emotional support during my grad-school angst, and provided a much-needed outside perspective that continues to save me from academic tunnel-vision: thank you, love. iv Table of Contents Dedication .................................................................................................................................................................... ii Acknowledgments .................................................................................................................................................... iii List of Tables .............................................................................................................................................................. vii List of Figures .......................................................................................................................................................... viii List of Appendices .................................................................................................................................................... ix Abstract ......................................................................................................................................................................... x Chapter I. Introduction ........................................................................................................................................... 1 Predatory interactions in the fossil record ................................................................................................... 1 Escalation and marine revolutions ............................................................................................................ 2 Crinoids and antagonistic biotic interactions ............................................................................................. 3 General anatomy and natural history of the taxon .............................................................................. 3 Basic crinoid taxonomy .................................................................................................................................. 4 Paleozoic & post-Paleozoic history of the Crinoidea ........................................................................... 4 History of predation and parasitism in crinoids ................................................................................... 5 Defensive adaptations ..................................................................................................................................... 9 Summaries of the following chapters .......................................................................................................... 10 Figures ...................................................................................................................................................................... 13 Chapter II. Growth, injury, and population dynamics in the extant cyrtocrinid Holopus mikihe (Crinoidea, Echinodermata) near Roatán, Honduras ................................................................. 15 Abstract ................................................................................................................................................................... 15 Introduction .......................................................................................................................................................... 16 Growth rates and regeneration frequencies in other crinoid taxa ............................................... 17 Data sources and methods ............................................................................................................................... 19 Location of dives ............................................................................................................................................. 19 Methods of data collection .......................................................................................................................... 19 Results ..................................................................................................................................................................... 20 Size and spatial distribution ...................................................................................................................... 20 Growth and regeneration rates .................................................................................................................. 21 Discussion .............................................................................................................................................................. 23 Growth and life history................................................................................................................................. 23 Habitat and behavior ..................................................................................................................................... 25 Acknowledgements ............................................................................................................................................ 26 Figures ...................................................................................................................................................................... 27 Tables ....................................................................................................................................................................... 31 v Chapter III. Temporal