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A Revision of the Hadrosauridae (Reptilia: Ornithischia) And Their Evolution During the Campanian and Maastrichtian By Michael Keith Brett-Surman B.A. 1972, University of Colorado M.A. 1975, University of California M.A. 1979, Johns Hopkins University M.Phil. 1985, George Washington University A Dissertation submitted to The Faculty of The Graduate School of Arts and Sciences of The George Washington University in partial satisfaction of the requirements for the degree of the Doctor of Philosophy February 19, 1989 Dissertation directed by Dr. Nicholas Hotton III Professorial Lecturer of Geology and Research Curator of Fossil Vertebrates at the National Museum of Natural History Department of Paleobiology Smithsonian Institution 1 Copyright 1989 Michael K. Brett-Surman All Rights Reserved ACKNOWLEDGMENTS I wish to thank the following people for their help during the course of this study; Richard Fox, Phil Gingerich, James Jensen, Dan Chure, the late Ted White, the late Russell King, Don Lindsey, Joanne Lindsey, the late Robert Makela, James Madsen, Christopher McGowan, William Morris, John Storer, William Turnbull, the late C.C. Young, Joseph Gregory, Robert Long, Fred Collier, Jann Thompson, Laurel Collins, and Tim Collins. Special thanks are owed to David Berman, John Bolt, A. Gordon Edmund, Eugene Gaffney, Erle Kauffman, Harold E. Koerner, Guy D. Leahy, Halska Osmolska, Dale Russell, Samuel P. Welles, Robert Purdy, and Raymond Rye. I am particularly indebted to the following individuals for their generous time and effort in aiding me during this study by providing data, equipment, photographs, helpful discussions and for their active support when I became quite sick of typing the Nth revision. They are Don Baird, Jennifer Clark, John Horner, Nicholas Hotton III, Douglas A. Lawson, George Olshevsky, John Ostrom, Gregory S. Paul, Phil Currie, Peter Galton, John S. McIntosh, Chip Clark, Hans-Dieter Sues, and Ralph Chapman. A special note of praise is due for my dissertation committee. Nicholas Hotton III is the dissertation director and Anthony Coates, David Atkins and Daryl Domning are the main readers. 2 A Revision of the Hadrosauridae (Reptilia: Ornithischia) And Their Evolution During the Campanian and Maastrichtian ABSTRACT A new taxonomy of the hadrosaurian dinosaurs is developed by a review of characters previously used to define genera, and an original analysis of the postcranial elements. Fifty-six morphological characters are commonly used for defining the hadrosaurids, and these are evaluated in terms of new data from ontogeny, paleopathology, and postcranial studies. Features once used to define taxa are now evaluated as the result of ontogeny. Forty-eight genera of hadrosaurids are taxonomically evaluated using both a "lumped" and "split" taxonomy. Synapomorphies are given for the family, two subfamilies and five tribes. Both "cladistic" and "evolutionary" principles are applied. Hadrosaurid macroevolution is discussed in light of paleobiogeography, morphological evolution and preservational biases in the fossil record. A new hypothesis of hadrosaurid extinction is proposed and named "Niche Assimilation". Its possible effects on other theories of dinosaur extinction are examined. 3 TABLE OF CONTENTS Acknowledgments Abstract Page 1 Table of Contents Pages 2 List of Tables Pages 3 - 4 List of Plates Pages 5 - 8 List of Abbreviations and Acronyms Pages 8 - 16 Chapter 1 Introduction Pages 17 - 48 Chapter 2 n Original Analysis of Hadrosaurid Postcrania Pages 49 - 61 Chapter 3 A Review of Cranial Features Pages 62 - 78 Chapter 4 A Review of Postcranial Features Pages 79 - 84 Chapter 5 Redefining the Hadrosauridae Pages 85 - 96 Tables 6 - 10 Pages 97 - 137 Chapter 6 The Genera of Hadrosaurids Pages 138 - 174 Chapter 7 An Overview of Hadrosaurid Evolution Pages 175 -186 Chapter 8 Speculations on Hadrosaurid Extinction Pages 