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Home , Abyssinian hare, African savanna hare, Alaskan hare, Amami rabbit, American pika, Appalachian cottontail

LAGOMORPHS

1709048_int_cc2015.indd 1 15/9/2017 15:59 1709048_int_cc2015.indd 2 15/9/2017 15:59 Lagomorphs , , and 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 on acid-­free paper 9 8 7 6 5 4 3 2 1

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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: . | BISAC: SCIENCE / Life Sciences / Biology / General. | SCIENCE / Life Sciences / Zoology / . | 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.

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1709048_int_cc2015.indd 4 15/9/2017 15:59 CONTENTS

Preface vii Ochotona macrotis, Large-­eared 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, 67 4 Introduced Lagomorphs 13 Ochotona pusilla, 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 ACCOUNTS Ochotona syrinx, Qinling Pika 82 Lagomorpha 29 Ochotona thibetana, Moupin Pika 83 Ochotona thomasi, Thomas’s Pika 84 Ochotonidae 31 Ochotona turuchanensis, Turuchan Pika 85 Ochotona alpina, Alpine Pika 31 Family 87 Ochotona argentata, Helan Shan Pika 33 The Rabbits 87 Ochotona cansus, Gansu Pika 34 Brachylagus idahoensis, Pygmy 87 Ochotona collaris, 36 Bunolagus monticularis, 90 Ochotona coreana, Korean Pika 39 Caprolagus hispidus, Hispid 93 Ochotona curzoniae, Plateau Pika 40 Nesolagus netscheri, Sumatran Striped Rabbit 95 Ochotona dauurica, 43 Nesolagus timminsi, Annamite Striped Rabbit 97 Ochotona erythrotis, Chinese Red Pika 46 Oryctolagus cuniculus, 99 Ochotona forresti, Forrest’s Pika 47 Pentalagus furnessi, 104 Ochotona gloveri, Glover’s Pika 48 Poelagus marjorita, 107 Ochotona hoffmanni, Hoffmann’s Pika 49 Pronolagus crassicaudatus, Natal 108 Ochotona hyperborea, 51 Pronolagus randensis, Jameson’s Red Rock Hare 110 Ochotona iliensis, 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, 114

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Sylvilagus 117 Lepus capensis, 176 Sylvilagus aquaticus, 117 Lepus castroviejoi, 179 Sylvilagus audubonii, 120 Lepus comus, Yunnan Hare 181 Sylvilagus bachmani, 122 Lepus coreanus, Korean Hare 182 Sylvilagus brasiliensis sensu stricto, Tapetí, Andean Lepus corsicanus, 184 Cottontail, Rio de Janeiro Dwarf Cottontail 125 Lepus europaeus, 187 Sylvilagus cognatus, Manzano Mountain Lepus fagani, Ethiopian Hare 190 Cottontail 130 Lepus flavigularis, 191 Sylvilagus cunicularius, 131 Lepus granatensis, Iberian Hare 193 Sylvilagus dicei, Dice’s Cottontail 135 Lepus habessinicus, 195 Sylvilagus floridanus, 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, 201 Sylvilagus mansuetus, San José Brush Rabbit 145 Lepus oiostolus, 202 Sylvilagus nuttallii, 147 Lepus othus, 204 Sylvilagus obscurus, 149 Lepus peguensis, 205 Sylvilagus palustris, 152 Lepus saxatilis, Cape Scrub Hare 206 Sylvilagus robustus, Davis Mountains Cottontail 154 Lepus sinensis, 208 Sylvilagus transitionalis, Cottontail 156 Lepus starcki, 209 Sylvilagus varynaensis, Venezuelan Lowland Lepus tibetanus, 211 Rabbit 157 Lepus timidus, 212 Lepus 159 Lepus tolai, 216 Lepus alleni, 159 Lepus townsendii, White-­tailed Jackrabbit 218 Lepus americanus, 163 Lepus victoriae, 220 Lepus arcticus, 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

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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 , 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.

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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, Lewiston, , United States

Sergio Ticul Álvarez-­Castañeda Amando Bautista Centro de Investigaciones Biológicas del Noroeste Centro Tlaxcala de Biología de la Conducta La Paz, Baja Sur, Mexico Universidad Autónoma de Tlaxcala Tlaxcala, Mexico Paulo C. Alves Faculdade de Ciências & CIBIO Penny A. Becker Universidade do Porto, Campus de Vairão 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, Northern Rocky Mountain Science Center Bozeman, , United States Anders Angerbjörn Department of Zoology Hichem Ben Slimen Stockholm University High Institute of Biotechnology of Béja Stockholm, Sweden Béja,

