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

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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, 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 87 Ochotona argentata, Helan Shan Pika 33 The Rabbits 87 Ochotona cansus, Gansu Pika 34 Brachylagus idahoensis, Pygmy 87 Ochotona collaris, Collared Pika 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, Daurian Pika 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, 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, 114

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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 , 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 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

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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 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, , 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, 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 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, 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 Forest 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

1709048_int_cc2015.indd 13 15/9/2017 15:59 This page intentionally left blank 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. CONSERVATION STATUS: 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 forests within its known range have been exten- sively logged, resulting in fragmentation. Cattle grazing accompanies the deforestation and intensifies the threat. Hunting 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 threatened species. 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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rabbit (S. bachmani), and the supraorbital process is fused and the jugal longer. SIZE: Head and body 303–340 mm; Tail 24–42 mm; Hind foot 72–82 mm; Ear 59–73 mm; Greatest length of skull 60.7–65.4 mm; Weight 620–1,300 g CURRENT DISTRIBUTION: The San José brush rabbit is an endemic rabbit species on San José Island in the Gulf of California, Baja California Sur, Mexico. The rabbit is only known from a restricted range of approximately 20 km² along the southwestern coastal plain of San José Island. TAXONOMY AND GEOGRAPHIC VARIATION: No sub- species. It is believed that the San José brush rabbit is closely related to the brush rabbit (S. bachmani) from mainland Baja California (Mexico). The San José brush Sylvilagus mansuetus habitat. Photo courtesy Tamara Rioja-­Paradela rabbit may be distinguished from the brush rabbit by its larger ears and paler pelage. pitaya (Stenocereus gummosus), sweet pitaya (S. thurberi), ECOLOGY: In 2008, the population density of the San José wild plum (Cyrtocarpa edulis), a local rue (Esenbeckia brush rabbit was estimated to be 25–35/km² in an area of flava), desert thorn (Lycium spp.), and ironwood (Olneya optimal habitat of approximately 4 km². In 2011, the aver- tesota). It is believed that the San José brush rabbit feeds age density was 11.4/km2 in xeric scrub in lowlands below only on green plants. 200 m asl. The current estimated number of individuals BEHAVIOR: The San José brush rabbit is solitary. The in this area is 2,087 (range 786–5,547). rabbit is most active between sunset and 0200 h and The San José brush rabbit shares its habitat with from about 0600 to 1000 h; therefore, it is nocturnal and other species of fauna that include mammals such as the crepuscular. During the day the rabbits rest under palo peninsula mule deer (Odocoileus hemionus peninsulae); verde, sour pitaya, sweet pitaya, ashy limberbush, cholla, several species of mice, including the San José kangaroo mesquite, cardon cactus, Adam’s tree, and goatnut. rat (Dipodomys merriami insularis), Bryant’s spiny pocket PHYSIOLOGY AND GENETICS: Diploid chromosome mouse (Chaetodipus spinatus bryanti), the northern Baja number 2n = 48 and FN = 80. Phylogenetic analysis deermouse (Peromyscus fraterculus cinereus), and the no- based on partial sequences (560 bp) of the gene 16S palera rat (Neotoma bryanti bryanti); carnivores, including showed that the San José brush rabbit is basal in position the northern cacomistle (Bassariscus astutus insulicola) with respect to seven other species of Sylvilagus, which and cats (Felis sylvestris); donkeys (Equus asinus) and goats suggests that the San José brush rabbit followed an evo- (Capra aegagrus hircus); reptiles such as whiptail lizards lutionary path that is different from that of most species (Aspidoscelis danheimae), rattlesnakes (Crotalus enyo evaluated, and that it shares this quality with the brush enyo, C. mitchellii mitchellii, C. ruber lucasensis), and go- rabbit because both species have the same diploid chro- pher (Pituophis catenifer bimaris, P. vertebralis); and mosome number (48). The basal position of the San José birds such as ospreys (Pandion haliaetus), red-­tailed hawks brush rabbit within the genus supports the proposal that (Buteo jamaicensis), peregrine falcons (Falco peregrinus), the karyotype of 2n = 48 is a fundamental element in the and American kestrels (F. sparverius). evolution of the Leporidae, particularly with respect to HABITAT AND DIET: San José Island hosts principally the diversification of forms within the genus Sylvilagus. xeric scrub; there is also a zone of reduced size where The minor nuclear and mitochondrial differences, and mangrove vegetation is found. The area of highest abun- color variation between S. mansuetus and S. bachmani, dance of the San José brush rabbit is characterized by a indicate that this form could be considered a subspecies of rich diversity of desert trees, cacti, and bushes: Adam’s S. bachmani. tree (Fouquieria diguetii), ashy limberbush (Jatropha cine­ REPRODUCTION AND DEVELOPMENT: The breeding rea), mesquite (Prosopis spp.), cardon cactus (Pachycereus phase in the San José brush rabbit apparently takes place pringlei), cholla (Opuntia cholla), copal tree (Bursera hind­ during the wet season, based on finding breeding sites in siana), elephant tree (B. microphylla), goatnut (Simmond­ November, pregnant females in June, and males with scro- sia chinensis), palo verde (Parkinsonia microphylla), sour tal testes in June. Museum specimens of collected