187 - 215 Appendices Pages 216 - 246 Bibliography Pages 247 - 266 Plates 4 LIST OF TABLES 1 - The ilia of hadrosaurines and lambeosaurines page 34 2 - Hadrosaurinae selected measurements pages 85-87 3 - Hadrosaurinae selected ratios pages 88-89 4 - Lambeosaurinae selected measurements pages 90-92 5 - Lambeosaurinae selected ratios pages 93-94 6 - Iguandontid and camptosaurid selected measurements and ratios, pages 95-96 7 - Iguanodontid versus hadrosaurid pectoral girdles pages 141-142 8 - Iguanodontid versus hadrosaurid forelimbs pages 143-145 9 - Iguanodontid versus hadrosaurid pelvic girdles pages 146-147 10 - Iguanodontid versus hadrosaurid hindlimbs pages 148-149 5 List of Plates 1 - Scapula, Coracoid, and Sternum 2 - Humerus, Radius, and Ulna 3 - Manus and Pes 4 - Ilia 5 - Pubes 6 - Ischia 7 - Femur, Tibia, and Tarsals 8 - Edmontosaurus skull 9 - Edmontosaurus restoration 10 - Anatotitan skull 11 - Anatotitan restoration 12 - Bactrosaurus skeleton 13 - Corythosaurus skeleton and restoration 14 - Kritosaurus skeleton 15 - Maiasaura restoration 16 - Parasaurolophus skeleton and restoration 17 - Prosaurolophus skeleton and restoration 18 - Tsintaosaurus skeleton 19 - Plot of the number of species per genus using a 'lumped' and 'split' taxonomy. 20 - The five clades of hadrosaurids, skulls and pelves. 6 List of Abbreviations For Plates and Tables ACF acromion fossa ACR acromion ridge ACT acetabulum AST astragalus CAL calcaneum CRF coracoid fossa CRL carpals CRR coracoid ridge DE dentary DLF deltoid Fossa DLP deltopectoral crest DLR deltoid ridge FEM femur FIB fibula FMH femoral head GL glenoid GTR greater trochanter H height HH humeral head HL/FL hindlimb/forelimb ratio HND sternal handle HUM humerus ILP iliac peduncle ISFH ischial foot - heel ISFT ischial foot - toe ISP ischial peduncle JU jugal L length LA lacrimal LTR lesser trochanter MC3 metacarpal III MC3/H metacarpal III/humerus ratio MT3 metatarsal III MT3/F metatarsal III/femur ratio NA nasal OBF obturator fossa OBP obturator process OLN olecranon notch OLP olecranon process PBH pubic blade height PBL pubic blade length PD predentary PDL sternal paddle PF prefrontal PM1 premaxilla - 1 PM2 premaxilla - 2 7 PO postorbital POEP post-orbital eye pocket POP post-acetabular process PP postpubis PRP pre-acetabular process PUB pubic blade PUN pubic neck PUP pubic peduncle QJ quadratojugal QU quadrate R/H radius/humerus ratio RAD radius RPL reflected premaxillary lips SCAP scapula SCBL scapular blade SHF shaft SQ sqamosal STB sternal body (paddle) STH sternal handle SUIC supra-iliac crest T/F tibia/femur ratio TIB tibia TR3 third trochanter TR4 fourth trochanter TRS tarsals U/H ulna/humerus ratio ULN ulna UNG ungual W width 8 List of Museum Acronyms American Museum of Natural History - AMNH New York British Museum (Natural History) - BM(NH) London Brigham Young University Museum - BYU Provo Carnegie Museum of Natural History - CM Pittsburgh Cleveland Museum of Natural History - CMNH Cleveland Museum of Natural History of the City of Denver - DMNH Ekalaka Museum - EM Ekalaka, Montana Field Museum of Natural History - FMNH Chicago Geol. Inst. Mongolian People's Republic - GSP Ulan Bator Inst. Vert. Palaeo. Palaeoanthro., Academic Sinica, Beijing - IVPP Los Angeles County Museum, Natural History - LACM Los Angeles Museum of Northern Arizona - MNA Flagstaff Museum National d'Histoire Naturelle - MNHN Paris Naatuurhistorisch Mus. Maastricht, Holland - NHMM National Museum of Natural Sciences - NMC Ottawa Palaeontological Institute, Academy of Natural Sciences - PIN, Moscow 9 Beijing Museum of Natural History - PMNH Beijing Palaeontological Museum - PMU.R Uppsala Royal Ontario Museum - ROM Toronto San Diego Museum of Natural History - SDMNH San Diego Senckenberg Museum, - SM Frankfurt am Main Texas Memorial Museum, - TMM Austin Tyrrell Museum of Palaeontology - TMP Drumheller University of Alberta, Dept. Geol. - UA Edmonton Museum