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 China 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

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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 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 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, 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, 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, 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, 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

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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, India 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, 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, 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, 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 Korea 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, 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, Canada 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, Japan

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

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impact infected rabbits. It is inferred that the rabbits have tremely long ears coupled with pearly white flanks, hips, either adapted to these parasites or were not infected with and rump. The long ears and lanky legs led to the formula- enough of them to be deleterious. tion of the famous ecogeographic “Allen’s Rule.” Dorsally : it is dappled in brown, gray, and black with distinctive IUCN Red List Classification: Data Deficient (DD) black nape markings and a throat patch of ochraceous MANAGEMENT: No studies have been conducted to de- buff. The white and black tail is short, and the huge ears termine any necessary management actions. It is assumed are nearly hairless and fringed in beige with little or no that the correlation between population declines of other black on the tips. The legs are long and slender, and the rabbit species in association with increased change resulting from agricultural activities would also negatively impact this species. Illegal hunting and increasing domes- ± tic and feral animal activity are also likely to have harmful impacts.

account author: Charlotte H. Johnston

Key References: Durrant and Guevara 2000a, 2000b, 2001.

Genus Lepus Linnaeus, 1758 Hares ( Lepus) are the largest of all lagomorphs. The 32 species of hare are nearly cosmopolitan, with many species being found on most continents (Africa, , , and ). They are not native to South America or Australia, although L. europaeus has been in- troduced on each of these continents. Some hare species are among the most northern in distribution of any terres- trial mammal; others inhabit deserts or open . Most features of hares represent an exaggeration of the features of the smaller lagomorphs: they have spectacu- larly large ears, hind feet, and eyes. They have four digits on both front and hind feet, although the first digit tends to be reduced. The dental formula is 2.0.3.3/1.0.2.3 = 28.

Lepus alleni Mearns, 1890 Antelope Jackrabbit

OTHER COMMON NAMES: Antelope hare, White-­sided jackrabbit, Allen’s jackrabbit; Liebre torda, Liebre blanca, Liebre appaloosa (Spanish) DESCRIPTION: The antelope jackrabbit has been con- sidered the handsomest and most striking of the North American hares. It was one of the last mammals in North America to be described, despite its large size. It has ex- Lepus alleni. Photo courtesy David E. Brown

