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Home , Alaskan hare, Amami rabbit, Bunyoro rabbit, Desert cottontail, Hispid hare, Mexican cottontail

LAGOMORPHS

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

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

JOHNS HOPKINS UNIVERSITY PRESS | baltimore

1709048_int_cc2015.indd 3 15/9/2017 15:59 © 2018 Johns Hopkins University Press All rights reserved. Published 2018 Printed in 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

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, 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, 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, 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, , 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, , 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 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 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 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 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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whereas the postorbital processes of the eastern cottontail Manzano Mountains are separated in the Tijeras Can- are homogeneously thick and fused with the frontal bone yon by a large freeway, which constitutes an impassable throughout a greater proportion of their length (~ 50%). barrier to the Manzano Mountain cottontail. Thus, the The dentition is characteristically distinct. The groove in Sandia and Manzano populations are functionally discon- the first upper incisor of the Manzano Mountain cotton­ tinuous. Additionally, suitable in the Sandia range tail is more lingually located than that of the eastern only covers ~ 100–110 km2, an area that is shrinking due cottontail; in the eastern cottontail, the ratio of labial to to the effects of global warming. Furthermore, the Sandia lingual lobes is between 50:50 and 60:40, whereas in the Range is bisected by the heavily trafficked road from the Manzano Mountain cottontail, the labial lobe is closer to town of Sandia Park on the east slope, to Sandia Peak and two-­thirds of the tooth. its ski area. The threats to the conservation and the well-­ SIZE: Head and body 384–462 mm; Tail 55–75 mm; being of this species are therefore somewhat severe. Hind foot 100–105 mm; Ear 67–73 mm; Greatest length Additional museum records identified as S. cognatus ex- of skull 68–72 mm; Weight > 1,100 g ist from the Capitan Mountains and the Mount Withing- CURRENT DISTRIBUTION: The Manzano Mountain ton Range, also in . These should be viewed cotton­tail occurs only in the Sandia and Manzano Moun- askance, particularly the latter, as it is separated from the tains of C New Mexico. Sandia and Manzano Ranges not only by extensive areas TAXONOMY AND GEOGRAPHIC VARIATION: No sub- of desert, but also by the Rio Grande River. The “moun- species. The monotypic S. cognatus was recently separated tain islands” geographical ecology of New Mexico has from the similar S. floridanus. likely resulted in interesting evolutionary phenomena ECOLOGY: Little is known about the natural history of the in Sylvilagus; e.g., the Guadalupe Mountains of S New Manzano Mountain cottontail. Mexico are occupied by the Davis Mountains cottontail HABITAT AND DIET: The Manzano Mountain cotton- (S. robustus). The taxonomy of Sylvilagus in New Mexico tail occurs at high elevations (2,300–3,100 m) in areas could bear further scrutiny. dominated by conifer , but may also include lower-­ altitude areas with montane scrub and subalpine-­montane account author: Luis A. Ruedas . It is likely limited at lower elevations by Piñon-­ Juniper woodlands, Juniper savanna, and Plains-­Mesa Key References: Frey et al. 1997; Ruedas 1998. grassland, which are dominated by desert cottontails. REPRODUCTION AND DEVELOPMENT: There are few data on reproduction in the Manzano Mountain cotton- tail. A juvenile individual and several subadults were collected in late July. A number of small adults have also Sylvilagus cunicularius been collected in January. It is likely that there is a pro- (Waterhouse, 1848) longed development to adulthood, certainly of at least one year, possibly longer. Mexican Cottontail : IUCN Red List Classification: Data Deficient (DD) OTHER COMMON NAMES: Conejo montés, Conejo de MANAGEMENT: Although the geographic range of the monte, Conejo serrano, Conejo (Spanish) species is depicted as being continuous across the Sandia DESCRIPTION: The Mexican cottontail is the largest and Manzano Mountains, this extent of range should be rabbit in Mexico, equaling medium-­sized jackrabbits in considered only as a conservative working hypothesis. The biomass. It has coarse pelage, a massive skull, and soft . two mountain chains are separated by the Tijeras Canyon, The dorsal color is dirty yellowish or grayish, without ru- which drops to an elevation of ~ 2,100 m. This is below fous, except on the nape, and the light sub-­terminal rings the generally preferred elevational range of the species, on the hairs are uniformly pale cream-­color. The ears are although it may range that low or even lower. However, as long as the head, and their backs are thinly haired and while suitable habitat may have existed during recent gla- gray with the extreme tips and outer edges darkening ciations, the current habitat in the pass is not suitable for to black. The orbital area is clear, deep buff; the sides of the Manzano Mountain cottontail, but rather more propi- the head are dark, dingy buff, and the nape is dull rusty tious to the desert cottontail. Furthermore, the Sandia and rufous. The forelegs are similar in color to the nape, but

