The PRINCETON FIELD GUIDE to DINOSAURS 2ND EDITION PRINCETON FIELD GUIDES Rooted in field experience and scientific study, Princeton’s guides to animals and plants are the authority for professional scientists and amateur naturalists alike. Princeton Field Guides present this information in a compact format carefully designed for easy use in the field. The guides illustrate every species in color and provide detailed information on identification, distribution, and biology. Albatrosses, Petrels, and Shearwaters of the World, by Derek Onley Birds of Southern Africa, Fourth Edition, by Ian Sinclair, Phil and Paul Scofield Hockey, Warwick Tarboton, and Peter Ryan Birds of Aruba, Curaçao, and Bonaire by Bart de Boer, Eric Birds of Thailand, by Craig Robson Newton, and Robin Restall Birds of the West Indies, by Herbert Raffaele, James Wiley, Birds of Australia, Eighth Edition, by Ken Simpson and Nicolas Orlando Garrido, Allan Keith, and Janis Raffaele Day Birds of Western Africa, by Nik Borrow and Ron Demey Birds of Borneo: Brunei, Sabah, Sarawak, and Kalimantan, by Carnivores of the World, by Luke Hunter Susan Myers Caterpillars of Eastern North America: A Guide to Identification Birds of Botswana, by Peter Hancock and Ingrid Weiersbye and Natural History, by David L. Wagner Birds of Central Asia, by Raffael Ayé, Manuel Schweizer, and Common Mosses of the Northeast and Appalachians, by Karl B. Tobias Roth McKnight, Joseph Rohrer, Kirsten McKnight Ward, and Birds of Chile, by Alvaro Jaramillo Warren Perdrizet Birds of the Dominican Republic and Haiti, by Steven Latta, Coral Reef Fishes, by Ewald Lieske and Robert Meyers Christopher Rimmer, Allan Keith, James Wiley, Herbert Dragonflies and Damselflies of the East, by Dennis Paulson Raffaele, Kent McFarland, and Eladio Fernandez Dragonflies and Damselflies of the West, by Dennis Paulson Birds of East Africa: Kenya, Tanzania, Uganda, Rwanda, and Mammals of Europe, by David W. Macdonald and Priscilla Barrett Burundi, by Terry Stevenson and John Fanshawe Mammals of North America, Second Edition, by Roland W. Birds of East Asia: China, Taiwan, Korea, Japan, and Russia, by Kays and Don E. Wilson Mark Brazil Marine Mammals of the North Atlantic, by Carl Christian Kinze Birds of Europe, Second Edition, by Lars Svensson, Dan Minerals of the World, by Ole Johnsen Zetterstrom, and Killian Mullarney Nests, Eggs, and Nestlings of North American Birds, Second Birds of the Horn of Africa, by Nigel Redman, Terry Stevenson, Edition, by Paul J. Baicich and Colin J. O. Harrison and John Fanshawe Palms of Southern Asia, by Andrew Henderson Birds of India, Pakistan, Nepal, Bangladesh, Bhutan, Sri Lanka, and Parrots of the World, by Joseph M. Forshaw the Maldives, by Richard Grimmett, Carol Inskipp, and Tim The Princeton Field Guide to Dinosaurs, Second Edition, by Inskipp Gregory S. Paul Birds of Kenya and Northern Tanzania: Field Guide Edition, by Raptors of the World, by James Ferguson-Lees and David A. Dale A. Zimmerman, Donald A. Turner, and David J. Christie Pearson Seeds of Amazonian Plants, by Fernando Cornejo and John Birds of Melanesia: Bismarcks, Solomons, Vanuatu, and New Janovec Caledonia, by Guy Dutson Sharks of the World, by Leonard Compagno, Marc Dando, and Birds of the Middle East, by R. F. Porter, S. Christensen, and Sarah Fowler P. Schiermacker-Hansen Stars and Planets: The Most Complete Guide to the Stars, Planets, Birds of Mongolia, by Sundev Gombobaatar, Axel Bräunlich, Galaxies, and the Solar System, Fully Revised and Expanded and Sh. Boldbaatar Edition, by Ian Ridpath and Wil Tirion Birds of Nepal, by Richard Grimmett, Carol Inskipp, and Tim Trees of Eastern North America, by Gil Nelson, Christopher J. Inskipp Earle, and Richard Spellenberg Birds of New Guinea, by Thane K. Pratt and Bruce M. Beehler Trees of Panama and Costa Rica, by Richard Condit, Rolando Birds of Northern India, by Richard Grimmett and Tim Inskipp Perez, and Nefertaris Daguerre Birds of Peru, by Thomas S. Schulenberg, Douglas F. Stotz, Trees of Western North America, by Richard Spellenberg, Daniel F. Lane, John P. O’Neill, and Theodore A. Parker III Christopher J. Earle, and Gil Nelson Birds of the Seychelles, by Adrian Skerrett and Ian Bullock Whales, Dolphins, and Other Marine Mammals of the World, by Birds of Southeast Asia, by Craig Robson Hadoram Shirihai and Brett Jarrett The PRINCETON FIELD GUIDE to DINOSAURS 2ND EDITION GREGORY S. PAUL Princeton University Press Princeton and Oxford Copyright © 2010, 2016 by Gregory S. Paul Requests for permission to reproduce material from this work should be sent to Permissions, Princeton University Press Published by Princeton University Press, 41 William Street, Princeton, New Jersey 08540 In the United Kingdom: Princeton University Press, 6 Oxford Street, Woodstock, Oxfordshire OX20 1TR nathist.press.princeton.edu All Rights Reserved ISBN 978-0-691-16766-4 Library of Congress Control Number: 2016933929 British Library Cataloging-in-Publication Data is available This book has been composed in Galliard, Goudy and Optima Printed on acid-free paper. ∞ Designed by D & N Publishing, Baydon, Wiltshire, UK Printed in China 10 9 8 7 6 5 4 3 2 1 CONTENTS Preface 6 Acknowledgments 6 Introduction History of Discovery and Research 9 What Is a Dinosaur? 