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mostly fulvous head is large. The lower parts of the largely three L. alleni subspecies produced no clusters, indicating white feet are densely furred in cinnamon brown. The a weak if any differentiation between subspecies. Similar sexes are similar in appearance, and the reddish orange species include the white-­tailed jackrabbit (L. townsendii), eyes are set high on the head, giving a motionless animal a the white-­sided jackrabbit, and the Tehuantepec jack­rabbit, greater than 300° view of its surroundings. The female has all of which are allopatric to the antelope jackrabbit. six mammae, two pectoral and four abdominal. Two anal ECOLOGY: The antelope jackrabbit is most often encoun- glands secrete a yellowish substance that emits a musky tered in areas of low relief, including level ground, gently odor when the animal is excited. sloping plains, alluvial fans, and mesas. Fewer observa- The skull of the antelope jackrabbit is proportion- tions occur in broken country, and steep slopes appear to ally large, as are the zygomatic breadth and nasal bones. be used mainly as travel routes between mesas and arroyo Generous premaxillaries and molars contrast with the bottoms. It normally occurs between 185 and 1,200 m asl, similar, but more fragile-­appearing, skull of the black-­ placing the animal in mostly tropic-­subtropic environ- tailed jackrabbit (L. californicus). The frontal area is broad ments receiving mean annual rainfall amounts between and the rostrum is long and heavy as is the supraorbital 200 and 450 mm, of which a minimum of 90 mm falls and postorbital processes. The auditory bullae are small, during April through September. Minimum temperatures and the basioccipital is long and not deeply constricted. appear to influence the distribution of the antelope jack- Only one molt occurs annually, and the hairs cannot be rabbit; it has been documented only in areas where mean differentiated from those of the black-­tailed jackrabbit by annual temperatures exceed 18°C, and where the average cuticular examination. number of days 0°C or colder is less than 60. SIZE: Head and body 430–700 mm; Tail 50–140 mm; Home ranges of the antelope jackrabbit in good habitat Hind foot 113–145 mm; Ear 137–180 mm; Greatest in Arizona average 29.14 ha. Males tend to have larger length of skull 85–102 mm; Weight 2,500–4,400 g home ranges (35.97 ha) than females (19.57 ha). Popu- PALEONTOLOGY: Determined largely on the basis of size, lation sizes fluctuate dramatically depending on habi- L. alleni first appeared in the Early Irvingtonian ~ 1.1 tat condition, predator densities, and other factors not million ybp, where it might have been conspecific with yet identified. Most mortality appears to occur during the then widespread L. giganteus. Reported to occur as far the first six months of life. Known predators include east as Florida during the Pleistocene, attributed (Canis latrans), (Lynx rufus), mountain to L. alleni have been reported from such SW Pleistocene (Puma concolor), (Panthera onca), the larger sites as Ventana Cave in Arizona and Burnet Cave in hawks (Buteo spp.), great-­horned (Bubo virginianus), S New Mexico, the latter animal possibly being the white-­ and golden eagles (Aquila chrysaetos). Probable preda- sided jackrabbit (L. callotis). tors of L. a. tiburonensis on Tiburon Island are coyotes CURRENT DISTRIBUTION: The antelope jackrabbit is (C. l. jamesi), which are present in high density, and unique to Arizona and NW Mexico. Its range in Mexico possibly gray foxes (Urocyon cinereoargenteus) and rattle- includes four states and Tiburon Island in the Gulf of snakes (Crotalus molossus estebanensis). The main compet- California. The full extent of its geographic range is es- itor there for forage is the mule (Odocoileus hemionus timated at 102,000 km², with 12.5% (12,700 km²) of the sheldoni). The antelope jackrabbit is reported to be an im- animal’s distribution being in Arizona, 76% (78,000 km²) portant disperser of seeds via its fur and especially in , 10.5% (9,500 km²) in Sinaloa, and the re- its feces. maining 1% (1,800 km²) in W and N Nayarit. The antelope jackrabbit is sympatric with the black-­ Within this range it is restricted to the plains, valleys, and tailed jackrabbit in Arizona and much of N Sonora, foothills of the Sonoran and Sinaloan biotic provinces. although mixed groups of the two species are rarely en- With the exception of the similar Tehuantepec jackrabbit countered. The black-­tailed jackrabbit ranges both lower (L. flavigularis), the antelope jackrabbit has the most re- and higher in elevation than the antelope jackrabbit. An- stricted range of any non-­insular North American hare. telope jackrabbits also occur more often in groups than do TAXONOMY AND GEOGRAPHIC VARIATION: Three sub- black-­tailed jackrabbits. Observations on Tiburon Island in- species: L. a. alleni (Arizona and N Sonora); L. a. palitans cluded only solitary individuals and groups of two animals. (southern extension of the species range); L. a. tiburonen­ HABITAT AND DIET: Antelope jackrabbits are most com- sis (isolated on Tiburon Island in the Gulf of California). monly seen in areas of present or former Sonoran savanna Genetic comparisons are needed to verify the integrity of or a Sonoran savanna grassland / semi-­desert these taxa, as an analysis of 12 cranial characters of the grassland transition. Other observations occur in vegeta-