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the tip and expand again toward the tip. The anterior edge ± of the palatal bridge is level with the front of the anterior premolar, and its posterior edge is level with the division between the last premolar and the first molar. The jugals are proportionally light and slightly grooved with a deep pit anteriorly. The supraorbital process is light and narrow and slightly raised above the plane of the frontals; the postorbital process is usually joined to the skull posteri- orly, enclosing a narrow flattened, oval foramen. The inter-­ parietal is triangular shaped, its antero-­posterior length is nearly two-­thirds its transverse diameter, and the occipital shelf is unusually broad. The ventral border of the ramus of the lower jaw usually sits on the posterior angle and the tip when resting on a plane, leaving the middle free; the middle ventral border outline of the skull of old individu- als sometimes becomes convex, raising its tip. SIZE: Head and body 485–515 mm; Tail 54–68 mm; Hind foot 108–111 mm; Ear 60–63 mm; Weight 1,800–2,355 g PALEONTOLOGY: The few known fossils of the Mexican cottontail from the United States and Mexico represent the Pliocene and the Pleistocene. Pliocene fossils (up- per premolars) have been collected in Graham County, SE Arizona. In Mexico, a broken skull and two right den- tary bones (one without teeth) of Pleistocene age were collected at “Barranca Seca,” Veracruz. It is thought that the remains were deposited when the environment was no more tropical than it is today. Other fossil records of the Mexican cottontail date from the Late Pleistocene in La Presita, San Luis Potosí, and Guilá Naquitz, Oaxaca. Additionally, a 12,000-­year-­old anklebone (calcaneum) of the Mexican cottontail was reported from Tlapacoya, in the state of Mexico. CURRENT DISTRIBUTION: The Mexican cottontail is a Mexican endemic species that ranges widely from the Pacific Coastal Plain from Sinaloa south to Oaxaca, and encompassing the Transverse Volcanic Axis from the highlands of Michoacán to Veracruz. Marginal records Sylvilagus cunicularius. Photo courtesy Jorge Vázquez in the northern part of its distribution include the states of Sinaloa, Nayarit, Jalisco, Michoacán, Mexico, Hidalgo, duller and less rufous; the hind legs and sides of the hind Puebla, and Veracruz. The Mexican cottontail occurs from feet are duller, more rusty brownish than the forelegs; the sea level to 4,300 m in elevation. tops of the hind feet are buffy whitish or pale, dull rusty. TAXONOMY AND GEOGRAPHIC VARIATION: Three The tail is short and grayish brown above and pure white subspecies: S. c. cunicularius (the central portion of their on the under surface. distribution; Neovolcanic Axis in the states of Michoacán, The skull is large, heavy, and broad across the brain- Guerrero, and Oaxaca); S. c. insolitus (northern distribu­ case. The rostrum is heavy with a massive base, flattened tion from mid-­Sinaloa along the Pacific Coastal Plain to in the frontal region, and arched along the upper outline; south C Jalisco); S. c. pacificus (S Jalisco south to the Pa- the nasals are sharply compressed into a dorso-­lateral, cific oastalC Plain in Oaxaca). Mexican cottontails from pit-­like indentation about one-­third of the distance from the Sierra Madre of Michoacán, in the western part

