BIOSTRATIGRAPHY AND DIVERSITY PATTERNS OF CENOZOIC ECHINODERMS FROM FLORIDA By CRAIG W. OYEN A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2001 Walter and Norma Oyen, who have always I dedicate this to my parents, expressed absolute confidence in my ability to succeed and supported all my this without their endeavors without question or hesitation. I could not have done it realize. support through all these years and I appreciate more than they ACKNOWLEDGMENTS This project could not have been completed without the aid of many people. Most important is the assistance and direction given by my dissertation committee chair, Dr. Douglas S. Jones. He encouraged me to work on whatever suggestions to improve my topic(s) I found interesting, and simply gave me approach in order to answer any of those questions. He also made my time in Gainesville enjoyable academically and socially by introducing me to other faculty and students, inviting me to his home or restaurants for dinners, and participating in pick-up basketball games and intramural games for relaxation. He (along with Roger Portell) accompanied me on many fascinating fieldtrips in Florida and other locations (often in association with GSA meetings) that expanded my scientific and other perspectives greatly. I thank the other members of my dissertation committee, Drs. Randazzo, Hodell, MacFadden, and Mature, for participating in my research and providing guidance whenever I it to had this group of faculty asked for their help. I consider a pleasure have members participating on my committee because they only treated me with respect and they openly provided suggestions they believed would serve me best in the context of completing my research and degree. Manager, IP Division, FLMNH) for I also thank Roger Portell (Collection his extensive help in most aspects of this dissertation. He spent much time with me during all phases of the research to help me become familiar with the invertebrate paleontology collection in the FLMNH, aided in nearly all fieldwork excursions, and accompanied me on visits to other museums or personal fossil collections. He also supplied me with many critical publications from his personal myself. His contacts with library and found others that I was able to purchase for non-professional fossil collectors in Florida enabled me to gather data I may not have been able to access otherwise. Roger’s help lessened some of the stress associated with my research. Financial assistance to complete the dissertation was provided by many Society of sources. Grant proposals I submitted were funded by the Geological America, the American Federation of Mineralogical Societies, Sigma Xi, the R. Jerry Britt Jr. Paleobiology Award (of the Florida Museum of Natural History), the Gary S. Morgan Student Research Award (of the Florida Paleontological Society), and the Mitchell Hope Scholarship Award (of the Southwest Florida Fossil Club). Employment by the Department of Geological Sciences provided teaching assistantships (under Dr. A. F. Randazzo, Chair) and research assistantships (under Dr. G.H. McClellan). The Florida Museum of Natural History provided research assistantships (under Dr. D.S. Jones). Members of my family also contributed significant financial assistance to my endeavors, particularly my parents (Walter and Norma Oyen) and my sister (Valerie Oyen- Larsen). IV of Barbara and Reed Toomey, I am very grateful for the thoughtfulness Roger and Anne Portell, Douglas and Sheila Jones, Jewel Pozefsky, Kendall Fountain, and Richard Hulbert. They gave me a place to stay (usually for extended periods of time) while traveling from Georgia or Pennsylvania during the holidays, long weekends, or summer breaks, so that I could finish my work. Fieldwork assistance was provided by many people during both the modern echinoderm study and fossil echinoderm collection phase of the Department) for giving me dissertation. I thank Dr. Frank Mature (Zoology guidance, equipment, and access to Seahorse Key during my study of the modern echinoids around the island. Dr. Douglas Jones, Roger Portell, and Kevin Schindler (FLMNH) helped construct the enclosures on site and also accompanied me during periodic visits to gather data. Field assistance on this part of the project (in addition to that listed above) was provided frequently by Karen Powers, Rich and Nicole Hisert, Len Fishkin, Kendall Fountain, and Ross Russell (all fellow students at UF). Assistance with fossil collecting was dominated by Dr. Douglas Jones, Roger Portell, and Kevin Schindler (of the FLMNH). A number of individuals made their personal fossil collections available for examination or donated specimens to the Florida Museum of Natural