OSTEOLOGY, PHYLOGENY, TAPHONOMY, AND ONTOGENETIC HISTOLOGY OF ORYCTODROMEUS CUBICULARIS, FROM THE MIDDLE CRETACEOUS (ALBIAN-CENOMANIAN) OF MONTANA AND IDAHO by L.J. Krumenacker A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Earth Sciences MONTANA STATE UNIVERSITY Bozeman, Montana May 2017 ©COPYRIGHT by Laurel James Krumenacker 2017 All Rights Reserved ii DEDICATION This dissertation is dedicated to my children, Elise and Shawn. You are everything to me and I will ALWAYS be here for you. The universe is full of wonderful unknowns. Find your own questions and seek the answers. Leave the world and the people you meet better than you found them. I love you. iii ACKNOWLEDGEMENTS Remember in “Jurassic Park” where the Tyrannosaurus eats a poorly dressed lawyer with thinning hair- that is the best scene… Despite all the opposition, I finished this PhD (and am working on cloning a T. rex). I am forever grateful to the people who walked me through these dark times: Cass Powell, you walked with me through the worst part of my life, being a true friend, helping me to find strength and worth in myself and reminding me how important it is to be active; thank you. Chancey Ringer, Liz Greenfield, Shardai Urdahl, Amanda Tebay, and all of FGHO, I genuinely wouldn’t have finished this degree without your patience, help, and friendship. Ash “Queen of Pine Bar” Ferguson, thanks for the encouragement and support. Thanks to my chair, Dr. Dave Varricchio, who helped me carve out a niche to work in the Cretaceous of Idaho, helped me learn to be a better scientist and writer, and who tirelessly helped me improve this work. Dr. Chris Organ and Dr. David Bowen provided valuable input and opportunities for improvement. Dr. Ted Dyman provided enjoyable learning in the field and input in all aspects of this project. Takuya Imai, Kelli Taddy, Eric Metz, and the crew of the Paleontology Field Course, thank you. Michael Holland, Bob Harmon, and Ellen Lamme, thank you for helping me in lab. Thanks to Haley Dunkel for friendship and employment for a large portion of my doctoral work. Steve Robison, Ali Abusaid, and Diane Wheeler provided permitting assistance while Leif Tapanila, Mary Thompson, and John Scannella allowed access to specimens. Josh Cotton provided awesome artwork for this dissertation and future publication. Thanks to Megan Maier for formatting help and Dan Lawver for computer help. iv FUNDING ACKNOWLEDGEMENT Funding for this research from the Jurassic Foundation and the Montana State University Graduate School were instrumental in aspects of this research and are gratefully acknowledged. Many thanks to Bob Simon for his generous personal donation that facilitated numerous aspects of this research as well. v TABLE OF CONTENTS 1. INTRODUCTION ...........................................................................................................1 Literature Cited ....................................................................................................................4 2. OSTEOLOGY AND PHYLOGENY OF THE NEORNITHISCHIAN DINOSAUR ORYCTODROMEUS CUBICULARIS FROM THE MIDDLE CRETACEOUS (ALBIAN-CENOMANIAN) OF MONTANA AND IDAHO..............................................................................................5 Contribution of Authors and Co-Authors .........................................................................5 Manuscript Information Page ...........................................................................................6 Abstract .............................................................................................................................7 Introduction .......................................................................................................................8 Geological Setting ...........................................................................................................10 Occurrence ......................................................................................................................11 Institutional Abbreviations ..............................................................................................12 Anatomical Abbreviations ..............................................................................................12 Terminology and Methodology ......................................................................................13 Systematic Paleontology .................................................................................................15 Description ......................................................................................................................18 Skull .......................................................................................................................18 Supraoccipital ............................................................................................19 Opisthotics and Exoccipital .......................................................................20 Basioccipital ...............................................................................................20 Prootic ........................................................................................................21 Basisphenoid and Parasphenoid .................................................................21 Laterosphenoid ...........................................................................................22 Premaxilla ..................................................................................................22 Maxilla .......................................................................................................23 Jugal ...........................................................................................................24 Supraorbital ................................................................................................25 Postorbital ..................................................................................................25 Frontal ........................................................................................................25 Quadrate .....................................................................................................26 Predentary ..................................................................................................26 Dentary .......................................................................................................27 Dentary Teeth.............................................................................................28 Axial Skeleton ........................................................................................................29 Cervical Vertebrae .....................................................................................29 Cervical Ribs ..............................................................................................30 vi TABLE OF CONTENTS CONTINUED Dorsal Vertebrae ........................................................................................31 Dorsal Ribs.................................................................................................33 Sacral Vertebrae .........................................................................................34 Caudal Vertebrae .......................................................................................35 Chevrons ....................................................................................................38 Ossified Tendons .......................................................................................38 Appendicular Skeleton ...........................................................................................40 Scapula .......................................................................................................40 Coracoid .....................................................................................................42 Humerus .....................................................................................................43 Ulna ............................................................................................................44 Radius ........................................................................................................44 Manus .........................................................................................................45 Ilium ...........................................................................................................46 Pubis ...........................................................................................................47 Ischium .......................................................................................................48 Femur .........................................................................................................49 Tibia ...........................................................................................................50 Fibula .........................................................................................................51 Tarsus .........................................................................................................51 Pes ..............................................................................................................52 Discussion .......................................................................................................................54 Phylogenetic Analysis and Review ........................................................................54