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of their range, are slightly larger in both skin and skull dimensions than those from elsewhere, but there are no color differences. Considering the wide range of this species under varied climatic conditions, the amount of variation is surprisingly small. A shared common ancestor with biogeographic and morphological affinities has beenypothesized h between S. cunicularius and the Tres Marías cottontail (S. graysoni). Phylogenetic studies of Sylvilagus using data from mito- chondrial sequences of the 16S gene show that S. cunicu­ larius and the desert cottontail (S. audubonii) constitute a monophyletic group, and that the Mexican cottontail is closely related to the Omiltemi rabbit (S. insonus). ECOLOGY: The population density of the Mexican cotton- Sylvilagus cunicularius habitat. Photo courtesy Jorge Vázquez tail averages 27 individuals/km² in C Mexico, but it varies according to the rainy and dry seasons. Individual home and grassland. In C Mexico it is abundant in pine and range size is ~ 3.6/ha, and home ranges overlap between pine-­oak forests, and in W Mexico it occurs in pastures, the sexes. The Mexican cottontail is sympatric with other dry deciduous forest, and areas of disturbed vegetation, lagomorphs in portions of its range, including the volcano where it selects characterized by grasses and rabbit (Romerolagus diazi), the desert cottontail, the east- herbs. In the state of Guerrero, this cottontail is usually ern cottontail (S. floridanus), the white-­sided jackrabbit found in upland valleys in both tropical deciduous and (Lepus callotis), the black-­tailed jackrabbit (L. californicus), pine-­oak zones. In Guerrero S. c. pacificus occupies the and the antelope jackrabbit (L. alleni). Mexican cottontails narrow coastal area below 457 m asl. In the Mexican from the highlands of C Mexico have been collected in Transvolcanic Belt it inhabits forests of pine and oak-­pine abandoned of Merriam’s pocket gophers (Cratoge­ with understory of clumped grasses. It rarely has been omys merriami) and in natural rocky hollows and crevices seen in conifer forests of “oyamel” (Abies religiosa). In on and underneath the ground. Ixtacuixtla, Tlaxcala, this cottontail is found in xeric scrub Fecal pellets of the Mexican cottontail are regularly dominated by kidneywood tree (Eysenhardtia polystachya), found next to the base of grasses, forming dunghills or shadbush (Amelanchier spp.), and sumac (Rhus spp.); alli- latrines that lie at least 20 m from each other. Fecal pellets gator juniper forests; and oak woodlands (Quercus spp.). of adults are characteristic and rarely confused with those In La Malinche National Park it inhabits pine forest and from other sympatric leporids, although their form and of macroura and Peruvian feath- size approach those of jackrabbits. Fecal pellets of Mexi- ergrass (Stipa ichu), being more abundant in grasslands can cottontails are brown, composed of regurgitated plant and mixed sites of pine forest and grasslands than in pine materials, and compact (flat and round at their widest forest alone. part). Fecal pellets are 1.0–1.5 cm in diameter. Mexican Mexican cottontails from C Mexico feed on clumped cottontails eat seeds of alligator juniper (Juniperus dep­ grasses, including Peruvian feathergrass, M. macroura, and peana) and deposit them in latrines away from the mother amplissima. They select tender shoots of grasses, tree, and therefore are a dispersing agent for this tree spe- young leaves of forbs, cortexes of shrubs, and cultivated cies. plants such as oats (Avena sativa) and corn (Zea mays). The most important predators of the Mexican cotton- In C Mexico the diet of the Mexican cottontail includes tail are mammalian such as grey foxes (Urocyon 23 species, such as M. virletii, Hall’s panicgrass (Panicum cinereoargenteus), (Canis latrans), (Lynx hallii), speargrass (Piptochaetium virescens), and Baccharis rufus), and pumas (Puma concolor); some birds of prey, conferta. including barn (Tyto alba) and hawks (Buteo spp.); BEHAVIOR: Mexican cottontails are solitary and active occasionally ; and American crocodiles (Crocodylus mainly at dusk and dawn, although they may be active at acutus). night and during the day. The periods of greatest visible HABITAT AND DIET: The Mexican cottontail occurs in activity in the Mexican cottontail are 0700–0900 h and tropical, semi-­arid, temperate, open forest, dense shrub, 1400–2000 h. They engage in a wide variety of individ-

