Table Of Content

The PRINCETON to FIELD GUIDE MESOZOIC SEA REPTILES 01 MMR – Intro pp001- 65.indd 1 16/05/2022 14:22 01 MMR – Intro pp001- 65.indd 2 16/05/2022 14:22 The PRINCETON to FIELD GUIDE MESOZOIC SEA REPTILES GREGORY S. PAUL Princeton University Press Princeton and Oxford 01 MMR – Intro pp001- 65.indd 3 16/05/2022 14:22 Copyright © 2022 by Princeton University Press Princeton University Press is committed to the protection of copyright and the intellectual property our authors entrust to us. Copyright promotes the progress and integrity of knowledge. Thank you for supporting free speech and the global exchange of ideas by purchasing an authorized edition of this book. If you wish to reproduce or distribute any part of it in any form, please obtain permission. Requests for permission to reproduce material from this work should be sent to [email protected] Published by Princeton University Press 41 William Street, Princeton, New Jersey 08540 99 Banbury Road, Oxford OX2 6JX press.princeton.edu All Rights Reserved Library of Congress Cataloging-in-Publication Data Names: Paul, Gregory S., author. Title: The Princeton fi eld guide to Mesozoic sea reptiles / Gregory S. Paul. Description: Princeton : Princeton University Press, [2022] | Series: Princeton fi eld guides | Includes bibliographical references and index. Identifi ers: LCCN 2022002212 (print) | LCCN 2022002213 (ebook) | ISBN 9780691193809 (hardback) | ISBN 9780691241456 (ebook) Subjects: LCSH: Marine reptiles, Fossil—Mesozonic—Identifi cation. | Paleontology—Mesozonic. Classifi cation: LCC QE861 .P39 2022 (print) | LCC QE861 (ebook) | DDC 567.9/37– dc23/eng20220420 LC record available at https://lccn.loc.gov/2022002212 LC ebook record available at https://lccn.loc.gov/2022002213 British Library Cataloging-in-Publication Data is available Editorial: Robert Kirk and Megan Mendonça Production Editorial: Kathleen Cioffi Jacket Design: Wanda España Production: Steven Sears Publicity: Matthew Taylor and Caitlyn Robson Copyeditor: Laurel Anderton This book has been composed in Goudy Old Style (introduction), ITC Galliard Pro (species section and headings), and Proxima Nova (labels) Printed on acid-free paper. ∞ Typeset and designed by D & N Publishing, Wiltshire, UK Printed in Italy 10 9 8 7 6 5 4 3 2 1 01 MSR – Intro pp001- 65.indd 4 18/05/2022 15:33 CONTENTS Preface 6 Acknowledgments 6 Introduction History of Discovery and Research 9 What Is a Mesozoic Sea Reptile? 10 Dating Sea Reptiles 12 The Evolution of Sea Reptiles and Their Mesozoic World 13 Extinction 23 After the Age of Sea Reptiles 25 Biology 26 General Anatomy 26 Swimming 32 On the Beach 40 The Salt Problem 40 Respiration and Circulation 40 Feeding Apparatus and Digestive Tract 41 Sea Reptiles as Food 42 Senses 43 Vocalization 45 Diseases, Pathologies, and Injuries 45 Behavior 45 Brains, Nerves, and Intelligence 45 Social Activities 45 Reproduction 46 Growth and Aging 47 Energetics 48 Gigantism 52 Mesozoic Marine Expedition 52 If Ancient Sea Reptiles Survived 54 Mesozoic Marine Reptile Conservation, Keeping, and Consumption 55 Where Fossil Sea Reptiles Are Found 55 Using the Group and Species Descriptions 61 Group and Species Accounts Diapsids 67 Neodiapsids 68 Index Mesozoic Sea Reptiles Taxa 203 Formations 207 01 MMR – Intro pp001- 65.indd 5 16/05/2022 14:22 PREFACE If I were, at about age twenty as a budding paleoresearcher and often misleading plethora of old names often based on poor artist, handed a copy of this book by a mysterious time traveler, material and errant assignments of fossil specimens. So the I would have been startled as well as delighted. The pages would name Steneosaurus, which had been applied to a number of sea reveal a world of new seagoing reptiles and ideas that I had only crocs, has now been obsolesced out, replaced by the likes of a hint of, if any such ideas existed at all. My head would spin at Macrospondylus, Yvridiosuchus, Proexochokefalos, Plagiophthalmo- the revelation that some of the sea reptiles that had long been suchus, and Bathysuchus. presumed to have low metabolic rates, like living turtles, lizards, Remaining frustratingly unresolved is the evolutionary origin and crocodilians, show signs of being highly energetic, more of some marine reptile groups. The specific origin of turtles as like seabirds and mammals. Also remarkable is the discovery a whole remains unresolved. Also still mysterious are the deriva- that many of the mosasaurs, which literally are marine lizards, tions of ichthyosaurs, thalattosaurs, and helveticosaurs. were not the tail-sculling crocodilian swimmers once thought Producing this volume has been satisfying in that it has given but