13 Dating Dinosaurs 14 The Evolution of Dinosaurs and Their World 15 Extinction 23 After the Age of Dinosaurs 24 Biology 25 General Anatomy 25 Skin, Feathers, and Color 33 Respiration and Circulation 34 Digestive Tracts 36 Senses 36 Vocalization 38 Disease and Pathologies 38 Behavior 39 Brains, Nerves, and Intelligence 39 Social Activities 39 Reproduction 40 Growth 45 Energetics 47 Gigantism 49 Mesozoic Oxygen 52 The Evolution—and Loss—of Avian Flight 53 Dinosaur Safari 54 If Dinosaurs Had Survived 55 Dinosaur Conservation 55 Where Dinosaurs Are Found 56 Using the Group and Species Descriptions 62 Group and Species Accounts Dinosaurs 69 Theropods 69 Sauropodomorphs 182 Ornithischians 239 Additional Reading 350 Index: Dinosaur Taxa 351 Formations 358 PREFACE If I were, at about age twenty as a budding paleoresearcher and me yet more reason to more fully achieve a long-term goal, to artist, handed a copy of this book by a mysterious time traveler, illustrate the skeletons of almost all dinosaur species for which I would have been shocked as well as delighted. The pages would sufficiently complete material is available. These have been used reveal a world of new dinosaurs and ideas that I barely had a hint to construct the most extensive library of side-view life studies of or had no idea existed at all. My head would spin at the revela- of dinosaurs in print to date. The result is a work that covers tion of the therizinosaurs such as the wacky feathered Beipiaosau- what is fast approaching two centuries of scientific investigation rus and at the biplane flying dromaeosaurids—or at the oversized into the group of animals that ruled the continents for over 150 shoulder spines of Gigantspinosaurus, the neck spines of Amargas- million years. Enjoy the travel back in time. aurus, the brow horns and atrophied arms of bulldog-faced Car- The author’s paleozoological website can be found at notaurus, the furry adornments of Tianyulong and Kulindadromeus, www.gspauldino.com, and includes a complete list of his techni- the bristly tail of Psittacosaurus, the bat-like membranous wings cal papers and other publications. of scansoriopterygids, and the often psychedelic frilled horns of the new stable of centrosaurine and chasmosaurine ceratop- sids. Even Triceratops has proven to have strange skin. It is a particular pleasure to at long last be able to restore the skeleton Acknowledgments of the once mysterious Deinocheirus, long known from only its colossal arms—the skull and the rest of its peculiar skeleton do A complaint back in the last decade on the online Dinosaur not disappoint. And who would have imagined it would become List by Ian Paulsen about the absence of a high-quality dino- possible to figure out the colors of some feathered dinosaurs? I saur field guide led to the production of the first edition of would note the new names for some old dinosaurs, including my this dinosaur guide, and its exceptional success, combined favorite, Giraffatitan. And that old Brontosaurus is back! There with the continuous flux of new discoveries and research, led would be the dinosaur-bearing beds with the familiar yet often to production of the second. Many thanks to those who have exotic names Tendaguru, Morrison, Nemegt, Great Oolite, Hell provided the assistance over the years that has made this book Creek, and Lance. Plus there are the novel formations, at least possible, including Peter Galton, Kenneth Carpenter, James to my eyes and ears, Yixian, Tiouraren, Dinosaur Park, Anacleto, Kirkland, Michael Brett-Surman, Philip Currie, Alex Downs, Fangyan, Portezuelo, and Maevarano. The sheer number of new John Horner, Xu Xing, Robert Bakker, Saswati Bandyopad- dinosaurs would demonstrate that an explosion in dinosaur dis- hyay, Rinchen Barsbold, Frank Boothman, David Burnham, coveries and research, far beyond anything that had previously Thomas Carr, Daniel Chure, Kristina Curry Rogers, Steven occurred, and often based on new high technologies, marked and Sylvia Czerkas, Peter Dodson, David Evans, James Farlow, the end of the twentieth century going into the twenty-first. Tracy Ford, Catherine Forster, John Foster, Mike Fredericks, Confirmed would be the paradigm shift already under way Peter Galton, Roland Gangloff, Donald Glut (whose encyclo- in the late 1960s and especially the 1970s that observed that pedia supplements made this work much easier), Mark Hallett, dinosaurs were not so much reptiles as they were near birds Jerry Harris, Scott Hartman, Thomas Holtz, Nicholas Hotton, that often paralleled mammals in form and function. Dinosaurs Hermann Jaeger, Peter Larson, Guy Leahy, Nicholas Long- were still widely seen as living in tropical swamps, but we now rich, James Madsen, Jordan Mallon, Charles Martin, Teresa know that some lived through polar winters so dark and bitterly Maryanska, Octavio Mateus, John McIntosh, Carl Mehling, cold that low-energy reptiles could not survive.