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in June also indicated reproductive activity in the wet hunting (sport or subsistence) that have been previously season: males with large (19 × 8 mm) scrotal testes and reported and considered within its management programs females containing advanced embryos. to eradicate exotic fauna and control hunting. It is neces- CONSERVATION STATUS: sary to develop research projects to determine population IUCN Red List Classification: Critically Endangered dynamics, reproduction, ethology, and other aspects of (CR)—B1ab(ii,iii,v) the rabbit’s biology and ecology that have been ignored. National-­level Assessments: Mexico (Endangered Offi- cial Norm NOM-059-ECOL-2010 (SEMARNAT)) account authors: Consuelo Lorenzo, Tamara Rioja-­ MANAGEMENT: All wildlife on San José Island is pro- Paradela, Arturo Carrillo-­Reyes, Mayra de la Paz, and tected under Mexican law, but local people and fishers Sergio Ticul Álvarez-­Castañeda often illegally kill protected mule deer and San José brush rabbits when they are legally hunting invasive goats. This Key References: Álvarez-­Castañeda et al. 2006; Case and Cody 1983; Cervantes et al. 1996, 1999; Chapman 1974; Gómez-­Nísino 2006; activity can be a threat to the entire rabbit population, González-­Cózatl et al. 2007; Hall 1981; López-­Forment et al. 1996; although more information is necessary to document the Lorenzo et al. 2011, 2014b, 2015; Nelson 1907; SEMARNAT 2010; extent of the hunting. Also, feral goats may negatively Thomas and Best 1994a; Worthington 1970. compete with the rabbits by eating and destroying native vegetation. Predation by native and non-­native species constitutes a threat to the entire population of the San José brush rabbit. The most serious threat is from feral cats that Sylvilagus nuttallii (Bachman, 1837) frequent the area of occupancy and commonly prey on Mountain Cottontail rabbits. Domestic dogs in areas occupied by people may also constitute a predation threat to the rabbits. Predation OTHER COMMON NAMES: Nuttall’s cottontail by cats and maybe dogs is especially critical given the pu- DESCRIPTION: The mountain cottontail is a small to tative tameness of this insular species. It is also believed medium-­sized Sylvilagus. The back and rump are gray or that the northern cacomistle may prey on rabbits, but buff, the sides are paler, and the belly is white. The nape this predator mainly lives in the rocky hills and not in the of the neck is orange. The hind legs are long, and the hind area of rabbit occupancy on the island; thus they may be feet are whitish or pale orange and are covered with long, only occasional and opportunistic predators. Several other dense hair. Both the front legs and the front feet are rusty-­ species on San José Island could prey on rabbits, including the three species of rattlesnake, both species of gopher , ospreys, red-­tailed hawks, peregrine falcons, and ± American kestrels. The gopher snake (P. vertebralis) has been observed eating a breeding female rabbit. The most important threat to the San José brush rabbit is related to human development activities, in spite of its protected status on San José Island under Mexican law. Plans for a resort are being developed with an accompany- ing golf course, private airport, and small marina in prime habitat occupied by the rabbit. This development would cause a loss of habitat (area of occupancy) for the rabbit, including breeding and reproduction sites. Additionally, a salt mine close to the area of highest rabbit density may be activated in the near future, and this facility could be used for worker housing and equipment storage. Free-­ranging dogs and cats belonging to the workers would then con- tribute to further predation on the rabbit. It is urgent to assess pressures caused by the destruc- tion of the habitat of the San José brush rabbit and illegal

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