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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, 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 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 87 Ochotona argentata, Helan Shan Pika 33 The Rabbits 87 Ochotona cansus, 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, 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, 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

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 , 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, 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, 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, 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

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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 Fish and Wildlife Conservation Commission Christy J. Bragg Ocala, Florida, United States Drylands Conservation Programme The Endangered Wildlife Trust John E. C. Flux South Africa 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, Oregon, 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, South Africa 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, , China Canberra, A.C.T., Australia Andrey Lissovsky Jeremy Holden Zoological Museum, Moscow State University Flora and Fauna International Moscow, Russia Cambridge, United Kingdom John A. Litvaitis Yeong-­Seok Jo Department of Natural Resources and the Environment Division of Research University of New Hampshire 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 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, Connecticut, 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 British Columbia 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 Wyoming 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, Italy InBIO Laboratório Associado & Faculdade de Ciências Universidade do Porto Porto, Portugal

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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 Idaho 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 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, Ethiopia 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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populations of the Japanese hare. In the north (Yamagata Key References: Abe et al. 2005; Fujimoto et al.1986; Fujita 2004; Prefecture), the breeding season extends from February Hirakawa 2001, Hirakawa et al. 1992; Imaizumi 1960; Kawamura to July with young born between April and August, mostly et al.1989; Nunome et al. 2010, 2014; Otsu 1974; Shimizu and Shimano 2010; Takada and Yamaguchi 1974; Taniguchi 1986; in May and June, while in the south (Kagoshima Prefec- Temminck 1844; Torii 1989, 1990; Tsuchiya 1979; Wu et al. 2005; ture) the breeding season is year-­round. The average litter Yamada 1987, 1990, 2014, 2015a; Yamada et al. 1988, 1989, 1990, size is 1.86 (range 1–4) in the north and 1.16 (range 1–3) 2002; Yamaguchi et al. 2008; Yamaguti 1935; Yatake et al. 2003. in the south. Gestation period is 42–43 days in the north and 45–48 days in the south. Weight at birth is 77–165 g in the north and 125–150 g in the south. Females become sexually mature at 8 to 10 months. Growth rate is faster Lepus californicus Gray, 1837 and adult body weight is 20% heavier in the north than in the south in order to survive winters. Females are in- Black-­tailed Jackrabbit duced ovulators, and ovulation occurs 12–15 hours after stimulation of copulation. Parturition takes place in a OTHER COMMON NAMES: California jackrabbit, Gray-­ shallow den (e.g., 5 cm in depth and 20 cm in diameter) sided jackrabbit, Narrow-­gauge mule, Jackass-­hare, Texas dug by the female, and the duration of parturition is short jack, Texan hare, Great Plains jackrabbit; Liebre de cola (2 minutes). Newborns can run approximately 1 hour negra, Liebre (Spanish) after delivery, owing to precocity. Young begin to feed on DESCRIPTION: The black-­tailed jackrabbit is the most plants on day 8, but are nursed until 1 month after birth, widely distributed North American jackrabbit and is suckling occurring only at midnight for approximately 2 medium-­sized for all measurements, relative to other minutes. Average life span is 1 year, and the maximum Lepus species. The pelage is buffy gray or sandy-­colored is around 4 years. Thirty percent of newborns survive to above, peppered with black, but white below. The tail year 1. is whitish or pale gray underneath and black above; the PARASITES AND DISEASES: The Japanese hare is known black coloration extends onto the rump. The ears are very to be infected by several cestodes, including Mosgovoyia long, conspicuously tipped with black, and white on the pectinata and Taenia pisiformis. Ectoparasites include ticks back. Eye-­shine at night is reddish. (Haemaphysalis flava, Ixodes nipponensis, and I. persulca­ The interparietal bone is fused to and indistinguishable tus). The Japanese hare can be infected by Francisella tu­ from the parietal bones. The rostrum is narrow and long, larensis causing tularemia, and it is an important zoonosis and tapers anteriorly. Occipito-­nasal length is greater than from the hare to . : IUCN Red List Classification: Least Concern (LC) ± National-­level Assessments: Japan Ministry of the En- vironment for L. b. lyoni (Near Threatened (NT)—criteria a and b) MANAGEMENT: The Japanese hare is a game species. The current status of the Japanese hare is declining due to the recent reduction of preferable . One subspecies, L. b. lyoni, endemic to Sado Island (855 km2) in Niigata Prefecture, is experiencing a severe decline due to the reduction of preferable habitat and the impact of introduced Japanese martens. During the 1950s and 1960s, 53 Japanese martens and seven red foxes were in- troduced from Honshu to the island for biological control to reduce feeding damage of tree seedlings by the hares. The martens covered all of the island, but the foxes did not successfully colonize.