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ual behaviors: resting, grooming, foraging, coprophagy, lowing nursing they reclose the . After the nursery and scent marking. These rabbits also participate in social burrow is opened, young come to the surface on their own behaviors such as group feeding, expulsion among indi- to nurse. Litters are nursed once a day in the afternoon, viduals, and running in pairs. Strongly aggressive behav- and nursing takes place until about 12 days following par- iors are displayed by Mexican cottontails. The maximum turition. number of that can be maintained in a colony Males possess one scrotal sac, containing the testis under semi-­natural conditions without severe fighting and and epididymis. The paired testes of adults are scrotal deaths is three to four females and two males in an enclo- throughout the year. The penis is cylindrical and covered sure of 530 m2 and two females and one male in a 108 m2 by the preputial skin structure. The male reproductive enclosure. system possesses four accessory sexual glands: ampulla, PHYSIOLOGY AND GENETICS: Diploid prostate, bulbourethral, and preputial. Females have a number = 42. The FN and morphology of the sexual lower level of testosterone than males in any condition of chromosome are unknown since only females have been reproduction. In males, testosterone levels are low four examined. months after the start of the breeding season, and then White blood cells of Mexican cottontail contain lym- soar before initiating breeding. phocytes (35% to 60%), monocytes (1%), eosinophils PARASITES AND DISEASES: Ectoparasites on Mexican (0—2%), basophils (0% to 2%), and neutrophils (37% to cottontails include fleas (Cediopsylla inaequalis interrupta, 62%). Euhoplopsyllus glacialis affinis, and Pulex irritans). Gut REPRODUCTION AND DEVELOPMENT: The Mexican cot- parasites identified in the Mexican cottontail include pro- tontail reproduces throughout the year. However, repro- tozoans (Eimeria), cestodes (Cittotaenia and Taenia), and ductive females have been recorded in greater numbers nematodes (Strongyloides, Passalurus, Trichostrongylus, and from March to October, even while in the same popula- Heterakis). tion males with descended testes and juveniles could be CONSERVATION STATUS: found throughout the year. The onset of breeding may IUCN Red List Classification: Least Concern (LC) be associated with an increase in day length and tem- MANAGEMENT: Mexican cottontails are hunted for food perature, whereas the birth of young is associated with and sport. They are also killed for being considered com- increased rainfall, hence with the availability of succulent petitors with livestock. This killing has caused a decline vegetation. Litter size ranges from 1 to 6, and the gesta- of some populations of the Mexican cottontail. Legal tion period is 28–31 days. Each female may reproduce hunting does not account for the reproduction season or several times a year. age structure of Mexican cottontails, which additionally Nursery burrows are extremely difficult to find be- contributes to the decrease of Mexican cottontail popu- cause mothers close the entrance with soil and cover it lations. Until a few years ago the Mexican cottontail was with leaves and grass like other lagomorphs. Burrows a common mammal near the coast in Jalisco, however, dug under semi-­natural conditions consist of a single at present it is rare in the area; local residents claim the tunnel with a simple entrance ranging from 15 to 53 cm decline is due to hunting. Similarly, the Mexican cotton- in length. These tend to be dug beneath grass tussocks tail is no longer abundant in the state of Morelos after in- (M. macroura), shrubs such as willow ragwort (Senecio tensive hunting by residents. In some parts of their range, salignus), or herbs including Eupatorium pazcuarense and fires and excessive logging have reduced drastically the pride of the mountain (Penstemon roseus). The nursery rabbit’s habitat (grasslands and pine forest). In C and W burrow ends in a small spherical chamber containing a Mexico the Mexican cottontail faces the threat of habitat nest 17 cm (range 13–22 cm) beneath the surface. Nests fragmentation. Additionally, it has been suggested that consist of dry grass, fragments of woody plants, pine deforestation, expansion of cattle pastures and agricul- needles and hay, fur pulled from the mother’s body, and ture, and introduced feral fauna and grasses are negatively fecal pellets probably also from the mother. affecting conservation of this species. Births occur at the nursery burrow entrance, and then newborns crawl into the nest. Young at birth are covered account authors: Consuelo Lorenzo, Jorge Vázquez, with fine hair, and their eyes are closed. Mothers open the Luisa Rodríguez-­Martínez, Amando Bautista, Antonio nursery burrow entrance to nurse their young, and fol- García-­Méndez, and Fernando A. Cervantes

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