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Scannellaj0515.Pdf (12.45Mb) ONTOGENETIC AND STRATIGRAPHIC CRANIAL VARIATION IN THE CERATOPSID DINOSAUR TRICERATOPS FROM THE HELL CREEK FORMATION, MONTANA by John Benedetto Scannella A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Earth Sciences MONTANA STATE UNIVERSITY Bozeman, Montana April 2015 ©COPYRIGHT by John Benedetto Scannella 2015 All Rights Reserved ii ACKNOWLEDGEMENTS Funding for this research was provided by grants from the Doris O. and Samuel P. Welles Research Fund of the University of California Museum of Paleontology, the Theodore Roosevelt Memorial Fund of the American Museum of Natural History, the Fritz Travel Grant of the Royal Ontario Museum, the Jurassic Foundation, and the Evolving Earth Foundation. Funding for travel to conferences was provided by the North American Paleontological Convention Student Travel Grant, the Society of Vertebrate Paleontology Jackson School of Geosciences Student Member Travel Grant, and the MSU Letters and Science Student Travel Grant. Many thanks also to Dr. Jack Horner, the Museum of the Rockies, the Sands Brothers, and Gerry Ohrstrom for graduate student funding and support. I thank my committee members Dr. Jack Horner, Dr. David Varricchio, Dr. David Roberts, Dr. Mark Goodwin and my graduate representative, Dr. Brendan Mumey, for their support and guidance. Thanks to all the curators, collections managers, graduate students, and volunteers who have permitted me to access their museum collections and displays. I thank my fellow MSU students, the staff and volunteers at the MOR, the MSU Earth Sciences Department, and the many friends I have made through SVP for exciting discussions and help along the way. I thank my parents, Ben and Mae Scannella, for taking me to the AMNH so many times when I was little. Thanks also to Angela Scannella, Regina Johnson, Preston Stewart, P.J. Stewart, and to Fred and Linda Baker. Special thanks to my sister, Christine Stewart, for taking me to the library and helping me take out books about dinosaurs when I was three. Finally, I thank my wife, Kari Scannella, for her love and support throughout the completion of this dissertation. iii TABLE OF CONTENTS 1. INTRODUCTION TO DISSERTATION .......................................................................1 A Brief Review of Early Studies of Ontogeny ...............................................................2 A Brief Survey of Studies of Dinosaur Cranial Ontogeny .............................................4 A Brief History of Triceratops Systematics .................................................................12 Triceratops Ontogeny ...................................................................................................15 Variation in Stratigraphic Context ................................................................................16 Summary of Dissertation ..............................................................................................18 Literature Cited .............................................................................................................23 2. TOROSAURUS MARSH, 1891 IS TRICERATOPS MARSH, 1889 (CERATOPSIDAE: CHASMOSAURINAE): SYNONYMY THROUGH ONTOGENY .....................................................................32 Contribution of Author and Co-Authors ........................................................................32 Manuscript Information Page ........................................................................................33 Abstract ..........................................................................................................................34 Introduction ....................................................................................................................34 Institutional Abbreviations .....................................................................................35 Materials and Methods ..................................................................................................36 Results ...........................................................................................................................37 Ontogenetic Development of Parietal Fenestrae ...................................................37 Squamosal Elongation ...........................................................................................39 The Ontogenetic Spectrum of Triceratops ............................................................40 Epiparietal and Episquamosal Variation ................................................................43 Osteohistology: a Test of Ontogenetic Hypotheses ...............................................45 Discussion ......................................................................................................................46 Ontogenetic Variation .............................................................................................46 Alternative Hypotheses ...........................................................................................47 Implications for Dinosaur Diversity ......................................................................49 Systematic Paleontology................... .............................................................................49 Revised Diagnosis ......................................................................................49 Valid Species .............................................................................................50 Locality and Horizon (YPM 1820) ............................................................50 Synonyms ...................................................................................................51 "Torosaurus" utahensis Gilmore, 1946 .....................................................51 Remarks .....................................................................................................51 Conclusions ....................................................................................................................52 Acknowledgments..........................................................................................................53 Appendix 2.1: Sample of Triceratops Cranial Material Collected by MOR .........65 Literature Cited ..............................................................................................................67 iv TABLE OF CONTENTS – CONTINUED 3. 'NEDOCERATOPS': AN EXAMPLE OF A TRANSITIONAL MORPHOLOGY .............................................................................72 Contribution of Author and Co-Authors ........................................................................72 Manuscript Information Page ........................................................................................73 Abstract ..........................................................................................................................74 Background ..............................................................................................................74 Methodology/Principal Findings ..............................................................................74 Conclusions/Significance ..........................................................................................75 Introduction ....................................................................................................................75 Institutional Abbreviations ......................................................................................78 Results ...........................................................................................................................79 Reassessment of USNM 2412 ................................................................................79 Nasal and Nasal Horn ................................................................................79 Postorbital Horn Cores ...............................................................................80 Parietal Fenestrae .......................................................................................81 Squamosal Morphology .............................................................................81 Episquamosal Number ...............................................................................82 Discussion ......................................................................................................................83 Taxonomic Status of 'Nedoceratops hatcheri' ........................................................83 Taxonomic Status of 'Torosaurus latus' .................................................................84 Variation in Epiossification Number and Position ....................................84 The parietal of USNM 2412 as Intermediate between Triceratops and 'Torosaurus' .....................................................................88 Juvenile 'Torosaurus latus' ........................................................................91 Transitional Morphologies and Dinosaur Systematics ...........................................93 Conclusions ....................................................................................................................94 Materials and Methods ...................................................................................................96 Acknowledgements ........................................................................................................96 References ....................................................................................................................106 4. A
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