rather were configured for stiffer-bodied, higher-speed me reason to illustrate the skeletons of almost all Mesozoic ma- swimming. Or how about Eretmorhipis with its platypus-like rine reptile species for which nearly complete material is avail- head, and Atopodentatus with its hammerhead? Plus there are able. These have been used to construct the most extensive the novel formations, at least to my eyes and ears—Sticky Keep, library of side-view life studies of these sea creatures to date in Loon River, Ziliujing, Hiccles Cove, Xiashaxiamo, Paja, Guan- print. An advantage of producing large sets of rigorous skeletals ling, Tamayama, Snow Hill Island, Katiki, Paso del Sapo, Lopez is that it can reveal information that would not otherwise arise. de Bertodano, Akrabou, Falang, Dukamaje, Arcadia Park, Mu- The overall result is a work that covers what is already well over waqqar Chalk, Hakel, Bet-Meir, Cerro del Pueblo, Romualdo, two centuries of scientific investigation into the groups of tetra- Phu Kradung, Snowshoe, Rosso Ammonitico. The sheer pods that inhabited the seas for up to 186 million years. Enjoy number of new sea reptiles with exotic names—Endennasaurus, the travel back in time. Miodentosaurus, Xinpusaurus, Helveticosaurus, Sinosaurosphargis, Majiashanosaurus, Borealonectes, Stenorhynchosaurus, Sachicasau- rus, Wapuskanectes, Attenborosaurus, Kawanectes, Futabasaurus, Acknowledgments Vegasaurus, Albertonectes, Kaiwhekea, Aristonectes, Morturneria, Thililua, Sclerocormus, Phantomosaurus, Barracudasauroides, Kyhy- tysuka, Qianichthyosaurus, Guanlingsaurus, Keilhauia, Undorosau- Thanks to Kenneth Carpenter, Michela Johnson, Judy Massare, rus, Simbirskiasaurus, Tethysaurus, Russellosaurus, Taniwhasaurus, Hilary Ketchum, Mark Young, Sven Sachs, Asier Larramendi, Kaikaifilu, Tenerasaurus, Aphanizocnemus, Haasiophis, Mexichelys, Marcela Gomez Perez, Leslie Noe, Michael Taylor, Michael Santanachelys, Indosinosuchus, Zoneait, Neptunidraco—would indi- Everhart, Nicholas Gardner, Ben Creisler, Wafa Alhalabi, cate that an explosion in discoveries and research, well beyond Sarah Chapman, Jane Davidson, John Schneiderman, Cornelis anything that had previously occurred, and often based on new Hazevoet, Vanessa Rhue, Mallory Theurer, Patrick Druckemiller, high technologies, marked the end of the twentieth century Stephen Poropat, Serjoscha Evers, David Cerny, and Robert Tel- going well into the twenty-first. leria. I would also like to thank all those who worked on this A major and coincidental aid to the production of this book book for Princeton University Press: Robert Kirk, Megan Men- was that while it was being created, a new wave of research at donça, Kathleen Cioffi, Wanda España, Steven Sears, Matthew long last cleared up much of what had been a perplexing and Taylor, Caitlyn Robson, and Laurel Anderton. 6 01 MMR – Intro pp001- 65.indd 6 16/05/2022 14:22 I N T R O D U C T I O N 01 MMR – Intro pp001- 65.indd 7 16/05/2022 14:22 Peloneustes philarchus 01 MMR – Intro pp001- 65.indd 8 16/05/2022 14:22 HISTORY OF DISCOVERY AND RESEARCH The remains of ancient sea reptiles have been found by humans Mesozoic forms during an age of new and exciting scientific and for millennia and may have helped form the basis for belief in technological progress. mythical beasts, including dragons and sea serpents. In the After the American Civil War, the United States entered the prescientific West the claim in the Genesis creation story that ancient maritime reptile scene with the discovery of a host of the planet and all life were formed just two or three thousand often well-preserved mosasaurs and plesiosaurs from Cretaceous years before the great Egyptian pyramids were built hindered the marine beds in the Great Plains. The first such fossil apparently scientific study of fossils. The peculiar, incomplete remains of sparked the fossil war between Othniel Marsh and an oversensi- plesiosaurs become public knowledge, albeit without any under- tive Edward Cope, when the former pointed out to the latter standing of what they were, well back in the 1600s and continu- that he had made the embarrassingly obvious anatomical error ing into the 1700s, when the also incomplete and fishlike fossils of mistaking the extraordinarily long neck of the famed plesio- of ichthyosaurs, thought to be those of fish, also came to public saur Elasmosaurus for the tail, and the short tail for the neck. attention. In the late 1700s mosasaurs began to show up, and The ensuing rivalry contributed to Cope and Marsh naming in the early 1800s a budding scientific community, increasingly a long list of species, again mostly invalid, although Tylosaurus aware that