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An Actualistic, CT-Based Study in Taphonomy

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An Actualistic, CT-Based Study in Taphonomy Heads and Skulls as Sediment Sorters: An Actualistic, CT-Based Study in Taphonomy A dissertation presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Doctor of Philosophy Joseph C. Daniel August 2012 © 2012 Joseph C. Daniel. All Rights Reserved. 2 This dissertation titled Heads and Skulls as Sediment Sorters: An Actualistic, CT-Based Study in Taphonomy by JOSEPH C. DANIEL has been approved for the Department of Biological Sciences and the College of Arts and Sciences by Lawrence M. Witmer Professor of Biomedical Sciences Benjamin M. Ogles Dean, College of Arts and Sciences 3 ABSTRACT DANIEL, JOSEPH C., Ph.D., August 2012, Biological Sciences Heads and Skulls as Sediment Sorters: An Actualistic, CT-Based Study in Taphonomy Director of Dissertation: Lawrence M. Witmer Reconstruction of extinct animals is hampered by lack of soft-tissue preservation. To test the hypothesis that sediment interacts with soft tissue during burial in predictable ways that may be of use in reconstructions, 29 ostrich heads in fresh, desiccated, rotten, or cleaned conditions were buried in two flumes; a short, deep-water flume with slow- moving water and a shallow-water flume with fast-moving water, designed to emulate common burial conditions in alluvial systems. After an initial CT scan, two heads from each setup (16 total) were reburied for seven months and then rescanned. Sediment was examined to confirm patterns detected by CT. Primary void space was retained in all conditions, especially within the tympanic recesses, increasing after prolonged burial, allowing crude estimates of burial conditions to be predicted. Sediment drapes overlay fresh and rotten heads in the shallow flume and to a lesser extent desiccated heads, but only rotten heads developed any drapes in the deep flume. Decompositional outgassing only partially disrupted external patterns in long-term-burial scans. Most internal soft- tissue patterns were obliterated by decomposition, although rostral conchae and auditory canals were still visible. Fat readily became adipocere, which is highly resistant to decay and can potentially be preserved as sediment traces. The best patterns were preserved in rotten or desiccated heads. These data indicate sediment patterns may be preserved in fossils that may reflect soft tissue and that may help to refine interpretations of fossils, 4 although not enough for detailed information for reconstructions. Void space origins and taphonomic implications for fossil preservation may be determined. Reassessments of published fossils include the preserved intestinal track of Scipionyx as a potential impacted bowel, preservation of a dinosaur “heart” is contraindicated, and possible conchae within Panoplosaurus. The presence of feathers on Sinornithosaurus and Sinosauropteryx is supported, but not strongly, and the chance of preserved internal organs in dinosaur mummies is low, although epidermal tissues and peripheral structures are high. Questions remain why some fossils show exceptional preservation while others in similar situations do not. Approved: _____________________________________________________________ Lawrence M. Witmer Professor of Biomedical Sciences 5 This work is dedicated to my wife Katharyn, without whose unflagging faith in me— even when I didn’t—and support this dissertation would never have been completed; and my children, Lorelai and Griffin, who provided the inspiration, hugs, and smiles to get me through. I would also like to dedicate this dissertation to my mother, for her gift of books and summers in the library; and my father, for seeing that I spent my youth in college, but did not live to see me finish. Today there remain but a few small areas on the world's map unmarked by explorers' trails. Human courage and endurance have conquered the Poles; the secrets of the tropical jungles have been revealed. The highest mountains of the earth have heard the voice of man. But this does not mean that the youth of the future has no new worlds to vanquish. It means only that the explorer must change his methods. —Roy Chapman Andrews, 1926 6 ACKNOWLEDGMENTS I am indebted to my advisor, Lawrence Witmer, for his guidance and aid throughout the project on both the science and the sociopolitical process of the career. I appreciate his patience with my idiosyncrasies and the roller-coaster life experiences as I dealt with the births, deaths, and other life experiences, both expected and unexpected, during my time at Ohio University. Thanks also to my supportive and unusually patient committee members, Patrick O’Connor, Susan Williams, and Greg Nadon. Thanks for the help and advice provided by Nancy Stevens and Don