Developments in Primatology: Progress and Prospects Series Editor: Louise Barrett Tracy L. Kivell Pierre Lemelin Brian G. Richmond Daniel Schmitt Editors The Evolution of the Primate Hand Anatomical, Developmental, Functional, and Paleontological Evidence Developments in Primatology: Progress and Prospects Series Editor Louise Barrett Lethbridge , Alberta , Canada More information about this series at http://www.springer.com/series/5852 Tracy L. Kivell • Pierre Lemelin Brian G. Richmond • Daniel Schmitt Editors The Evolution of the Primate Hand Anatomical, Developmental, Functional, and Paleontological Evidence Editors Tracy L. Kivell Pierre Lemelin Animal Postcranial Evolution (APE) Lab Division of Anatomy Skeletal Biology Research Centre Department of Surgery School of Anthropology and Conservation Faculty of Medicine and Dentistry, University of Kent University of Alberta Canterbury, UK Edmonton , AB , Canada Department of Human Evolution Daniel Schmitt Max Planck Institute for Evolutionary Department of Evolutionary Anthropology Anthropology Duke University Leipzig , Germany Durham , NC , USA Brian G. Richmond Division of Anthropology American Museum of Natural History New York , NY , USA Department of Human Evolution Max Planck Institute for Evolutionary Anthropology Leipzig, Germany ISSN 1574-3489 ISSN 1574-3497 (electronic) Developments in Primatology: Progress and Prospects ISBN 978-1-4939-3644-1 ISBN 978-1-4939-3646-5 (eBook) DOI 10.1007/978-1-4939-3646-5 Library of Congress Control Number: 2016935857 © Springer Science+Business Media New York 2016 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifi cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfi lms 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. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specifi c statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. Printed on acid-free paper This Springer imprint is published by Springer Nature The registered company is Springer Science+Business Media LLC New York Foreword Rarely are we privileged to witness the appearance of a book that strikes out in a completely new, groundbreaking direction and will accelerate a major fi eld of research. The new direction of this book on the evolution of the primate hand is toward a comprehensive, highly informed, critical review of the subject in chapters, not by a single author but by a team of experts actively involved in the varied fi elds that contribute to the understanding of the subject. The team approach ensures uni- form high quality of the review together with a shared underlying theme (the vari- ability of primate hands upon a shared primitive pattern). Acceleration of research on the evolution of the hand will be the inevitable result of the team’s having laid a deep, broad foundation of knowledge and ideas for the production of new investigations. This highly readable collection of chapters will be a welcomed resource to all who are interested in the human hand, its functions, and its origins. There is a fasci- nation with hands and their expression of human behavior, manifest in representa- tions ranging from the walls of prehistoric caves to the Social Programs Bas-relief created by Robert Graham at the Roosevelt Memorial on the Washington D.C. mall. Hand surgeons marvel to me at the dexterity and adaptability of the human hand to its varied roles, and teachers lament the trend away from “hands-on” activities that enhance learning in K-12 science courses and even in medical anatomy laborato- ries. It seems to them almost as if our hand was freed from locomotion only to become captive to computer keyboards! All these readers will fi nd that most of our remarkable manipulative capabilities originated early in our primate ancestry and may be understood in the context of nonhuman primate locomotor and manipulative behavior, including the constant interaction of touch and proprioceptive cues in learning about the physical and social environment. For the fi rst time we fi nd in a single book descriptions of all currently known human and nonhuman primate fossil hand bones, together with detailed descriptive and quantitative data on living primate musculoskeletal hand anatomy, develop- ment, and uses that inform functional and phylogenetic analyses of the fossils. In addition, we are introduced to the most recent developments in approaches to fi ne- tuning these analyses. Three chapters review experiments involving new techniques v vi Foreword for imaging, motion capture, and hand pressure recording, along with sophisticated modeling of joint movements and stresses in locomotor and manipulative behavior. Readers learn about the wide variety and impressive fl exibility of primate hand postures during locomotion as well as manipulation and are encouraged to expand studies to include active haptic sensing of shapes, weights, and textures of objects with which primates interact in their environment. A chapter on hand development brings the reader up to date on the current understanding of genetic and develop- mental factors in phenotypic variation and addresses potential for and constraints on phylogenetic change in the hands of primates. Especially welcome in a new book on primate hand evolution are chapters on the comparative morphology of hand integ- ument and on neural control of the hand. Most impressive is the tremendous effort made by the authors not only to review the literature critically in each area but also to summarize extensive, detailed infor- mation in tables and to provide drawings and photographs that are informative and relevant to the text. Careful attention is given to the defi nitions and uses of terms for anatomical features, grips, thumb and fi nger movements, and taxonomic categories, which should at last reduce confusion in the literature. Here is a springboard for new research that will enable us to communicate knowledgeably and effectively about how our future fi ndings relate to the fi ndings of our current and former colleagues. In their suggestions for future directions in research, the authors echo persistent calls in the literature for more fossils, especially for associated elements of the hand and evidence from the early evolutionary stages of the genus Homo . They strongly emphasize in addition the need for comparative studies and functional analyses of morphological variability within and among a much larger range of primate species. However, their book also reveals the large store of data already available for func- tional and phylogenetic analysis of living and fossil primate hands, and the chal- lenge now will be to keep the book up to date as the future research they propose comes to fruition. It is a great personal pleasure to introduce the book, which will be an invaluable resource for all whose work in evolutionary biology and human health care focuses on the fascinating diversity of primate hands. The editors deserve con- gratulations for their powerful concept and for their monumental achievement. Mary W. Marzke School of Human Evolution and Social Change Arizona State University Tempe, AZ, USA Contents 1 Introduction ............................................................................................. 1 Tracy L. Kivell , Pierre Lemelin , Brian G. Richmond , and Daniel Schmitt 2 On Primitiveness, Prehensility, and Opposability of the Primate Hand: The Contributions of Frederic Wood Jones and John Russell Napier .............................................................. 5 Pierre Lemelin and Daniel Schmitt Part I Anatomical and Developmental Evidence 3 The Primate Wrist .................................................................................. 17 Tracy L. Kivell 4 Morphological Diversity in the Digital Rays of Primate Hands ......... 55 Biren A. Patel and Stephanie A. Maiolino 5 The Role of Genes and Development in the Evolution of the Primate Hand ................................................................................ 101 Campbell Rolian 6 Organization and Evolution of the Neural Control of the Hand in Primates: Motor Systems, Sensory Feedback, and Laterality .......................................................................................... 131 Andrey Verendeev , Chet C. Sherwood , and William D. Hopkins 7 Anatomy, Function, and Evolution of the Primate Hand Musculature .................................................................................. 155 Pierre Lemelin and Rui Diogo 8 Comparative and Functional Morphology of the Primate Hand Integument .................................................................................... 195 Stephanie A. Maiolino , Amanda K. Kingston , and Pierre Lemelin vii viii Contents Part II Biomechanical, Experimental and Behavioral Evidence