account author: Fumio Yamada

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tis, L. c. merriami, L. c. richardsonii, L. c. sheldoni, L. c. texi­ anus, L. c. wallawalla, and L. c. xanti. The integrity of some of these “ecotypes” is questionable, and the phylogenetic relationships of these subspecies need to be evaluated. Genetic analyses highlight the distribution of subspecies of L. californicus on the Baja California Peninsula, Mexico, into three groups: L. c. xanti from La Paz, Santa Anita, and San Jose del Cabo; L. c. magdalenae from the La Paz Isth- mus to S Vizcaino Desert; and L. c. martirensis from the Vizcaino Desert and northward. Similar species include the white-­tailed jackrabbit, the antelope jackrabbit (L. alleni), and the white-­sided jackrabbit. The black-­tailed jackrabbit does not inter- breed with the closely related and sympatric antelope and white-­sided jackrabbits. The specific status of the black jackrabbit (L. insularis) has been questioned by research- ers who consider that it may be only a melanistic form of L. californicus. The nuclear and mitochondrial differences, Lepus californicus. Photo courtesy Randall D. Babb color variation, and small differences in the structure of a single bone in the skull (jugal) between L. insularis and 75 mm. The pronounced supraorbital process has an ante- L. californicus suggest that the two lineages may have di- rior projection, and its posterior projection is usually long, verged recently. Ongoing research may further clarify the tapering, and fused to the cranium. relationship between these two forms. SIZE: Head and body 430–700 mm; Tail 50–140 mm; ECOLOGY: The black-­tailed jackrabbit is a shrub-­ Hind foot 113–145 mm; Ear 137–180 mm; Greatest associated hare that has extraordinary thermoregulatory, length of skull 85.4–101.9 mm; Weight 1,300–3,600 g physiological, and behavioral for surviving in PALEONTOLOGY: Fossil remains and DNA analysis show hot, dry regions on low-­quality foods. Population densities that the black-­tailed jackrabbit is genetically distinct from can vary from 0.1 to 1.2 /ha in natural and the sympatric white-­tailed jackrabbit (L. townsendii) and from 1.5 to 208/ha adjacent to and on agricultural lands. more closely related to the white-­sided jackrabbit (L. cal­ Home range sizes typically vary between 10 and 20 ha, but lotis) clade, this taxon evolving from a common ancestor can range from 4 to 140 ha. Population sizes can fluctuate some 1.1 million ybp. Centered in the Intermountain Re- dramatically depending on weather, land-­use practices, gion of W North America, the species greatly expanded its and predator-­control activities. Although population range during the Holocene. fluctuations may be influenced by precipitation amounts CURRENT DISTRIBUTION: The black-­tailed jackrabbit oc- and variations in recruitment rate, most fluctuations are curs across the southern two-thirds­ of the W and C United the result of changes in mortality rate. The black-­tailed States, as well as in 15 states in Mexico (down Baja Cali- jackrabbit is a keystone prey species for (Canis fornia and the central northern portion of Mexico). Dis- latrans), foxes (Vulpes spp., Urocyon cinereoargenteus), junct populations occur in SW Montana, SE Oklahoma, badgers (Taxidea taxus), (Lynx rufus), large buteos and several islands off Baja California. The black-­tailed (Buteo spp.), great-­horned (Bubo virginianus), and jackrabbit has been successfully introduced in New Jersey, especially golden eagles (Aquila chrysaetos). Massachusetts, Maryland, Virginia, and Florida. The spe- The black-­tailed jackrabbit molts only once annually. It cies ranges in elevation from –84 m to 3,750 m. Another is also an important disperser of plant seeds, on its and form recently introduced (1960–1991) to Cerralvo Island, especially via its feces. Baja California Sur, Mexico, is considered L. c. xanti. HABITAT AND DIET: The black-­tailed jackrabbit inhab- TAXONOMY AND GEOGRAPHIC VARIATION: Seventeen its more-­arid areas as diverse as dry , savannas, subspecies: L. c. altamirae, L. c. asellus, L. c. bennettii, , deserts, croplands, shrublands, and dune areas L. c. californicus, L. c. curti, L. c. deserticola, L. c. eremicus, (in island populations). It typically inhabits open country L. c. festinus, L. c. magdalenae, L. c. martirensis, L. c. melano­ with scattered thickets or patches of shrubs; consequently,

