MaMMals of China Andrew T. Smith and Yan Xie Editors Robert S. Hoffmann Darrin Lunde John MacKinnon Don E. Wilson W. Chris Wozencraft Contributing Authors Federico Gemma Illustrator Professor Wang Sung Honorary Editor PRinCETon UnivERSiTY PRESS PRinCETon AnD oXFoRD Wildlife of Australia, by iain Campbell and Sam Woods Wildlife of East Africa, by Martin B. Withers and David Hosking Wildlife of the Galápagos, by Julian Fitter, Daniel Fitter, and David Hosking Wildlife of Southern Africa, by Martin B. Withers and David Hosking Coral Reef Fishes: Indo-Pacific and Caribbean, Revised Edition, by Ewald Lieske and Robert Myers A Field Guide to the Birds of New Zealand, by Julian Fitter The Kingdon Pocket Guide to African Mammals, by Jonathan Kingdon Mammals of China, edited by Andrew T. Smith and Yan Xie Reptiles and Amphibians of East Africa, by Stephen Spawls, Kim M. Howell, and Robert C. Drewes Copyright © 2013 by Princeton University Press Published by Princeton University Press, 41 William Street, Princeton, new Jersey 08540 in the United Kingdom: Princeton University Press, 6 oxford Street, Woodstock, oxfordshire oX20 1TW press.princeton.edu Cover illustrations © Federico Gemma All Rights Reserved iSBn (pbk.) 978-0-691-15427-5 Library of Congress Control number 2013930079 British Library Cataloging-in-Publication Data is available This book has been composed in Minion Pro Printed on acid-free paper Printed in China 1 3 5 7 9 10 8 6 4 2 Contents List of Figures 5 List of Maps 6 List of Habitat Images 6 Preface 7 Contributors 7 Introduction 8 China’s Geography and Mammal Biogeography 9 Mammal Conservation 12 China’s Protected-Area System 14 How to Use This Book 15 Acknowledgments 18 Class MaMMalia—The Mammals 35 oRDER PRoBosCiDEa 36 Family Elephantidae—Elephants 36 oRDER siREnia 37 Family Dugongidae—Dugongs 37 oRDER sCanDEnTia 38 Family Tupaiidae—Tree Shrews 38 oRDER PRiMaTEs 39 Family Lorisidae—Lorises 39 Family Cercopithecidae 41 Subfamily Cercopithecinae—old World Monkeys 42 Subfamily Colobinae—Leaf-eating Monkeys 46 Family Hylobatidae—Gibbons and Siamangs 52 oRDER RoDEnTia 58 Family Sciuridae 58 Subfamily Ratufinae—Giant Squirrels 59 Subfamily Sciurinae—True Squirrels 60 Subfamily Callosciurinae—oriental Tree Squirrels 69 Subfamily Xerinae—Afro-Asian Ground Squirrels 76 Family Gliridae—Dormice 84 Family Castoridae—Beavers 85 Family Dipodidae 87 Subfamily Allactaginae—Four- and Five-toed Jerboas, Fat-tailed Jerboas 87 Subfamily Cardiocraniinae—Dwarf Jerboas 91 Subfamily Dipodinae—Three-toed Jerboas 93 Subfamily Euchoreutinae—Long-eared Jerboa 96 Subfamily Sicistinae—Birch Mice 96 Subfamily Zapodinae—Jumping Mice 98 Family Platacanthomyidae—Spiny Dormice 99 Family Spalacidae 100 Subfamily Myospalacinae—Zokors 100 Subfamily Rhizomyinae—Bamboo Rats 103 4 — Contents Family Cricetidae 105 Subfamily Arvicolinae—voles, Lemmings, and Water voles 106 Subfamily Cricetinae—Hamsters 129 Family Muridae 138 Subfamily Gerbillinae—Gerbils 138 Subfamily Murinae—Murine Rodents 143 Family Hystricidae—old World Porcupines 164 oRDER laGoMoRPha 166 Family ochotonidae—Pikas 166 Family Leporidae—Rabbits and Hares 178 oRDER ERinaCEoMoRPha 184 Family Erinaceidae 184 Subfamily Erinaceinae—Hedgehogs 184 Subfamily Galericinae—Gymnures 187 oRDER soRiCoMoRPha 189 Family Soricidae 189 Subfamily Crocidurinae—Crocidurine Shrews 189 Subfamily Soricinae—Red-toothed Shrews 195 Family Talpidae 211 Subfamily Scalopinae—American Moles 211 Subfamily Talpinae—True Moles 212 Subfamily