LAGOMORPHS

1709048_int_cc2015.indd 1 15/9/2017 15:59 1709048_int_cc2015.indd 2 15/9/2017 15:59 Lagomorphs Pikas, Rabbits, and Hares of the World

edited by Andrew T. Smith Charlotte H. Johnston Paulo C. Alves Klaus Hackländer

JOHNS HOPKINS UNIVERSITY PRESS | baltimore

1709048_int_cc2015.indd 3 15/9/2017 15:59 © 2018 Johns Hopkins University Press All rights reserved. Published 2018 Printed in China on acid-­free paper 9 8 7 6 5 4 3 2 1

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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: Lagomorpha. | BISAC: SCIENCE / Life Sciences / Biology / General. | SCIENCE / Life Sciences / Zoology / Mammals. | SCIENCE / Reference. Classification: LCC QL737.L3 L35 2018 | DDC 599.32—dc23 LC record available at https://lccn.loc.gov/2017004268

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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 Pika 58 Contributors ix Ochotona mantchurica, Manchurian Pika 60 Ochotona nubrica, Nubra Pika 61 1 Introduction 1 Ochotona opaca, Kazakh Pika 62 2 Evolution of Lagomorphs 4 Ochotona pallasii, Pallas’s Pika 65 3 Systematics of Lagomorphs 9 Ochotona princeps, American Pika 67 4 Introduced Lagomorphs 13 Ochotona pusilla, Steppe Pika 72 5 Diseases of Lagomorphs 18 Ochotona roylei, Royle’s Pika 75 6 Conservation of Lagomorphs 22 Ochotona rufescens, Afghan Pika 77 Ochotona rutila, Turkestan Red Pika 79 SPECIES ACCOUNTS Ochotona syrinx, Qinling Pika 82 Order Lagomorpha 29 Ochotona thibetana, Moupin Pika 83 Ochotona thomasi, Thomas’s Pika 84 Family Ochotonidae 31 Ochotona turuchanensis, Turuchan Pika 85 Ochotona alpina, Alpine Pika 31 Family Leporidae 87 Ochotona argentata, Helan Shan Pika 33 The Rabbits 87 Ochotona cansus, Gansu Pika 34 Brachylagus idahoensis, Pygmy Rabbit 87 Ochotona collaris, Collared Pika 36 Bunolagus monticularis, Riverine Rabbit 90 Ochotona coreana, Korean Pika 39 Caprolagus hispidus, Hispid Hare 93 Ochotona curzoniae, Plateau Pika 40 Nesolagus netscheri, Sumatran Striped Rabbit 95 Ochotona dauurica, Daurian Pika 43 Nesolagus timminsi, Annamite Striped Rabbit 97 Ochotona erythrotis, Chinese Red Pika 46 Oryctolagus cuniculus, European Rabbit 99 Ochotona forresti, Forrest’s Pika 47 Pentalagus furnessi, Amami Rabbit 104 Ochotona gloveri, Glover’s Pika 48 Poelagus marjorita, Bunyoro Rabbit 107 Ochotona hoffmanni, Hoffmann’s Pika 49 Pronolagus crassicaudatus, Natal Red Rock Hare 108 Ochotona hyperborea, Northern Pika 51 Pronolagus randensis, Jameson’s Red Rock Hare 110 Ochotona iliensis, Ili Pika 53 Pronolagus rupestris, Smith’s Red Rock Hare 111 Ochotona koslowi, Koslov’s Pika 55 Pronolagus saundersiae, Hewitt’s Red Rock Hare 113 Ochotona ladacensis, Ladak Pika 57 Romerolagus diazi, Volcano Rabbit 114