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tion characteristic of either the Arizona Upland subdivi- believed that the consumption of cacti pulp and fruits al- sion or Lower Colorado River subdivision of the Sonoran lows the hare to survive without free water and causes the Desert. These jackrabbits also use mesquite bosques and animal to pass urine in white, -­like splashes. thornscrub, and infrequently are seen in retired farmlands The diet varies with location and vegetation availabil- and buffelgrass (Pennisetum ciliare) pastures. Velvet mes- ity. Antelope jackrabbits feed intensely and often with quites (Prosopis velutina) are often a dominant over-­story green grasses and forbs being the primary food items plant, and no other plant is more often associated with an- during winter-­spring and summer monsoon rains when telope jackrabbit observations than this species. Relatively grasses constitute more than 80% of the diet. At other few observations occur in temperate oak woodlands and times the velvet mesquite is the preferred item, the hares semi-­desert grasslands more than 1,220 m in elevation. eating the leaves, bark, buds, and tufts of newly emerging Percentages of various classes of vegetation in the diet green leaves during the appropriate seasons. So heavy is of the antelope jackrabbit are grasses and forbs (45%), the use of this plant that browse-­lines can often be seen mesquite (36%), and cacti (8%). Important grasses in heavily populated areas, where the hares stand on include spidergrass (Aristida ternipes), purple threeawn their hind legs to reach green leaves and stem tips. Cacti, (A. purpurea), cane bluestem (Bothriochloa barbinodis), particularly barrel cacti, and prickly pear pads and fruit jackrabbit grass ( barbata), sideoats grama are fed on during dry seasons, presumably for the water (B. curtipendula), Rothrock’s grama (B. rothrockii), feather content as such use appears to decline where stock tanks fingergrass (Chloris virgata), bush muhly (Muhlenber­ or catchments are present. Cactus roots, mineral dirt, and gia porteri), witchgrass (Panicum capillare), whiplash the feces of other jackrabbits are also eaten. pappus­grass (Pappophorum vaginatum), and sand drop- BEHAVIOR: The antelope jackrabbit’s skipping run with seed (Sporo­bolus cryptandrus), all of which are believed to head held high at first appears comical and inefficient, provide both hiding cover and food. Interestingly, these but an especially energetic fourth or fifth bound enables grasses average 64 cm in height, approximately the mean the hare’s bulging eyes to survey the brushy terrain ahead length of an adult antelope jackrabbit. Antelope jackrab- while checking for any pursuers behind. When pressed, bits also occupy dense stands of tropical grasses found the animal pursues an all-­out run that can attain a speed along drainages and other more mesic areas, including of more than 70 km/h and may encompass leaps of up to sideoats grama, needle grama (Bouteloua aristidoides), vine 1.5 m in height. Rarely does the animal run in a straight mesquite (Panicum obtusum), Bermuda grass (Cynodon line, and if more than one animal is flushed each tends to dactylon), spear grass (Heteropogon contortus), sprangle- go its own way. Otherwise, when feeding, the hare hops tops (Leptochloa spp.), and Arizona cottontop (Digitaria along in a rather ungainly manner, often holding its hind- californica). quarters higher than its forequarters. The principal habitat requirement for the antelope Active year-­round, the antelope jackrabbit is primarily jackrabbit in Arizona appears to be a potentially her- a nocturnal and crepuscular animal that can be observed baceous understory within a velvet mesquite or other at any hour. During the day, individuals rest in shallow Sonoran vegetation matrix typical of a tropic-­subtropic depressions known as forms, typically under the cover of environment. Important forbs include such winter an- a mesquite or other shade-­providing . These forms, nuals as globe mallow (Sphaeralcea emoryi), native and which range from 8 to 15 cm wide and 28 to 46 cm long, non-­native mustards (Brassica spp., Sisymbrium spp., Les­ are typically free of rocks and sticks; these are temporary querella spp., Descurainia spp.), nievitas (Cryptantha spp.), structures that may be used repeatedly or only occasion- plantains (Plantago spp.), filaree (Erodium cicutarium), ally. Here the hare may be seen resting on its haunches or California poppy (Eschscholzia mexicana), broadfruit laid out like a dog with feet resting fore and aft. The ears combseed (Pectocarya platycarpa), and red-­maids (Calan­ may be erect or laid back, depending on the alertness of drinia ciliata). Summer rains result in significant areas the animal. Generally solitary, resting animals may also be occupied by Mexican poppy (Kallstroemia grandiflora), seen in pairs and less often in groups of up to a half-­dozen amaranths (Amaranthus spp.), and purselanes (Portulaca animals or more. spp.). While usually silent, individuals may squeal or scream Cacti are also frequently associated with the antelope when in distress, and thump their hind feet when jackrabbit, these succulents being present in about one-­ alarmed. Antelope jackrabbits are difficult to handle and third of the observations. Based on the evidence of tracks, maintain in captivity, the animals struggling or throwing fecal pellets, and teeth marks at the base of the cacti, it is themselves against the pen until serious injury occurs.