Asma Awadi Joel Berger Faculty of Sciences of Tunis Department of Fish, Wildlife and Conservation Biology University El Manar State University Tunis, Tunisia Fort Collins, Colorado, United States

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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, Illinois, United States Miguel Delibes-­Mateos Sabuj Bhattacharyya CIBIO, Universidade do Porto Molecular Ecology Laboratory Campus Agrario de Vairão Centre for Ecological Sciences Vairão, Vila do Conde, Portugal Indian Institute of Science Bangalore, Karnataka, Robert C. Dowler Department of Biology, Angelo State University Jorge Bolaños San Angelo, Texas, United States El Colegio de la Frontera Sur Departamento de Conservación de la Biodiversidad Ricardo Farrera-­Muro San Cristóbal de Las Casas, Chiapas, Mexico Instituto Tecnológico de Estudios Superiores de Monterrey Never Bonino Campus Puebla, Puebla, Mexico Instituto Nacional de Tecnologica Agropecuaria Bariloche, Argentina Craig Faulhaber Florida Fish and Wildlife Conservation Commission Christy J. Bragg Ocala, Florida, United States Drylands Conservation Programme The Endangered Wildlife Trust John E. C. Flux South 23 Hardy Street, Waterloo Lower Hutt, Wellington, New Zealand David E. Brown School of Life Sciences Johnnie French Arizona State University Department of Biology Tempe, Arizona, United States Portland State University Portland, , United States Arturo Carrillo-­Reyes Oikos: Conservación y Desarrollo Sustentable Antonio García-­Méndez San Cristóbal de Las Casas, Chiapas, Mexico El Colegio de la Frontera Sur Departamento de Conservación de la Biodiversidad Fernando A. Cervantes San Cristóbal de Las Casas, Chiapas, Mexico Instituto de Biología, UNAM Colección Nacional de Mamíferos Fernando Gopar-­Merino Distrito Federal, Mexico Centro de Investigaciones en Geografia Ambiental Universidad Nacional Autonoma de Mexico Kai Collins Morelia, Michoacán, Mexico Research Institute Department of Zoology & Entomology Thomas Gray University of Pretoria WWF—Greater Mekong Pretoria, Phnom Penh, Cambodia

Brian D. Cooke Klaus Hackländer Institute for Applied Ecology University of Natural Resources and Life Sciences Vienna University of Canberra Institute of Wildlife Biology and Game Management Canberra, Australia Vienna, Austria

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David C. D. Happold Weidong Li Research School of Biology Natural Ecological Protection Studio Australian National University Ürümqi, Xinjiang, China Canberra, A.C.T., Australia Andrey Lissovsky Jeremy Holden Zoological Museum, Moscow State University Flora and Fauna International Moscow, Cambridge, John A. Litvaitis Yeong-­Seok Jo Department of Natural Resources and the Environment Division of Research University of National Institute of Biological Resources Durham, New Hampshire, United States Incheon, South Marian Litvaitis Charlotte H. Johnston Department of Natural Resources and the Environment School of Life Sciences University of New Hampshire Arizona State University Durham, New Hampshire, United States Tempe, Arizona, United States Shaoying Liu Patrick A. Kelly Sichuan Academy of Forestry Department of Biological Sciences Chengdu, Sichuan, China California State University, Stanislaus Turlock, California, United States Consuelo Lorenzo El Colegio de la Frontera Sur Howard Kilpatrick Departamento Conservación de la Biodiversidad 391 Rt. 32 San Cristóbal de Las Casas, Chiapas, Mexico North Franklin, , United States Debbie Martyr Adrienne Kovach Fauna & Flora International-­Indonesia Programme University of New Hampshire Ragunan, Jakarta, Indonesia Durham, New Hampshire, United States Conrad A. Matthee Charles J. Krebs Department of Botany and Zoology Department of Zoology Stellenbosch University University of Stellenbosch, South Africa Vancouver, British Columbia, Jennifer L. McCarthy Hayley C. Lanier University of Delaware, Associate of Arts Program Department of Zoology and Physiology Wilmington, Delaware, United States University of Casper, Wyoming, United States Robert McCleery Department of Wildlife Ecology and Conservation John W. Laundré University of Florida James San Jacinto Mountains Natural Reserve Gainesville, Florida, United States Idyllwild, California, United States José Melo-­Ferreira Antonio Lavazza CIBIO, Centro de Investigação em Biodiversidade e Virology Department, IZSLER Recursos Genéticos Brescia, InBIO Laboratório Associado & Faculdade de Ciências Universidade do Porto Porto, Portugal