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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 taxonomy, world. This work is designed to expand coverage of the we have maintained a conservative systematic approach. world’s lagomorphs and update the 1990 LSG Lagomorph Nevertheless, we highlight relevant taxonomic issues that Action Plan (Rabbits, Hares and Pikas: Status Survey and require attention. Conservation Action Plan, compiled and edited by Joseph A. This work has been a team effort, with 82 specialists Chapman and John E. C. Flux). The Action Plan has contributing to species accounts. We especially thank served as the most thorough single source of informa- Rachel Fadlovich for her work on the references, and Aryn tion on lagomorphs for biologists, but it was never widely Musgrave for constructing all the species range maps. ATS available to the public and it has become outdated. In this thanks Harriet Smith for her insightful editorial work on book we present updated range maps of all lagomorph his chapters. We are grateful to all the photographers who species, high-­quality images of most species, as well as provided their work for free. We appreciate the meticulous current information on identification, systematics, ecol- copy editing by Maria E. denBoer. Finally, the collabora- ogy, behavior, reproduction, genetics, physiology, and con- tion with our editorial team from Johns Hopkins Univer- servation and management of the pikas, rabbits, and hares sity Press, Vincent Burke, Tiffany Gasbarrini, Debby Bors, of the world. The book also summarizes key components and Meagan M. Szekely, is highly appreciated.

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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, 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, 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 Mammal 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 Animal 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

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suggested that they prefer clearings in the forests. Not- the brightest in color. The lateral parts of the rabbit are withstanding, when forest is cleared, the areas formerly pale reddish; the whole venter is whitish; there is a brown occupied by Gabb’s cottontail become dominated, or section on the throat. even exclusively occupied, by the eastern cottontail. The size of the skull is medium-­large, presenting a long Because much of Central America’s formerly forested rostrum, a long diastema, and long incisive foramina. The area has become deforested or converted to agricul- maxillary tooth row is relatively short, and the basioccipi- ture, the conservation status of Gabb’s cottontail and tal is narrow. As in the marsh rabbit (S. palustris), the pos- its subspecies should be viewed with more than a little terior extensions of the supraorbital processes are fused to concern. In particular, these animals are elevationally the braincase throughout most of their length. restricted to forests in the tropical lowlands, which also SIZE: Head and body 437–480 mm; Tail 50–51 mm; Hind have seen the greatest degree of deforestation and hu- foot 91–99 mm; Ear 57–64 mm; Greatest length of skull man alteration of habitat. 78–80 mm CURRENT DISTRIBUTION: The Tres Marías cottontail account author: Luis A. Ruedas occurs on the four islands of the Tres Marías Islands Archipelago, off the state of Nayarit, Mexico. The extent Key References: Diersing 1981; Eisenberg 1989; Hall 1981; Hershko- of its known range is less than 500 km². The subspecies vitz 1950; Ruedas and Salazar-­Bravo 2007; Yates et al. 1979. S. g. graysoni occurs on the islands María Madre, María Magdalena, and María Cleofas, while S. g. badistes is found only on San Juanito Island. TAXONOMY AND GEOGRAPHIC VARIATION: Sylvilagus graysoni is closely related to the adjacent mainland species Sylvilagus graysoni (J. A. Allen, 1877) the Mexican cottontail (S. cunicularius). It is hypothesized Tres Marías Cottontail that the ancestral group of S. graysoni may have reached the islands during some early period of land connection OTHER COMMON NAMES: Tres Marías rabbit; Conejo de to the island. After the population became established on las Islas Marías, Conejo (Spanish) all four islands, the population on San Juanito Island must DESCRIPTION: The Tres Marías cottontail has relatively have become isolated from the others, resulting in charac- short ears, and its size varies from medium to large. The ters distinct enough to warrant subspecific recognition as dorsal pelage is rufous, and the nape and rump areas are S. g. badistes. ±