of Paleontology, Univ. Calif. - UCMP Berkeley University of Michigan Mus. Paleontology - UMMP Ann Arbor University of Nebraska State Museum - UNSM Lincoln National Museum of Natural History, Smithsonian Institution - USNM,Washington, D.C. Peabody Museum of Natural History, Yale Univ. - YPM New Haven Peabody Museum of Natural History, Yale Univ., (Princeton Collection) - YPM(PU, New Haven) Inst. Paleobiol., Polish Acad. Sci. - ZPAL Warsaw 10 CHAPTER 1 INTRODUCTION The purpose of this study is to make an original analysis of hadrosaurid postcrania and its effect on hadrosaurid taxonomy at the generic level. A revised diagnosis of all supra-generic clades is based on these new data, and the origin of the family and its palaeobiogeography are discussed. Major evolutionary trends are examined and hadrosaurid extinction is reviewed in light of extinction of all dinosaurs. The morphological features used to define the hadrosaurids, in both the past and the present, are reviewed and discussed. New finds, especially in studies on the postcrania, establish the taxonomic validity of both cranial and postcranial characters and show how they relate to ontogeny. Forty-eight genera of the family Hadrosauridae are reviewed. Synapomorphies are provided for each recognized subfamily and for each of the five clades newly recognized in this report. Both "cladistic" and "evolutionary" principles are used to delineate clades. A new definition of each family, subfamily and tribe is given. An original overview of hadrosaurid paleobiology, paleobiogeography, and macroevolution is presented. A decline in diversity and numbers of individuals, from the Campanian through the Maastrichtian, is newly demonstrated to be consistent and independent of 'lumping'
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  • Competition Structured a Late Cretaceous Megaherbivorous Dinosaur Assemblage Jordan C

    Competition Structured a Late Cretaceous Megaherbivorous Dinosaur Assemblage Jordan C

    www.nature.com/scientificreports OPEN Competition structured a Late Cretaceous megaherbivorous dinosaur assemblage Jordan C. Mallon 1,2 Modern megaherbivore community richness is limited by bottom-up controls, such as resource limitation and resultant dietary competition. However, the extent to which these same controls impacted the richness of fossil megaherbivore communities is poorly understood. The present study investigates the matter with reference to the megaherbivorous dinosaur assemblage from the middle to upper Campanian Dinosaur Park Formation of Alberta, Canada. Using a meta-analysis of 21 ecomorphological variables measured across 14 genera, contemporaneous taxa are demonstrably well-separated in ecomorphospace at the family/subfamily level. Moreover, this pattern is persistent through the approximately 1.5 Myr timespan of the formation, despite continual species turnover, indicative of underlying structural principles imposed by long-term ecological competition. After considering the implications of ecomorphology for megaherbivorous dinosaur diet, it is concluded that competition structured comparable megaherbivorous dinosaur communities throughout the Late Cretaceous of western North America. Te question of which mechanisms regulate species coexistence is fundamental to understanding the evolution of biodiversity1. Te standing diversity (richness) of extant megaherbivore (herbivores weighing ≥1,000 kg) com- munities appears to be mainly regulated by bottom-up controls2–4 as these animals are virtually invulnerable to top-down down processes (e.g., predation) when fully grown. Tus, while the young may occasionally succumb to predation, fully-grown African elephants (Loxodonta africana), rhinoceroses (Ceratotherium simum and Diceros bicornis), hippopotamuses (Hippopotamus amphibius), and girafes (Girafa camelopardalis) are rarely targeted by predators, and ofen show indiference to their presence in the wild5.