Miles and other faculty members that provided assistance. I would also like to thank Willem Roosenberg for serving on my comprehensive exam committee and, when asked later why he avoided certain topics, answering, “Because I already knew you knew all that.” I thank Dr. Gar Rothwell (OU Environmental & Plant Biology) for advice, assistance, and access to his thin-sectioning equipment and masonry saw and Dr. Mark Stoy of the Ohio Coal Research Center for advice on and use of the Malvern Mastersizer 2000. Special thanks go to Ryan Ridgely (graphics guru), and Casey Holliday, Tobin Hieronymus, Dave Dufeau, and Justin Tickhill of the Witmer lab for assistance in many aspects of the project. Thanks go to Angie Nielson for knowing everything about how to navigate the OU bureaucracy and getting things done. This work would not have been possible without access to the extensive collection of CT scans made available to me by Lawrence Witmer, for which I am very grateful. For loan of specimens for CT scanning and/or for providing existing CT datasets, I thank Mark Norell & Carl Mehling (American Museum); Michael Ryan, Bruce Latimer, & the 7 late Mike Williams (Cleveland Museum); and Matt Lamanna (Carnegie Museum). Thanks to Heather Rockhold, RT(CT), O’Bleness Memorial Hospital (Athens, OH), Ron Beshears (NASA Marshall Space Flight Center), and Tim Rowe & Rich Ketcham (UTCT, Austin) for CT scanning. Support for this work was provided by grants from Sigma Xi, the Jurassic Foundation, the Department of Biological Sciences and the Graduate Student Senate at Ohio University, as well as National Science Foundation awards IBN-9601174, IBN- 0343744, IOB-0517257 to LMW. Finally, I would like to thank my family for all their help. My wife, Katharyn, provided support and encouragement when I hit the inevitable moments of despair. My kids, Lorelai and Griffin, provided the joy and inspiration to see it through. I am indebted to my in-laws, John and Shannon Chamberlin for making all of this even possible and encouraging me—and so many others— to “follow your bliss.” 8 TABLE OF CONTENTS Page Abstract ............................................................................................................................... 3 Acknowledgments............................................................................................................... 6 List of Tables ...................................................................................................................... 9 List of Figures ................................................................................................................... 10 Chapter 1: Void space within buried ostrich heads and its implications for the interpretation of fossil patterns: an experimental, actualistic, CT-based study in taphonomy......................................................................................................................... 13 Chapter 2: Sediment/carcass interactions at the integument and their implications for fossil interpretation: A CT-based actualistic taphonomy study ........................................ 53 Chapter 3: The utility of CT-based, taphonomic study of sediment/carcass interactions for soft-tissue reconstruction of extinct organisms................................................................. 95 Chapter 4: New interpretations of fossils based on CT-based, actualistic taphonomy studies ............................................................................................................................. 158 References ....................................................................................................................... 209 Appendix A: Grain-size estimation techniques .............................................................. 230 Appendix B: Grain-size estimation in this study ............................................................ 234 Appendix C: Grain-size data ........................................................................................... 240 Appendix D: Thigh circumference estimation ................................................................ 343 9 LIST OF TABLES Page Table 1: Ohio University Vertebrate Collections (OUVC) specimens of ostrich used in this study, arranged by treatment condition for short-term burial set .............................51 Table 2: Standard flume sediment mixture .....................................................................51 Table 3: Statistical summary of void space comparisons ...............................................52 Table 4: Grain-size categories for sediment distribution estimates. ...............................94 Table 5: MANOVA results using full data set from both flumes ...................................94 Table 6: Endpoints for original void space within
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