Sylvilagus graysoni. Photo courtesy Jorge Antonio Castrejón Pineda

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ECOLOGY: The Tres Marías cottontail occurs in decidu- San Juanito Island. They are common in old fields covered ous tropical forest up to 305 m asl where the vegetation is with a scattered growth of bushes. They become active predominantly composed of the false tamarind (Lysiloma around 1500 h. divaricatum), Cyrtocarpa procera, and Bursera spp. The Tres PHYSIOLOGY AND GENETICS: Diploid chromosome Marías cottontail has only a few historical predators: rac- number = 42 and FN = 78. The X chromosome is a small coons (Procyon lotor), red-­tailed hawks (Buteo jamaicensis), acrocentric or subtelocentric, whereas the Y chromo- and other prey birds such as caracaras (Polyborus plancus). some is a small acrocentric. The Tres Marías cottontail A few non-­native species have been introduced to the shares the diploid chromosome number with the Mexican islands through human activity, threatening the existence cottontail, the eastern cottontail (S. floridanus), and the of the cottontails, especially due to the competition for desert cottontail (S. audubonii). resources. White-­tailed deer (Odocoileus virginianus), pigs REPRODUCTION AND DEVELOPMENT: Little is known of (Sus scrofa), and domestic goats (Capra aegagrus hircus) reproduction in the Tres Marías cottontail. Eleven females were introduced deliberately to María Magdalena Island, were lactating and 2 had 75 mm embryos in March. while the introduction of the black rat (Rattus rattus) to CONSERVATION STATUS: all the islands was purely accidental. IUCN Red List Classification: Endangered (EN)— HABITAT AND DIET: The habitat on Tres Marías Islands B2ab(iii,v) is more arid than on the mainland. Average yearly rainfall National-­level Assessments: Mexico (Endangered Offi- is 635 mm, most of which falls in summer, resulting in a cial Norm NOM-059-ECOL-2010 (SEMARNAT)) tropical dry deciduous and moist forest habitat. Tempera- MANAGEMENT: The Tres Marías cottontail was reported tures are moderate although recorded extremes are 4.6°C to be abundant on Tres Marías Islands in 1897. However, and 37.5°C; monthly averages range from 20.3°C in Janu- almost a century later no evidence of activity of rabbits ary and February to 28.1°C in July and August. was found on María Madre, María Magdalena, and María San Juanito Island is covered with dense stands of Cleofas, although some individuals were observed on San trees, bushes, and cacti 3–4 m high. The natural vege- Juanito Island. tation on coastal areas of María Madre Island is tropical Exotic species and farming have had strong negative deciduous forest with a canopy height of 4 m. Most of this ecological impacts on the rabbit and contributed to its forest was removed by humans for agricultural and gov- population decline, although the main cause has been ernmental purposes, due to the presence of a penal colony human activities on the islands. An example of this is the there of 2,000–3,000 habitants. The island originally was situation on María Cleofas, which has been stripped of forested with trees up to 30 m, but most of that vegetation about half of its native vegetation in preparation for hu- has been logged and removed. man settlement. María Magdalena Island was designated an ecological Some vital actions to ensure survival of the Tres Marías reserve by the Mexican government in 2000. This island cottontail are to cease all forms of hunting, turn San has a less disturbed forest and a higher vegetation density Juanito Island into an ecological reserve, create a biolog- than found on María Madre Island. Since there are no ical reserve on María Madre Island, and support scien- permanent residents in this area, the abundance of the tific investigation to assess the rabbit’s present status and Tres Marías cottontail there is higher than on the other is- specific habitat conditions. A detailed long-­term study has lands. On María Cleofas Island, the forest appears to be in strongly recommended the need to identify the current good condition; most of the vegetation is open woodland state of conservation and the basic biological characteris- around the coast that grows thicker than that of the cen- tics of the species. ter of the island. The eastern side of the island has been Since 1905, the Tres Marías Islands have housed a fed- cleared in preparation for plantations. In addition, a small eral penal colony. In 1998, the archipelago was listed as military detachment stays near their camp on that side of a priority area for conservation; in 2000, the archipelago the island. was declared a protected Natural Area under the category BEHAVIOR: The Tres Marías cottontail is an easy-­to-­catch of biosphere reserve based on the good state of conserva- rabbit since, surprisingly, it does not fear humans. Rabbits tion of its ecosystems and the degree of endemism of the from all islands show far less fear of people than do their species of flora and fauna that it contains. mainland counterparts. This lack of escape behavior is On 2 June 2010, the International Coordinating particularly apparent in the Tres Marías cottontails from Council of UNESCO’s Man and the Biosphere Pro-

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

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