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its numbers are often high in intensively grazed areas ad- or tularemia (Francisella tularensis), and other bacterial jacent to cultivated fields. This animal is one of the most pathogens have been found in a number of western U.S. widespread and numerically common inhabitants of Great populations. Jackrabbits are prone to harbor the tick Basin, Mohave, Sonoran, and Chihuahuan desert-­scrub Dermacentor parumapertus and other vectors for tulare- communities, as well as plains, intermountain, and semi-­ mia (Q-­fever, Coxiella burnetii). Rocky Mountain spotted desert grasslands. fever (Rickettsia rickettsii) has been reported as occurring The diet varies with vegetation availability and lo- in western U.S. populations, although actual investiga- cation. This jackrabbit forages on grasses, forbs, crops, tions have failed to find any serious disease outbreaks and hay in summer, and buds, bark, and leaves of woody in western U.S. jackrabbits. An investigation of a high-­ plants in winter. Individuals obtain water both from vege- density population of black-­tailed jackrabbits near Battle tation and by re-­ingesting soft fecal pellets; drinking free Mountain, Nevada, in 1951 failed to incriminate either water has also been reported. tularemia or plague in the “die-­off.” heT only pathogens BEHAVIOR: Active year-­round, the black-­tailed jack­rabbit noted were antibodies for Colorado tick fever and west- is often crepuscular. Most feeding occurs at night, yet ern equine encephalomyelitis, neither of which could be the species can be observed at any hour. During the day, implicated as a serious cause of mortality. individuals rest in shallow depressions (forms), typically Parasite infestations have also been suggested as con- under the shade of a shrub or small tree. In the hottest tributing to reduced jackrabbit numbers, especially after regions, these are extended to shallow , which are a population experiences a rapid decline. But although used for 3–5 hours per day. Exceptional individuals may high infestations and the presence of botflies travel up to 16 km round-­trip to feed in fields. Generally (Cuterebra spp.), tapeworms (Taenia spp.), ticks, and lice solitary yet feeding in loose and surprisingly large groups, often accompany high jackrabbit densities, most para- black-­tailed jackrabbit individuals will typically “freeze” site infestations wax and then wane with the cessation of when approached. However, when one knows it has been warm weather. seen, it will jump up suddenly and flee at up to 64 km/h, CONSERVATION STATUS: occasionally stopping to see if it is pursued. When chased IUCN Red List Classification: Least Concern (LC) by predators at moderate speeds, black-­tailed jackrabbit MANAGEMENT: The black-­tailed jackrabbit is hunted for individuals will periodically bounce in an exceptionally subsistence, sport, and food. Skins have been used for high “observation leap,” which may reach 6 m in height. hats, and live individuals have been used to train racing Although usually silent, individuals may squeal or scream . Because during years of high abundance the species when in distress and thump their hind feet when alarmed. is regarded as a pest in certain areas, due to damage to PHYSIOLOGY AND GENETICS: Diploid rangeland, hayfields, and cultivated crops, local control number = 48 and FN = 82 efforts may reduce numbers temporarily. From the late REPRODUCTION AND DEVELOPMENT: Breeding is 1800s to the mid-­1950s, tens of thousands of black-­tailed promiscuous, with females accepting the first interested jackrabbits were poisoned or killed in “jackrabbit drives” male. The breeding season varies with latitude and en- to protect crops and provide more forage for livestock. vironmental factors. In Idaho, breeding is restricted to However, these efforts generally ceased or were much February to May; in the SW United States, it can last from reduced after 1980. December to September. Gestation ranges from 40 to 47 Because the black-­tailed jackrabbit is not designated days. Litter size varies from 3.8 to 4.9 in the north to 1.8 a game animal in most states, trends in annual harvests to 2.2 in the south, giving a total output per female of and hunt success trends are only available regionally. De- about 10–14 precocial offspring per year. Actual recruit- clines in hunter and harvest numbers have been reported ment rates, however, are much lower; the ratio of young in California (strongly so), Colorado, Nebraska, Nevada, to adult animals does not usually exceed 1:1. Early-­born and Oklahoma. Abundance surveys in Nebraska suggest females can breed in their year of birth. Average life span a declining trend in the black-­tailed jackrabbit there, and averages 1.4–1.8 years, but one individual in captivity Washington closed the season on all jackrabbits in 2001. lived 6.75 years. Vehicle collisions can constitute significant local PARASITES AND DISEASES: Diseases and parasites have sources of mortality, as can forest fires, drought, hail, long been suspected as being agents capable of reduc- tularemia outbreaks, and extremely cold weather. Be- ing black-­tailed jackrabbit numbers, and rabbit fever, cause the species favors open habitats such as croplands,