1709048_int_cc2015.indd 11 15/9/2017 15:59 xii Contributors

José M. Mora Luisa Rodríguez-­Martínez Instituto Internacional en Conservación y Manejo de Vida Centro Tlaxcala de Biología de la Conducta Silvestre Universidad Autónoma de Tlaxcala Universidad Nacional Tlaxcala, Mexico Heredia, Costa Rica Luis A. Ruedas Sanjay Molur Department of Biology Zoo Outreach Organization Portland State University Coimbatore, Tamil Nadu, India Portland, Oregon, United States

Dennis L. Murray Eugenia C. Sántiz-­López Department of Biology El Colegio de la Frontera Sur Trent University Departamento de Conservación de la Biodiversidad Peterborough, Ontario, Canada San Cristóbal de Las Casas, Chiapas, Mexico

P. O. Nameer Stéphanie Schai-­Braun Centre for Wildlife Studies University of Natural Resources and Life Sciences Vienna Kerala Agricultural University Institute of Wildlife Biology and Game Management Thrissur City, Kerala, India Vienna, Austria

Clayton K. Nielsen Lisa A. Shipley Cooperative Wildlife Research Laboratory and School of the Environment Department of Forestry Washington State University Southern Illinois University Pullman, Washington, United States Carbondale, Illinois, United States Andrew T. Smith Janet L. Rachlow School of Life Sciences Department of Fish and Wildlife Sciences Arizona State University University of Tempe, Arizona, United States Moscow, Idaho, United States Adia Sovie Juan Pablo Ramírez-­Silva Department of Wildlife Ecology and Conservation Programa Académico de Biología University of Florida Unidad Académica de Agricultura Gainesville, Florida, United States Universidad Autónoma de Nayarit Xalisco, Nayarit, Mexico Andrew Tilker Leibniz Institute for Zoo and Wildlife Research Chris Ray Berlin, Germany, and Institute for Arctic and Alpine Research Global Wildlife Conservation University of Colorado Austin, Texas, United States Boulder, Colorado, United States Zelalem Gebremariam Tolesa Tamara Rioja-­Paradela Department of Biology Sustentabilidad y Ecología Aplicada Hawassa University Universidad de Ciencias y Artes de Chiapas Hawassa, Tuxtla Gutiérrez, Chiapas, Mexico Myles B. Traphagen Terry J. Robinson Westland Resources, Inc. Department of Botany and Zoology Tucson, Arizona, United States Stellenbosch University Matieland, South Africa

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Julieta Vargas Rafael Villafuerte Instituto de Biología, UNAM IESA-­CSIC Colección Nacional de Mamíferos Campo Santo de los Mártires 7 Distrito Federal, Mexico Códoba, Spain

Jorge Vázquez Fumio Yamada Centro Tlaxcala de Biología de la Conducta Department of Wildlife Biology Universidad Autónoma de Tlaxcala Forestry and Products Research Institute Tlaxcala, Mexico Matsunosato 1, Tsukuba, Ibaraki,

Alejandro Velázquez Centro de Investigaciones en Geografia Ambiental Universidad Nacional Autonoma de Mexico Morelia, Michoacán, Mexico