History that, in turn, allowed me to incorporate the information into my dissertation. I would like to thank those people who provided a significant number of specimens. The individuals with an academic or research affiliation have that information included in the parentheses after their name: Dr. Burchard Carter (Georgia Southwestern V state University), Dr. Jonathon Bryan (Okaloosa-Walton Community College), Dr. Thomas Scott (Florida Geological Society), Drs. Emily and Harold Vokes (Tulane University), Dr. Lyle Campbell (University of South Carolina at Spartanburg), Harley Means (Florida Geological Survey), Dr. Sherwood Wise (Florida State University), Muriel Hunter (formerly of Coastal Petroleum Co.), Jules DuBar, Phil Whisler, Tim Cassady (deceased), Byron Shumaker, Wendy Conway, Gary Schmelz, and Charles Hewlett. Access to quarries and property is not a simple process any longer, and several mine operators or owners have been generous in permitting me and other paleontologists to collect from their sites. This study benefited from the assistance given by Larry Rogers (Limestone Products, Inc.), C.T. Williams (Florida Rock Industries), Tom Jones (formerly of DoLime Products, Inc.), Fred Pirkle (formerly of DuPont Corporation), Richard Brown (Quality Aggregates, Inc.), Hugh Cannon (formerly of Quality Aggregates, Inc.), and Jimmy Philman (Handyphil, Inc.). work for the 1 appreciate assistance in completing the photographic dissertation document, particularly when I was nearing completion of the manuscript and didn’t have time to do it myself. Two people, in particular, allowed this to occur more easily and rapidly: George Hecht (FLMNH) photographed many of the fossils and Terry Lott (FLMNH) printed most of the photos. at University of Florida for I thank many of my fellow students the providing entertainment and camaraderie while engaged in the academic, VI athletic, and other endeavors during my time here. In particular, this includes all the “Psychotic Basement Troll” intramural basketball team players (Jose Garrido, Kendall Fountain, Len Fishkin, Greg Ferris, Rich Hisert, Joe Stoner, Stan Crownover, Robin Graves, Chris Saum, Neil Johnson, and Dr. Douglas Jones, among others) over the years. We weren’t pretty, but we had fun. My physicians benefited financially from this activity as well, as a result of my multiple sports injury visits. Norma), brothers (Lance and Finally, I thank my parents (Walter and Mitch), and sister (Valerie) for never asking “why” I was doing the research, and only asked” when” they could come for graduation. VII TABLE OF CONTENTS page ACKNOWLEDGMENTS iii KEY TO SYMBOLS xi ABSTRACT xiii CHAPTERS 1 INTRODUCTION 1 General Overview 1 Previous Work 2 Purpose and Goals 5 Methods and Materials Studied 8 2 STRATIGRAPHY OVERVIEW AND GEOLOGIC SETTING 11 Introduction 'll Eocene Stratigraphy 12 Oligocene Stratigraphy 14 Miocene Stratigraphy 14 Pliocene Stratigraphy 16 Pleistocene Stratigraphy 17 3 SYSTEMATIC PALEONTOLOGY OF FLORIDA ECHINODERMS 19 Introduction 19 Class Echinoidea Fossils 21 Eocene Echinoids 21 Oligocene Echinoids 92 Miocene Echinoids 108 Pliocene Echinoids 150 Pleistocene Echinoids 220 viii Class Crinoidea Fossils 237 Lower Ocala Limestone Crinoids 237 Upper Ocala Limestone Crinoids 239 Class Asteroidea Fossils 241 Eocene Asteroids 241 Oligocene Asteroids 242 Miocene Asteroids 243 Pliocene Asteroids 244 Class Ophiuroidea Fossils 247 Eocene Ophiuroids 247 Miocene Ophiuroids 248 Pliocene Ophiuroids 249 4 ECHINODERM DIVERSITY PATTERNS AND BIASES 259 Taxonomic and Biostratigraphic Discussion 259 Eocene Echinoids 260 Oligocene Echinoids 265 Miocene Echinoids 268 Pliocene Echinoids 272 Pleistocene Echinoids 276 Crinoids 277 Asteroids 279 Ophiuroids 282 Biases in the Cenozoic Echinoderm Record 284 Resolution of Data 284 Stratigraphic Nomenclature 285 Outcrop Exposure and Relief 286 Carbonate Versus Siliciclastic Environments 288 Age of Stratigraphic Units 292 Epoch Duration 292 Taxonomic Nomenclature 295 Collector Bias 298 Substrate and Facies Preferences of Echinoids 299 The Eocene-Oligocene Diversity Change 308 Early Paleogene Oceanographic Conditions 310 Early to Middle Paleogene Transitions 311 Late Eocene Extinctions and Biogeographic Patterns 314 Chapter Summary 322 5 ALLOMETRIC HETEROCHRONY IN MELLITID ECHINOIDS: A CASE STUDY FROM FLORIDA 325 Preface to the Biometric Analysis 325 Original Research Objectives 325 Growth Study Procedure 326 IX Introduction and Heterochrony Overview 329 Materials and Methods 333 Materials Examined 333 Data Acquisition Methods 334 Biometric Traits Evaluated 336 Data Analysis Methods 339 Results 340 6 SUMMARY AND CONCLUSIONS 349 New Echinoderm Diversity Patterns 349 Taxonomic Implications