LAGOMORPHS 1709048_int_cc2015.indd 1 15/9/2017 15:59 1709048_int_cc2015.indd 2 15/9/2017 15:59 Lagomorphs Pikas, Rabbits, and Hares of the World edited by Andrew T. Smith Charlotte H. Johnston Paulo C. Alves Klaus Hackländer JOHNS HOPKINS UNIVERSITY PRESS | baltimore 1709048_int_cc2015.indd 3 15/9/2017 15:59 © 2018 Johns Hopkins University Press All rights reserved. Published 2018 Printed in China 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: Smith, Andrew T., 1946–, editor. Title: Lagomorphs : pikas, rabbits, and hares of the world / edited by Andrew T. Smith, Charlotte H. Johnston, Paulo C. Alves, Klaus Hackländer. Description: Baltimore : Johns Hopkins University Press, 2018. | Includes bibliographical references and index. Identifiers: LCCN 2017004268| ISBN 9781421423401 (hardcover) | ISBN 1421423405 (hardcover) | ISBN 9781421423418 (electronic) | ISBN 1421423413 (electronic) Subjects: LCSH: Lagomorpha. | BISAC: SCIENCE / Life Sciences / Biology / General. | SCIENCE / Life Sciences / Zoology / Mammals. | SCIENCE / Reference. Classification: LCC QL737.L3 L35 2018 | DDC 599.32—dc23 LC record available at https://lccn.loc.gov/2017004268 A catalog record for this book is available from the British Library. Frontispiece, top to bottom: courtesy Behzad Farahanchi, courtesy David E. Brown, and © Alessandro Calabrese. Special discounts are available for bulk purchases of this book. For more information, please contact Special Sales at 410-516-6936 or specialsales @press .jhu .edu. 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. 1709048_int_cc2015.indd 4 15/9/2017 15:59 CONTENTS Preface vii Ochotona macrotis, Large- eared Pika 58 Contributors ix Ochotona mantchurica, Manchurian Pika 60 Ochotona nubrica, Nubra Pika 61 1 Introduction 1 Ochotona opaca, Kazakh Pika 62 2 Evolution of Lagomorphs 4 Ochotona pallasii, Pallas’s Pika 65 3 Systematics of Lagomorphs 9 Ochotona princeps, American Pika 67 4 Introduced Lagomorphs 13 Ochotona pusilla, Steppe Pika 72 5 Diseases of Lagomorphs 18 Ochotona roylei, Royle’s Pika 75 6 Conservation of Lagomorphs 22 Ochotona rufescens, Afghan Pika 77 Ochotona rutila, Turkestan Red Pika 79 SPECIES ACCOUNTS Ochotona syrinx, Qinling Pika 82 Order Lagomorpha 29 Ochotona thibetana, Moupin Pika 83 Ochotona thomasi, Thomas’s Pika 84 Family Ochotonidae 31 Ochotona turuchanensis, Turuchan Pika 85 Ochotona alpina, Alpine Pika 31 Family Leporidae 87 Ochotona argentata, Helan Shan Pika 33 The Rabbits 87 Ochotona cansus, Gansu Pika 34 Brachylagus idahoensis, Pygmy Rabbit 87 Ochotona collaris, Collared Pika 36 Bunolagus monticularis, Riverine Rabbit 90 Ochotona coreana, Korean Pika 39 Caprolagus hispidus, Hispid Hare 93 Ochotona curzoniae, Plateau Pika 40 Nesolagus netscheri, Sumatran Striped Rabbit 95 Ochotona dauurica, Daurian Pika 43 Nesolagus timminsi, Annamite Striped Rabbit 97 Ochotona erythrotis, Chinese Red Pika 46 Oryctolagus cuniculus, European Rabbit 99 Ochotona forresti, Forrest’s Pika 47 Pentalagus furnessi, Amami Rabbit 104 Ochotona gloveri, Glover’s Pika 48 Poelagus marjorita, Bunyoro Rabbit 107 Ochotona hoffmanni, Hoffmann’s Pika 49 Pronolagus crassicaudatus, Natal Red Rock Hare 108 Ochotona hyperborea, Northern Pika 51 Pronolagus randensis, Jameson’s Red Rock Hare 110 Ochotona iliensis, Ili Pika 53 Pronolagus rupestris, Smith’s Red Rock Hare 111 Ochotona koslowi, Koslov’s Pika 55 Pronolagus saundersiae, Hewitt’s Red Rock Hare 113 Ochotona ladacensis, Ladak Pika 57 Romerolagus diazi, Volcano Rabbit 114 1709048_int_cc2015.indd 5 15/9/2017 15:59 vi Contents Sylvilagus 117 Lepus capensis, Cape Hare 176 Sylvilagus aquaticus, Swamp Rabbit 117 Lepus castroviejoi, Broom Hare 179 Sylvilagus audubonii, Desert Cottontail 120 Lepus comus, Yunnan Hare 181 Sylvilagus bachmani, Brush Rabbit 122 Lepus coreanus, Korean Hare 182 Sylvilagus brasiliensis sensu stricto, Tapetí, Andean Lepus corsicanus, Corsican Hare 184 Cottontail, Rio de Janeiro Dwarf Cottontail 125 Lepus europaeus, European Hare 187 Sylvilagus cognatus, Manzano Mountain Lepus fagani, Ethiopian Hare 190 Cottontail 130 Lepus flavigularis, Tehuantepec Jackrabbit 191 Sylvilagus cunicularius, Mexican Cottontail 131 Lepus granatensis, Iberian Hare 193 Sylvilagus dicei, Dice’s Cottontail 135 Lepus habessinicus, Abyssinian Hare 195 Sylvilagus floridanus, Eastern Cottontail 137 Lepus hainanus, Hainan Hare 196 Sylvilagus gabbi, Gabb’s Cottontail 140 Lepus insularis, Black Jackrabbit 198 Sylvilagus graysoni, Tres Marías Cottontail 142 Lepus mandshuricus, Manchurian Hare 200 Sylvilagus insonus, Omiltemi Rabbit 144 Lepus nigricollis, Indian Hare 201 Sylvilagus mansuetus, San José Brush Rabbit 145 