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gramme (MAB-­ICC) added the Tres Marías Islands to the buffy brown. There is a significant amount of white on the UNESCO’s World Network of Biosphere Reserves, with dorsal surface of the hind feet. the aim of integrating conservation of biodiversity and The skull is medium-­sized with a long palate; it has management of natural resources. a broad braincase, a narrow breadth across the nasals, large maxillary and mandibular tooth rows, short incisive account authors: Consuelo Lorenzo, Juan Pablo foramina, a short diastema, a narrow basioccipital, and Ramírez-­Silva, Fernando A. Cervantes, and Ricardo medium-­sized auditory bullae, and is broad across the Farrera-­Muro carotid foramina with a narrow breadth across the infra- orbital canals. The shield-­bulla depth is shallow as well as Key References: Acevedo 1995; Ceballos and Navarro 1991; Cer- the skull depth. The supraoccipital shield is squared, and vantes 1997; Chapman and Ceballos 1990; Chapman et al. 1990; Diersing and Wilson 1980; Dooley 1988; Hall 1981; Lorenzo et al. the posterior section of the supraorbital process is slender 1993; Nelson 1899; SEMARNAT 2010; Wilson 1991. and flat. SIZE: Head and body 398–440 mm; Tail 40–45 mm; Hind foot 89–104 mm; Ear 60–76 mm; Greatest length of skull 77–78 mm CURRENT DISTRIBUTION: The Omiltemi rabbit has a Sylvilagus insonus (Nelson, 1904) very restricted distribution, less than 500 km² within the Sierra Madre del Sur, Guerrero, Mexico. This rabbit lives Omiltemi Rabbit in a highly fragmented area within a natural reserve, the Omiltemi State Park (Parque Ecológico Estatal Omiltemi) OTHER COMMON NAMES: Omiltemi cottontail; Conejo in the Sierra Madre del Sur. It occurs at elevations be- de Omiltemi, Conejo (Spanish) tween 2,133 and 3,505 m. DESCRIPTION: The Omiltemi rabbit is a medium-­sized TAXONOMY AND GEOGRAPHIC VARIATION: No sub- cottontail that has a short tail, medium-­sized hind feet, species. The phylogenetic relationship of S. insonus with and medium-­sized ears. The dorsal and nape pelage is other species of Sylvilagus is unclear. Morphological rufous black. The ears are black-­brown, the sides are comparisons between the tapetí (S. brasiliensis), Dice’s grayish black, and the tail is reddish black on the dorsal cottontail (S. dicei), and the Omiltemi rabbit indicate that surface and dingy buffy on the ventral surface. The sides the tapetí and Dice’s cottontail are more closely related of the nose and the orbital area have dingy buffy grayish to each other than either is to the Omiltemi rabbit. The coloration. The ventral surface of the is Omiltemi rabbit is one of the most poorly known of all mammals; it is represented by only five museum speci- ± mens, four complete and one with only a skin. ECOLOGY: Very little is known about the natural history of the Omiltemi rabbit. It is sympatric with the Mexican cottontail (S. cunicularius). It creates runways within the dense undergrowth, and occupies under rocks or other similar shelters. AND DIET: The preferred habitat of the Omiltemi rabbit is dense cloud forest, where pines (Pinus), (Quercus), and alders (Alnus) are abundant. BEHAVIOR: The Omiltemi rabbit is mainly a nocturnal species. PHYSIOLOGY AND GENETICS: Comparative phylogenetic analysis based on partial sequences (560 bp) of the gene 16S between eight species of Sylvilagus showed that the Omiltemi rabbit is most closely related to the desert cot- tontail (S. audubonii) and the Mexican cottontail instead of the tapetí as had been proposed previously. The sym- patric distribution between the Omiltemi rabbit and the

1709048_int_cc2015.indd 144 15/9/2017 16:00 Sylvilagus mansuetus 145 Mexican cottontail (in Guerrero, Mexico) seems to sup- ± port the close relationship between these species. Genetic analysis using partial sequences of cytochrome b gene revealed an average genetic divergence of 15% between the Omiltemi rabbit and the Mexican cottontail. : IUCN Red List Classification: Endangered (EN)— B2ab(ii,iii) National-­level Assessments: Mexico (Endangered Offi- cial Norm NOM-059-ECOL-2010 (SEMARNAT)) MANAGEMENT: Scientists consider the Omiltemi rabbit to be one of the most endangered mammals in the world. Deforestation is a serious threat to the Omiltemi rabbit, as the within its known range have been exten- sively logged, resulting in . Cattle grazing accompanies the deforestation and intensifies the threat. of the Omiltemi rabbit remains a concern even though most of the known range has been incorporated into a natural reserve area. The type local- ity of the Omiltemi rabbit is located within the Omiltemi State Ecological Park, which has been declared a natural reserve area (~ 3,613 ha). Intensive studies are necessary to determine whether the Omiltemi rabbit still exists and to confirm its current local distribution. Research and a risk assessment analysis should focus on habitat require- ments and the survival of this . Until such information is available, a comprehensive conser- vation strategy for this species and its habitat cannot be developed.

account authors: Consuelo Lorenzo, Fernando A. Cervantes, Julieta Vargas, and Ricardo Farrera-­Muro

Key References: AMCELA 2008c; Cervantes and Lorenzo 1997a; Cervantes et al. 2004; Chapman and Ceballos 1990; Diersing 1981; González-­Cózatl et al. 2007; Hall 1981; Jiménez-­Almaraz et al. 1993; Lorenzo et al. 2015; Nelson 1904, 1909; SEMARNAT 2010. Sylvilagus mansuetus. Photo courtesy Arturo Carrillo-­Reyes

obscured by the black tip of each hair. The sides of the body are paler gray than the back. The ears are brown to Sylvilagus mansuetus Nelson, 1907 pale mixed with gray and have whitish hairs on the inside. San José Brush Rabbit The sides of the neck are paler than the sides of the body, and the throat is gray mixed with white. The front legs are OTHER COMMON NAMES: Conejo de la Isla San José ochre-­colored clay, and the hind legs are yellowish white. (Spanish) The ventral part of the body is white mixed with gray, and DESCRIPTION: The San José brush rabbit is a medium-­ the ventral part of the tail is pale white. sized rabbit. The dorsal part is pale brown mixed with Proportionally, the skull of the San José brush rabbit is yellowish gray at the top of the back and head, slightly longer and narrower than that of the closely related brush

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