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Males may fight by engaging in “boxing matches” on tual recruitment rates appear much lower, however, and their hind legs, pummeling each other for up to a min- population size appears determined more by variation in ute or more. These contests, which are really scratching juvenile mortality than variation in recruitment rate. matches, continue until both participants are bloodied PARASITES AND DISEASES: Antelope jackrabbits are sus- and one of them leaves the field. ceptible to a variety of pathogens, and animals have been PHYSIOLOGY AND GENETICS: Diploid chromosome observed suffering from tumors, sores, and mange. Para- number = 48. Genetic analyses indicate a common an- site loads, especially during the warmer months, may be cestor of the white-­sided jackrabbit clade evolved with excessive, with up to 50% of the animals harboring bot- the increasing climatic isolation attendant with the (Cuterebra princeps, C. americana) in the throat and Pleistocene into the temperate white-­sided jackrabbit, rump areas. Tapeworm cysts (Taenia multiceps, Cittotaenia the tropic-­subtropic antelope jackrabbit, and the tropical spp.) are also common, as are an array of nematodes (e.g., Tehuantepec jackrabbit. Dermatoxys veligera, Nematodirus arizonensis, Passalurus The antelope jackrabbit possesses thermoregulatory, ambiguus). Ectoparasites include chiggers (Hexidionis physiological, and behavioral for surviving allredi), fleas (e.g., Hoplopsyllus glacialis affinis), and ticks in hot, arid environments and feeding on low-­quality (Haemaphysalis leporispalustris, Dermacentor albipictus, foods. Able to dissipate heat via its lanky extremities, the D. parumapertus). antelope jackrabbit also employs heat-­avoiding activity CONSERVATION STATUS: patterns such as feeding at night, use of shade, and solar IUCN Red List Classification: Least Concern (LC) alignment when resting. Having a normal body tem- MANAGEMENT: The antelope jackrabbit remains a com- perature of ~ 37.9°C, the antelope jackrabbit is able to mon animal within suitable habitat, but habitat losses withstand external temperatures of up to 51°C for up to within its limited range have been extensive due to agri- 4 hours. Its relatively high lethal body temperature of cultural and residential development. Because the species 45.4°C is avoided by an efficient mean water evaporation shuns open , extensive croplands, clean farms rate of less than 0.10% of body mass / h, which is ob- maintained by chemicals, and cleared pastures can limit tained through respiration rates that may be up to 350/ or eliminate populations. Additionally, vehicle collisions minute. The tolerance of antelope jackrabbits for cold is can constitute significant local sources of mortality, as less pronounced, animals showing symptoms of hypother- can predator irruptions, extremely cold weather, and, mia when temperatures were below 20°C. probably, disease. The animal has no legal status other The antelope jackrabbit does not require preformed than as “wildlife” in either Arizona or Mexico. Antelope water and is one of the largest “desert” animals thought jack­rabbits are shot for subsistence meat, sport, dog food, able to subsist entirely on metabolic water obtained from and bait to trap fur-­bearing carnivores. In the past, live plants. This is thought to be due to the large individuals have been used to train racing greyhounds. percentage of succulent cacti in the animal’s diet. Previously the species was regarded as a pest during years REPRODUCTION AND DEVELOPMENT: Antelope jack­ of high abundance on certain rangelands and in certain rabbits are thought to be promiscuous, with receptive agricultural fields. Such conditions are now of rare or no females mating with any sexually mature male encoun- occurrence. tered. Under favorable conditions they breed as early as 6 Research and monitoring are needed to address long-­ months of age. The breeding season extends from late De- term status and trend of populations, the animal’s long-­ cember to September, with one peak in spring and another term relationship with the black-­tailed jackrabbit, and, in summer. Timing of breeding appears to depend on tem- most important, to understand the driving forces relating perature and forage conditions. Mating activity consists of to its population dynamics. Understanding the relation- males fighting, chasing, and jumping, and a short copu- ships of individuals within groups would also be of scien- lation is sometimes accompanied by low-­pitched growls tific interest. or chucks. Gestation is about 6 weeks, and neonates are precocial, weighing from 133 to 184 grams at birth. The account authors: David E. Brown, Consuelo Lorenzo, fur-­lined nests are usually well concealed in a depression and Maria Altemus or vegetation. The young, which have a white spot in the Key References: Alcalá-­Galván and Miranda-­Zarazuá 2012; Allen center of the forehead, are more wary, more nocturnal, 1890, 1906; Altemus 2016; AMCELA 2008b; Armstrong and Jones, and more sedentary than adults. The mean litter size is 1.9 Jr. 1971; Arnold 1942; Best and Henry 1993; Brown et al. 2014; (range 1–5), with a potential of 3 to 4 litters/year. Ac- Caire 1978; Dawson and Schmidt-­Nielsen 1966; Dixon et al. 1983;

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