Theagarten Lingham-Soliar The Vertebrate Integument Volume 1 Origin and Evolution The Vertebrate Integument Volume 1 The arid and barren landscape walled by plateaus, represents the hostile environment of the Cape Karoo in South Africa (a region pioneered by the author’s maternal family). The rocks of the Karoo System were deposited between the Carboniferous (360–286 million years ago) and Early Jurassic (208–187 million years ago) and are known for some of the most important finds of mammal-like reptiles in the world (Chap. 8). Photo, B Lingham (circa 1922, family archive) Theagarten Lingham-Soliar The Vertebrate Integument Volume 1 Origin and Evolution 123 Theagarten Lingham-Soliar Life Sciences University of KwaZulu-Natal Durban South Africa Present address Environmental Sciences Nelson Mandela Metropolitan University Port Elizabeth South Africa ISBN 978-3-642-53747-9 ISBN 978-3-642-53748-6 (eBook) DOI 10.1007/978-3-642-53748-6 Springer Heidelberg New York Dordrecht London Library of Congress Control Number: 2013957128 Ó Springer-Verlag Berlin Heidelberg 2014 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. Exempted from this legal reservation are brief excerpts in connection with reviews or scholarly analysis or material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Duplication of this publication or parts thereof is permitted only under the provisions of the Copyright Law of the Publisher’s location, in its current version, and permission for use must always be obtained from Springer. Permissions for use may be obtained through RightsLink at the Copyright Clearance Center. Violations are liable to prosecution under the respective Copyright Law. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. While the advice and information in this book are believed to be true and accurate at the date of publication, neither the authors nor the editors nor the publisher can accept any legal responsibility for any errors or omissions that may be made. The publisher makes no warranty, express or implied, with respect to the material contained herein. Printed on acid-free paper Springer is part of Springer Science+Business Media (www.springer.com) Dedicated to my mother LL-S, Who gave us dignity amidst the undignified world of pre-1994 South Africa Preface The need for the present book became increasingly apparent as I worked on the integument and its structures over the past 14–15 years, first in fossil groups and later in extant vertebrates. Even among living vertebrates, in particular marine groups, it was only in the last quarter of the twentieth century that we were beginning to understand the structure of the dermis and its enormous biome- chanical contribution to locomotion. Coming right up to 2013 my colleagues and I were able to demonstrate how little the biomechanical microstructure of the feather was known given that such knowledge would have important implications for bird flight. Yet we need to go back over 450 million years to the very beginning of vertebrate evolution to put our understanding of the vertebrate integument today in proper context. I was fortunate that my undergraduate lecturer and later Ph.D. supervisor, Beverly Halstead, at the University of Reading in the UK, was one of the key workers involved in research on vertebrate origins and that his seminal contributions on the extinct jawless vertebrates, on the origin of bone and on the origin of teeth formed a major part of his zoology lectures. This background characterises the direction and function of the book, which is aimed at both undergraduate and postgraduate students. The topic of the book at times may appear complex but the profusely illustrated text with diagrams, photos and some of my own artwork may help make the book easy to use and understand. While many of the illustrations are original I owe a debt to many authors cited whose illustrations have been used to a greater or lesser extent, unmodified or modified (if there are errors, they are mine). Fossilised integument and integumental structures although scarce have been preserved in vertebrate fossils, perhaps nowhere more so than in the famous Liaoning region in China. Although much of the integumental material on dino- saurs from China have, in recent years, been concerned with possible feather origins or feather-like structures, I hope the book will encourage investigations on a broader level because there is, I am sure, much material that may be neglected with respect to knowledge on the integument in general (i.e. not connected with feather origins). For instance, I have demonstrated this in the dermis of the dinosaur Psittacosaurus from Liaoning and with a colleague, Prof. Gerhard Plowdowski, on colour in the epidermis of another specimen, also from Liaoning. Feather origins are important, hence there is no suggestion that the search needs to be downplayed. vii viii Preface It seems safe to say that despite the importance of the integument in the lives of vertebrates and its changing morphological and functional role over time there is no other book that deals with the topic specifically, in particular its mechano- structural properties. Volume 2 will discuss the biomechanics of the integument in much greater detail. Acknowledgments There are many individuals who have made my science possible, yet none more than my mother. My mother died unexpectedly before I received my Ph.D. She had an exciting early life in the Karoo—family racehorses, driving her parents Rolls Royce aged 14 and achieving recognition as a talented classical pianist while still in her teens. It all ended with the ‘legalized’ expropriation of our family’s assets and wealth during the apartheid years in South Africa, when I was just a boy. Despite the unimaginable hardships of those years, she single-handedly filled our lives with culture, kindness and hope. My brother Sagaren Soliar is 8 years older than I but thankfully he failed to recognise that when I was growing up. An avid reader, he would shove books at me to read. At age about 8, most memorably was Mika Waltari’s Sinuhe the Egyptian, regarded by authorities today as among the greatest historical novels written. It started my passion for Egyptology to the extent that when a few years later my brother raved about a book on hominid origins by the Leakey’s, I scathingly said, verbatim, that I would rather search for the tombs of pharaohs, even with the prospect of failure, than sift sand in a barren landscape only to find a few dried-up, old bones after 30 years—famous last words. His passion for knowledge and vibrant exchange of ideas had the most profound effect on my development and perceptions. My early career as an industrial research chemist in the UK, I owe to one man, Les Leedham, head of the research laboratory. He fostered my ability and arranged day-release to enable me to continue my part-time university studies (in chemistry). He was a giant of a man both in stature (over 6 feet 6 inches tall) and in human decency and confirmed why as an exile from South Africa I was proud to have chosen England as my new home. Several years later I met Beverly Halstead, who lectured the first year zoology undergraduate course at Reading University. I was not the keenest of students, preferring the fine pubs in Berkshire and interesting company to diligent course studies. However, Beverly Halstead’s passion for paleontology got through and in spite of myself I gained first class grades in it. Kindly, he recommended my study for a Ph.D. to the head of department with the words that he would rather chose me than someone with a double-first from Cambridge. Beverly Halstead died in a traffic accident before I could complete my Ph.D. ix x Acknowledgments During my Ph.D. I met Richard Estes (San Diego State University), one of the top herpetologists in the world who was on a year’s sabbatical at the Natural History Museum, London. We liked each other instantly and he had a major effect on my research. Before he flew back to the USA we agreed that he would for- malize himself as my second supervisor. I would never see him again. Shortly after returning home, Richard was diagnosed with inoperable intestinal cancer and he died within the year. Tom Kemp at Oxford University took over as bona fide supervisor. For his help I am grateful but also for his bravery, given that I had already got through two supervisors within 4 months of each other. Wolf-Ernst Reif was my host at Tubingen University on two successive Royal Society Post-doctoral Fellowships. He had a significant effect on my research on marine vertebrates, in particular on sharks, in which he was a world expert. I benefitted too from his extensive knowledge of philosophy. Alexi Yu Rozanov, a principled and generous man, was my host over 5 years at the Paleontological Institute of the Russian Academy of Sciences. He was extre- mely helpful to my research and instrumental in my appointment during this time as editor of Paleontological Journal.