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increasingly intensive chemical farming, pest control, and reddish. The white-­sided jackrabbit undergoes one molt removal of all native cover may limit populations. There is a year during early summer from front to rear. Except for concern about low encounter rates of the species during the shorter pelage in summer, there is little difference thousands of kilometers of roadside surveys in western between the summer and winter coats. Females are larger Oregon. Rates of sightings along dirt roads across Nevada than males. during 1995–2015 crepuscular and nocturnal surveys The skull is similar to that of the more common black-­ were two-­thirds to one-­sixth of those reported in field tailed jackrabbit (L. californicus), but with a higher nasal notes from the 1930s to 1950s. aperture, a smaller and more inclined supraorbital sur- Research and monitoring are needed to address the face, and a lesser breadth across the auditory bullae. long-­term status and trend of the black-­tailed jackrabbit, SIZE: Head and body 432–598 mm; Tail 47–90 mm; including its relationship with management objectives Hind foot 113–145 mm; Ear 137–180; Greatest length of and broad-­scale drivers of population dynamics, genetics skull 85–93 mm; Weight 1,500–3,200 g (e.g., more-­resolved subspecies definitions), and gaps in distribution and synecological relationships between the black-­tailed jackrabbit and other sympatric jackrabbits. ±

account authors: Erik A. Beever, David E. Brown, and Consuelo Lorenzo

Key References: AMCELA 2008a; Best 1996; Bowen et al. 1960; Cervantes et al. 1999–2000; Clemons et al. 2000; Desmond 2004; Fagerstone et al. 1980; Flinders and Chapman 2003; Flux and Angermann 1990; French et al. 1965; Grayson 1977; Hall 1981; Hinds 1977; Johnson and Anderson 1984; Knick and Dyer 1997; Lechleitner 1958a, 1958b, 1959; Lorenzo et al. 2014b; Marín et al. 2003; Nelson 1909; Palmer 1897; Rice and Westoby 1978; Schmidt-­ Nielsen et al. 1965; Simes et al. 2015; Smith 1990; Smith and Ny- degger 1985; Smith et al. 2002; Stoddart 1985; Vorhies and Taylor 1933; Wywialowski and Stoddart 1988.

Lepus callotis Wagler, 1830 White-­sided Jackrabbit

OTHER COMMON NAMES: Antelope rabbit, Beautiful-­ eared jackrabbit, Gaillard’s jackrabbit, Snowsides; Liebre torda, Liebre pinta (Spanish) DESCRIPTION: The white-­sided jackrabbit is a medium to large hare with a buffy cinnamon brown dorsum merg- ing to iron gray on the rump and hips, with distinctive white sides and underparts below the median line. The ears, while large, are not exaggerated and are tipped with a white fringe and sport dusky patches along the poste- rior edges. A nape patch ranges from brown to black, and the limbs are white with buff on the upper surfaces. The gular area is cinnamon to ochraceous with much white on the head. The tail is moderately short with a black upper surface and white underparts. The eye-­shine at night is Lepus callotis. Photo courtesy Martha Marina Gomez Sapiens

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