Lepus oiostolus, Woolly Hare 202 Sylvilagus nuttallii, Mountain Cottontail 147 Lepus othus, Alaskan Hare 204 Sylvilagus obscurus, Appalachian Cottontail 149 Lepus peguensis, Burmese Hare 205 Sylvilagus palustris, Marsh Rabbit 152 Lepus saxatilis, Cape Scrub Hare 206 Sylvilagus robustus, Davis Mountains Cottontail 154 Lepus sinensis, Chinese Hare 208 Sylvilagus transitionalis, New England Cottontail 156 Lepus starcki, Ethiopian Highland Hare 209 Sylvilagus varynaensis, Venezuelan Lowland Lepus tibetanus, Desert Hare 211 Rabbit 157 Lepus timidus, Mountain Hare 212 Lepus 159 Lepus tolai, Tolai Hare 216 Lepus alleni, Antelope Jackrabbit 159 Lepus townsendii, White- tailed Jackrabbit 218 Lepus americanus, Snowshoe Hare 163 Lepus victoriae, African Savanna Hare 220 Lepus arcticus, Arctic Hare 165 Lepus yarkandensis, Yarkand Hare 222 Lepus brachyurus, Japanese Hare 168 Lepus californicus, Black- tailed Jackrabbit 170 References 225 Lepus callotis, White- sided Jackrabbit 173 Index 259 1709048_int_cc2015.indd 6 15/9/2017 15:59 PREFACE he IUCN Species Survival Commission Lagomorph of topics of broad interest across all lagomorph species: Specialist Group (LSG) and the World Lagomorph evolution, systematics, lagomorph diseases, introduced Society (WLS) are pleased to present this compre- lagomorphs, and conservation and management. Despite Thensive compendium of all the lagomorphs in the several ongoing controversies in lagomorph taxonomy, world. This work is designed to expand coverage of the we have maintained a conservative systematic approach. world’s lagomorphs and update the 1990 LSG Lagomorph Nevertheless, we highlight relevant taxonomic issues that Action Plan (Rabbits, Hares and Pikas: Status Survey and require attention. Conservation Action Plan, compiled and edited by Joseph A. This work has been a team effort, with 82 specialists Chapman and John E. C. Flux). The Action Plan has contributing to species accounts. We especially thank served as the most thorough single source of informa- Rachel Fadlovich for her work on the references, and Aryn tion on lagomorphs for biologists, but it was never widely Musgrave for constructing all the species range maps. ATS available to the public and it has become outdated. In this thanks Harriet Smith for her insightful editorial work on book we present updated range maps of all lagomorph his chapters. We are grateful to all the photographers who species, high- quality images of most species, as well as provided their work for free. We appreciate the meticulous current information on identification, systematics, ecol- copy editing by Maria E. denBoer. Finally, the collabora- ogy, behavior, reproduction, genetics, physiology, and con- tion with our editorial team from Johns Hopkins Univer- servation and management of the pikas, rabbits, and hares sity Press, Vincent Burke, Tiffany Gasbarrini, Debby Bors, of the world. The book also summarizes key components and Meagan M. Szekely, is highly appreciated. 1709048_int_cc2015.indd 7 15/9/2017 15:59 This page intentionally left blank CONTRIBUTORS Pelayo Acevedo Fernando Ballesteros Instituto de Investigación en Recursos Cinegéticos Sistemas Naturales Ciudad Real, Spain c/ Santa Susana 15 3° C Oviedo, Spain Maria Altemus School of Natural Resources and the Environment Ron Barry University of Arizona P. O. Box 471 Tucson, Arizona, United States Lewiston, Maine, United States Sergio Ticul Álvarez- Castañeda Amando Bautista Centro de Investigaciones Biológicas del Noroeste Centro Tlaxcala de Biología de la Conducta La Paz, Baja California Sur, Mexico Universidad Autónoma de Tlaxcala Tlaxcala, Mexico Paulo C. Alves Faculdade de Ciências & CIBIO Penny A. Becker Universidade do Porto, Campus de Vairão Washington Department of Fish and Wildlife Vairão, Vila do Conde, Portugal Olympia, Washington, United States Nguyen The Truong An Erik A. Beever Leibniz Institute for Zoo and Wildlife Research U.S. Geological Survey Berlin, Germany Northern Rocky Mountain Science Center Bozeman, Montana, United States Anders Angerbjörn Department of Zoology Hichem Ben Slimen Stockholm University High Institute of Biotechnology of Béja Stockholm, Sweden Béja, Tunisia Asma Awadi Joel Berger Faculty of Sciences of Tunis Department of Fish, Wildlife and Conservation Biology University El Manar Colorado State University Tunis, Tunisia Fort Collins, Colorado, United States 1709048_int_cc2015.indd 9 15/9/2017 15:59 x Contributors Leah K. Berkman Mayra de la Paz Cooperative Wildlife Research Laboratory Centro de Investigaciones Biológicas del Noroeste Southern Illinois University La Paz, Baja California Sur, Mexico Carbondale,