“Ever since Nick Hotton’s book from the 1980s SCIENCE · PALEONTOLOGY we have needed an update on the biology of therapsids, and it has been Anusuya Chinsamy- C R ADIATION HISTOLOGY BIOLOGY About 320 million years ago a Turan and her students and associates who H group of reptiles known as the synapsids through their bone histological I N emerged and forever changed Earth’s work have made the S greatest progress in A F O R E R U N N E R S ecological landscapes. This book discusses M the origin and radiation of the synapsids this fi eld.” Y from their sail-backed pelycosaur ancestor Martin Sander, - T to their diverse descendants, the therapsids Steinmann U O F R or mammal-like reptiles, which eventually Institute, A N gave rise to mammals. It further showcases University the remarkable evolutionary history of the of Bonn F synapsids in the Karoo Basin of South Africa M A M M A L S O and the environments that existed at the R E time. By highlighting studies of synapsid R Anusuya Chinsamy-Turan is a U bone microstructure, it offers a unique N perspective of how such studies are utilized paleobiologist and global expert on fossil N EDITED BY ANUSUYA CHINSA MY-TUR AN to reconstruct various aspects of biology, bone microstructure. She is Professor and E R such as growth dynamics, biomechanical Fellow of the University of Cape Town and S function, and the attainment of sexual and is former director of the National History O skeletal maturity. A series of chapters outline Collections, Iziko Museums of Cape Town. F the radiation and phylogenetic relationships She is author of The Microstructure of M of major synapsid lineages and provide Dinosaur Bone: Deciphering Biology through A M direct insight into how bone histological Fine Scale Analysis and Famous Dinosaurs M analyses have led to an appreciation of these of Africa. A enigmatic animals as once-living creatures. L S The penultimate chapter examines the early radiation of mammals from their nonmam- malian cynodont ancestors, and the book concludes by engaging the intriguing ques- tion of when and where endothermy evolved among the therapsids. I N D I A N A Bloomington & Indianapolis iupress.indiana.edu Jacket illustrations: Luis Rey 1-800-842-6796 Life of the Past James O. Farlow, editor ForerunnersMmec.indd 1 9/21/11 10:39 AM Forerunners oF M a MM als Life of the Past James O. Farlow, editor Forerunners oF M a MM als Radiation · Histology · Biology Edited by Anusuya Chinsamy-Turan Indiana University Press Bloomington and Indianapolis This book is a publication of Manufactured in the United States of America Indiana University Press 601 North Morton Street Library of Congress Bloomington, Indiana 47404-3797 USA Cataloging-in-Publication Data iupress.indiana.edu Forerunners of mammals : radiation, histology, biology / edited by Anusuya Telephone orders 800-842-6796 Chinsamy-Turan. Fax orders 812-855-7931 p. cm. — (Life of the past) Includes bibliographical references and © 2012 by Indiana University Press index. ISBN 978-0-253-35697-0 (cloth : alk. All rights reserved paper) 1. Reptiles, Fossil. 2. Mammals—Evolution. No part of this book may be reproduced 3. Bones—Histology. I. Chinsamy-Turan, or utilized in any form or by any means, Anusuya. electronic or mechanical, including QE861.F67 2012 photocopying and recording, or by any 567.9’3—dc23 information storage and retrieval system, 2011016622 without permission in writing from the publisher. The Association of American 1 2 3 4 5 17 16 15 14 13 12 University Presses’ Resolution on Permissions constitutes the only exception to this prohibition. The paper used in this publication meets the minimum requirements of the American National Standard for Information Sciences—Permanence of Paper for Printed Library Materials, ANSI Z39.48-1992. This book is dedicated to the memory of James W. Kitching (1922–2003) Gondwanan Paleontologist An Exceptional Fossil Hunter and an Inspiration to Many Contents C ix Preface xi Acknowledgments xiii List of Contributors 1 The Origin and Radiation of Therapsids 3 Tom S. Kemp Therapsid Biodiversity Patterns and Paleoenvironments 2 of the Karoo Basin, South Africa 31 Roger Smith, Bruce Rubidge, and Merrill van der Walt The Microstructure of Bones and Teeth 3 of Nonmammalian Therapsids 65 Anusuya Chinsamy-Turan The Paleobiology and Bone Microstructure 4 of Pelycosaurian-Grade Synapsids 91 Adam K. Huttenlocker and Elizabeth Rega 5 Dicynodont Growth Dynamics and Lifestyle Adaptations 121 Sanghamitra Ray, Jennifer Botha-Brink, and Anusuya Chinsamy-Turan Biological Inferences of the Cranial Microstructure of 6 the Dicynodonts Oudenodon and Lystrosaurus 149 Sandra C. Jasinoski and Anusuya Chinsamy-Turan Bone and Dental Histology of Late Triassic 7 Dicynodonts from North America 179 Jeremy L. Green Bone Histology of Some Therocephalians and Gorgonopsians, 8 and Evidence of Bone Degradation by Fungi 199 Anusuya Chinsamy-Turan and Sanghamitra Ray The Radiation and Osteohistology of 9 Nonmammaliaform Cynodonts 223 Jennifer Botha-Brink, Fernando Abdala, and Anusuya Chinsamy-Turan 10 The Radiation, Bone Histology, and Biology of Early Mammals 249 Jørn H. Hurum and Anusuya Chinsamy-Turan 11 The Evolution of Mammalian Endothermy 273 John A. Ruben, Willem J. Hillenius, Tom S. Kemp, and Devon E. Quick 289 References 321 Index Preface P This book brings together a group that has over many years researched various aspects of the evolution and paleobiology of the synapsids. Many of us have collaborated in our research endeavors, and all of us have at some stage shared information and had many hearty discussions about the biology of our distant relatives. The book comprises eleven chapters. The first two chapters provide an introduction to the predecessors of mammals and their relatives, and an assessment of the ancient world in which they radiated. The opening chapter sets the scene, providing a guide of “who” the synapsids were and how they are related to one another. In this chapter, Tom Kemp provides an up to date assessment of the radiation of the synapsids from their earli- est pelycosaur members, to the diverse nonmammalian therapsids, and later to the increasingly more mammal-like cynodonts. All this is done from a global perspective. The second chapter of this book deals more specifically with the Ka- roo Basin of South Africa and documents an unparalleled track record of the evolution and radiation of the therapsids. Here, Karoo paleontologists, Roger Smith and Bruce Rubidge, together with a recent PhD graduate Merrill van der Walt, provide a unique perspective of therapsid biodiver- sity and paleoenvironmental analysis of the Karoo Basin of South Africa. For the first time, faunal turnover in the Karoo Basin is provided through a lens of absolute numbers of genera and have permitted detailed trophic level analyses for each of the biozones. The third chapter by yours truly sets the scene for the bone microstruc- ture chapters that follow. The first part of this chapter takes the form of an atlas of bone microstructure that will enable a novice to identify particular types of bone tissues in synapsids. The second part examines the biologi- cal implications of particular types of bone microstructures and how these can be utilized to deduce various aspects of the biology of extinct animals. The next seven chapters focus on particular synapsid lineages. Each of these chapters is structured to provide a phylogenetic and paleobio- logical context before delving into the bone microstructure of that par- ticular group. The first in this series of chapters (chapter 4) predictably deals with the earliest members of the Synapsida, the pelycosaurs. Here, Adam Huttenlocker and Elizabeth Rega present an overview of the bone microstructure of pelycosaurian-grade synapsids from both “normal” and pathological skeletal elements. The fifth chapter is by Sanghamitra Ray, my former postdoctoral fellow, Jennifer Botha-Brink, my former PhD student, and me. Here we ix
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