Snakes of Central and Western Africa i-xii.indd 1 2019-03-20 7:25 AM i-xii.indd 2 2019-03-20 7:25 AM Snakes of Central and Western Africa Jean- Philippe Chippaux and Kate Jackson Johns Hopkins University Press • Baltimore i-xii.indd 3 2019-03-20 7:25 AM © 2019 Johns Hopkins University Press All rights reserved. Published 2019 Printed in Canada on acid- free paper 9 8 7 6 5 4 3 2 1 Johns Hopkins University Press 2715 North Charles Street Baltimore, Maryland 21218-4363 www.press.jhu.edu Library of Congress Cataloging- in- Publication Data Names: Chippaux, Jean- Philippe, author. | Jackson, Kate, 1972– author. Title: Snakes of Central and Western Africa / Jean- Philippe Chippaux and Kate Jackson. Description: Baltimore : Johns Hopkins University Press, 2019. | Includes bibliographical references and index. Identifiers: LCCN 2018023756 | ISBN 9781421427195 (hardcover : alk. paper) | ISBN 1421427192 (hardcover : alk. paper) | ISBN 9781421427201 (electronic) | ISBN 1421427206 (electronic) Subjects: LCSH: Snakes—Africa, Central—Identification. | Snakes—Africa, West—Identification. Classification: LCC QL666.O6 C5352 2019 | DDC 597.960967—dc23 LC record available at https://lccn.loc.gov/2018023756 A catalog record for this book is available from the British Library. Frontispiece: Dipsadoboa viridis defensive posture, Republic of Congo. By K. Jackson Special discounts are available for bulk purchases of this book. For more information, please contact Special Sales at 410-516-6936 or [email protected]. Johns Hopkins University Press uses environmentally friendly book materials, including recycled text paper that is composed of at least 30 percent post- consumer waste, whenever possible. i-xii.indd 4 2019-03-20 7:25 AM For Monique Bourgeois, in honor of her paradigm- shifting recognition of the true phylogenetic affiliations of Atractaspis, an independently derived lineage of front- fanged venomous snakes previously assumed to be vipers. Bourgeois made this discovery using graph paper and a dissecting microscope, at the University of Lubumbashi in the early 1960s, decades before technological advances made it possible for molecular systematics to prove her right. i-xii.indd 5 2019-03-20 7:25 AM This page intentionally left blank i-xii.indd 6 2019-03-20 7:25 AM Contents Acknowledgments ix 1 • Identification of African Snakes 1 2 • Evolution of African Snakes 28 3 • Biogeography of African Snakes 40 4 • Snakebite in Sub- Saharan Africa 49 5 • Families Typhlopidae and Leptotyphlopidae 53 6 • Families Boidae and Pythonidae 60 7 • Family Viperidae 72 8 • Family Elapidae 104 9 • Family Lamprophiidae 136 Subfamilies Atractaspidinae and Aparallactinae 10 • Family Lamprophiidae 189 Subfamilies Lamprophiinae, Pseudoxyrhophiinae, and Pseudaspidinae 11 • Family Lamprophiidae 241 Subfamilies Psammophiinae and Prosymninae 12 • Family Colubridae 288 Subfamily Natricinae 13 • Family Colubridae 302 Subfamilies Colubrinae and Grayiinae Bibliography 385 Index 411 i-xii.indd 7 2019-03-20 7:25 AM This page intentionally left blank i-xii.indd 6 2019-03-20 7:25 AM Acknowledgments We acknowledge the following people and institutions for their help in making this book possible. We thank the National Science Foundation (grant 1145437 to KJ) and Whitman College for funding. We thank the museums whose collections we referred to in the course of research for this book, and the curators and collections managers who facilitat- ed our efforts, particularly: Garin Cael and Danny Meirte at the Royal Museum of Central Africa, Tervuren; Jeremy Jacobs, Roy McDiarmid, James Poindexter, Rob Wilson, Addison Wynn, and George Zug at the Smithsonian Institution, Washington, DC; Alan Resatar at the Field Museum of Natural History, Chicago; Patrick Campbell and Colin McCarthy at the Natural History Museum, London; David Kizirian and Chris Raxworthy at the American Museum of Natural History, New York; Andreas Schlüter and Günter Stephan at the Staatliches Muse- um für Naturkunde, Stuttgart; Ivan Ineich, Laure Paul, and Nicolas Vidal at the Museum National d’Histoire Naturelle, Paris. We thank the students from Whitman College (United States) and from Uni- versité Marien Ngouabi (Republic of Congo) who worked with KJ on various aspects of this project: Jordan Benjamin, Sylvestre Boudzou- mou, Nat Clarke, Evan Conner, Andrew Hill, Eric Hsu, Willie Kunkel, Lise- Bethy Mavoungou, Ange Mboungou- Louiki, Kevin Moore, Katie Moyer, Anna Ripley, Claire Snyder, Ange- Ghislain Zassi- Boulou. We thank colleagues and coworkers who lent their expertise in geogra- phy and geographic information system (GIS) technology to aspects of this project: Austun Ables, Nick Bader, Bob Carson, Amy Molitor. We thank friends and colleagues who gave feedback and suggestions as we developed this book, and we thank fellow herpetologists and pho- tographers of African snakes who allowed us to use their photographs in our book. We thank Tuhin Giri for applying his artistic talent and eye for scientific accuracy to the head scale drawings used throughout this book. We thank the peer reviewer for constructive comments and suggestions provided. Finally, we thank our editor, Tiffany Gasbarrini, and the staff of Johns Hopkins University Press for bringing it all together. Jean- Philippe Chippaux and Kate Jackson ix i-xii.indd 9 2019-03-20 7:25 AM