many fossils represented creatures no longer alive on and Platecarpus are real. Also uncovered were late Mesozoic sea the planet, began to realize that plesiosaurs, ichthyosaurs, and turtles, including the gigantic Archelon. mosasaurs represented exotic extinct groups of aquatic reptiles, From the late 1800s into the early 1900s, Mesozoic sea rep- often quite large. Mosasaurs in particular were recognized—by tile discoveries continued at a good clip, with Samuel Williston the great anatomist Georges Cuvier, who helped establish paleo- being the lead researcher in the United States. And in Australia, zoology—as true lizards specialized for a marine lifestyle, while the Cretaceous plesiosaur then known as Kronosaurus (currently the systematic position of the other groups remained obscure Eiectus) set a new size standard. Popular interest in marine rep- other than that they were some form of reptile, not the fish, am- tiles remained substantial, although they had been somewhat phibians, crocodilians, marine mammals, or preflood humans displaced by the dinosaurs of the land and the pterosaurs of some had been mistaken for. the sky. The world wars and Great Depression put a damper Western Europe, with its increasingly scientific orientation, on the science of sea reptiles, so that they came to be seen, and with plenty of ancient marine Jurassic and Cretaceous sedi- along with dinosaurs and pterosaurs, as evolutionary dead ends ments to produce aquatic fossils, remained the center of the of limited importance—good for getting crowds into museum first generation of sea reptile discoveries and research. Along halls, but not worthy of deep intellectual attention. Even so, with flying pterosaur fossils, which became available in the late skeletons of the whale-sized Triassic ichthyosaur Shonisaurus, first 1700s, the new marine reptiles including Mosasaurus, Plesiosau- recognized in the 1920s, were excavated in Nevada from the rus, and Ichthyosaurus played an important role in overturning 1950s to 1970s. In the 1960s Dale Russell produced a major the faith-based belief in a very young earth in which life had not study on the mosasaurs. undergone major transformations or extinctions over deep time, The great revival and revolution of scientific and popular and they did so before the existence of dinosaurs was realized interest in dinosaurs and pterosaurs—which began in the late in the 1820s. Of particular importance to the first generation 1960s, picked up in the 1970s, and went into the 1980s and of modern maritime reptile finds was the renowned Mary An- beyond—at first passed marine reptiles by, as they continued ning, who collected major Jurassic plesiosaur and ichthyosaur to be seen as conservative reptilians. This began to change in skulls and skeletons on the southwest coast of England from the 1990s and continued into the current century. In British the 1810s to the 1840s. Anning prospected, often under dan- Columbia the remains of an exceptionally big early ichthyosaur gerous winter conditions at the base of unstable coastal cliffs, were excavated. In this century it was realized that the last of the even though the severe societal limitations on the working class mosasaurs had evolved compact bodies and more vertical, sym- as well as her gender hindered the analytical side of her work. metrical tails that indicate they were high-speed swimmers of the But in part because she was a very atypically female fossil col- open ocean. It is also being discovered that—much as dinosaurs lector, she became widely known (and remains so today) for her have proven to have had high, rather than reptilian, metabolic major contributions by both the populace and the otherwise rates—ichthyosaurs, plesiosaurs, and mosasaurs had elevated me- patriarchal scientific establishment of the time. Also entering tabolisms and warm bodies. And turning up in recent years are the scene were major early official paleozoologists, most notably marine reptiles with heads like hammers, and others with heads Richard Owen, who named hundreds of species, albeit inval- like that of the duck-billed platypus. idly based mostly on fragmentary remains. Also revealed were After centuries of research, much of it using the sophisticated other sea reptiles—placodonts, nothosaurs, and crocodile rela- technologies of the last few decades, what is known about the tives such as Teleosaurus. Marine reptiles became highly popular basic paleobiology of Mesozoic marine reptiles is unlikely to un- with a public fascinated by these sometimes big, usually bizarre dergo a dramatic change in the future. Even so, the research and 9 01 MMR – Intro pp001- 65.indd 9 16/05/2022 14:22