GIS Based Biogeography of Cincinnatian (Upper Ordovician) Brachiopods with Special Reference to Hebertella A thesis presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Master of Science Brandon C. Klingensmith August 2011 © 2011 Brandon C. Klingensmith. All Rights Reserved 2 This thesis titled GIS Based Biogeography of Cincinnatian (Upper Ordovician) Brachiopods with Special Reference to Hebertella by BRANDON C. KLINGENSMITH has been approved for the Department of Geological Sciences and the College of Arts and Sciences by Alycia L. Stigall Associate Professor of Geological Sciences Howard Dewald Interim Dean, College of Arts and Sciences 3 Abstract KLINGENSMITH, BRANDON C., M.S., August 2011, Geological Sciences GIS Based Biogeography of Cincinnatian (Upper Ordovician) Brachiopods with Special Reference to Hebertella Director of Thesis: Alycia L. Stigall Geographic ranges of type-Cincinnatian brachiopod species were reconstructed using Geographic Information Systems in order to analyze geographic distributions through space and time. Area polygons were digitized around species occurrence points plotted on the Cincinnatian outcrop belt (Ohio, Kentucky, Indiana), resulting in almost 100 individual maps spanning four depositional sequences. Individual species displayed patterns of habitat tracking as well as range expansion and contraction in response to regional sea level fluctuations. Statistical results show that species that established large geographic ranges preferentially survived the influx of extra-basinal species during the Richmondian Invasion. A systematic revision of the North American articulated brachiopod genus Hebertella Hall and Clarke is proposed. Specimens representing 13 species were coded for morphologic character analysis. A single most parsimonious tree produced from analysis of character data shows the evolution of characters. Biogeographic patterns derived from the cladogram as well as those analyzed in the previous study suggest east to mid-continent dispersal of Hebertella species from the Middle to Late Ordovician. Approved: _____________________________________________________________ Alycia L. Stigall Associate Professor of Geological Sciences 4 Acknowledgments Research was funded in part by grants from the Ohio University Alumni Association and the Geological Society of America. Research assistance and access to types was provided generously by archivists and collection managers at numerous institutions including the Cincinnati Museum Center, the Orton Geological Museum at the Ohio State University, the Limper Museum at Miami University, the Carnegie Museum of Natural History, the American Museum of Natural History, The Academy of Natural Sciences, the Museum of Comparative Zoology at Harvard University, and the National Museum of Natural History. For good company in the lab and field I am grateful to Katilin Maguire and Kristen Everman. In my academic training I was inspired by the entire faculty in the Department of Geological Sciences at Ohio University and hope them the very best. For friendship and moral support during the writing of this thesis I thank John F. Taylor, Jeremy Bader, and our cronies back in PA. Thanks are also given to Tom Dutro and John Pojeta for help at the NMNH; Jim Loch, John Repetski, Dave Brezinski, Jim Miller, Joe Clark, Charlie Burger, and Bob Kervin for their good wishes. Naturally, I am deeply indebted to my advisor Alycia L. Stigall. She presents quality science, while demonstrating patience and an unparalleled love and mastery of the science in which she is a pioneer. I owe the gratitude of an advisee whose life has been forged by opportunities made possible by her generosity. I wish you and your family joy and happiness. To my family and close friends, I owe just about everything else. 5 Table of Contents Page Abstract ............................................................................................................................... 3 Acknowledgments............................................................................................................... 4 List of Tables ...................................................................................................................... 7 List of Figures ..................................................................................................................... 8 Chapter 1: Introduction ....................................................................................................... 9 Chapter 2: GIS Based Analysis of Brachiopod Paleobiogeographic Patterns Across the Maysvillian to Richmondian Transition ........................................................................... 11 Abstract ......................................................................................................................... 11 Introduction ................................................................................................................... 12 Background ................................................................................................................... 15 Geologic Setting ........................................................................................................ 15 The Biota ................................................................................................................... 20 The Richmondian Invasion and Extinction Interval ................................................. 20 Methods ........................................................................................................................ 21 Geographic Extent .................................................................................................... 21 Temporal Range ........................................................................................................ 23 Species Occurrence Information ............................................................................... 24 Range Reconstruction ............................................................................................... 25 Stratigraphic Biases .................................................................................................. 27 Statistical Analysis .................................................................................................... 27 Results ........................................................................................................................... 29 Sea-Level Versus Geographic Range ....................................................................... 29 Geographic Range Versus Survivorship ................................................................... 30 Habitat Tracking ....................................................................................................... 32 Discussion ..................................................................................................................... 35 Habitat Tracking vs. Speciation ................................................................................ 36 Geographic Range and Sea Level ............................................................................. 38 Geographic Range and Survival ............................................................................... 39 Conclusions ................................................................................................................... 40 6 Chapter 3: Phylogenetic Revision of the Middle and Late Ordovician Brachiopod Hebertella From North America ....................................................................................... 43 Abstract ......................................................................................................................... 43 Introduction ................................................................................................................... 43 Phylogenetic Analysis ................................................................................................... 44 Description of Genus ................................................................................................ 44 Investigated Taxa ...................................................................................................... 45 Characters and Character States ................................................................................ 47 Characters Analyzed ................................................................................................. 49 Parsimony Analysis .................................................................................................. 55 Results ........................................................................................................................... 57 Evolutionary and Biogeographic Implications ......................................................... 59 Systematic Paleontology ............................................................................................... 63 Chapter 4: Conclusions ..................................................................................................... 70 References ......................................................................................................................... 74 Appendix 1: Cincinnatian Brachiopod Database .............................................................. 84 Appendix 2: Geographic Area of Species Range Polygons............................................ 141 7 List of Tables Tables Page 1. Species based stratigraphic range chart……………………………………………...22