Uropsilinae—Shrew Moles 216 oRDER ChiRoPTERa 217 Family Pteropodidae—old World Fruit Bats 217 Family Rhinolophidae—Horseshoe Bats 224 Family Hipposideridae—old World Leaf-nosed Bats 233 Family Megadermatidae—False vampire Bats 238 Family Emballonuridae—Sheath-tailed Bats 239 Family Molossidae—Free-tailed Bats 240 Family vespertilionidae 242 Subfamily vespertilioninae—vesper Bats 243 Subfamily Myotinae—Little Brown Bats 260 Subfamily Miniopterinae—Long-fingered Bats 269 Subfamily Murininae—Hairy-winged and Tube-nosed Bats 270 Subfamily Kerivoulinae—Woolly Bats 274 oRDER PholiDoTa 275 Family Manidae—Pangolins 275 oRDER CaRniVoRa 277 Family Felidae 278 Subfamily Felinae—Purring Cats 278 Subfamily Pantherinae—Roaring Cats 287 Family viverridae 292 Subfamily Hemigalinae—Hemigaline Civets 293 Subfamily Paradoxurinae—Palm Civets 294 Subfamily Prionodontinae—Linsangs 297 Subfamily viverrinae—Civet Cats 298 Family Herpestidae—Mongooses 301 Family Canidae—Canids 303 List of figures — 5 Family Ursidae—Bears 308 Family otariidae—Eared Seals 313 Family Phocidae—True Seals 315 Family Mustelidae 318 Subfamily Lutrinae—otters 318 Subfamily Mustelinae—Weasels, Wolverine, Martens, and Badgers 321 Family Ailuridae—Red Panda 333 oRDER PERissoDaCTYla 334 Family Equidae—Horses, Zebras, and Asses 334 oRDER aRTioDaCTYla 337 Family Suidae—Pigs 338 Family Camelidae—Camels 339 Family Tragulidae—Chevrotains and Mouse-deer 340 Family Moschidae—Musk Deer 341 Family Cervidae 345 Subfamily Capreolinae—Deer, Moose, and Reindeer 345 Subfamily Cervinae—Deer and Muntjacs 349 Subfamily Hydropotinae—Water Deer 359 Family Bovidae 360 Subfamily Antilopinae—Gazelles and Saiga 360 Subfamily Bovinae—Cattle and Spiral-horned Antelopes 365 Subfamily Caprinae—Goats and Sheep 367 oRDER CETaCEa 377 Family Delphinidae—Dolphins 378 Family Phocoenidae—Porpoises 379 Family iniidae—River Dolphins 380 APPENDIXES 381 i. Cetaceans Found off the Coast of China 381 ii. Hypothetical Mammals 382 iii. introduced Alien, Feral, or Free-ranging Domestic Mammals 383 GLOSSARY 384 FURTHER READING 387 Index to Scientific Names 388 Index to Common Names 391 List of Figures Figure 1. Standard external mammal measurements 16 Figure 2. Standard external bat measurements 16 Figure 3. Feet of the Bengal Slow Loris 40 Figure 4. Feet of the Eurasian Beaver 86 Figure 5. Feet of five-toed and three-toed jerboas 87 Figure 6. Feet of dwarf hamsters Cricetulus and Phodopus 131 Figure 7. Wing of flying fox (Pteropus) 221 Figure 8. Foot pad of bamboo bat (Tylonycteris) 257 Figure 9. Tails of indian and Chinese pangolins 276 6 — List of Maps Figure 10. Front paws of Asian Black and Brown bears 311 Figure 11. Tails of Sable, Beech Marten, and Yellow-Throated Marten 324 Figure 12. Skull and horn profiles of Goitered, Mongolian, Tibetan, and Przewalski’s gazelles 361 List of Maps Front Cover (inside). Administrative Provinces and Autonomous Regions of China Map 1. Major Rivers and Landforms of China 19 Map 2. Biogeographic Regions of China 20 Map 3. China’s Physical Geographic Regions and Major Biotic Divisions 21 Map 4. Mammal Biogeographic Divisions and Regions of China 21 Back Cover (inside). Protected Areas of China List of Habitat Images Habitat 1. Tian Shan Grassland and Forest; Xinjiang 22 Habitat 2. Tamarisk Riparian Habitat; Xinjiang 22 Habitat 3. Bogda Feng from Turpan Basin; Xinjiang 23 Habitat 4. Gobi Desert Habitat; Gansu 23 Habitat 5. nei Mongol Grasslands; nei Mongol 24 Habitat 6. Tibetan Plateau Grasslands; Qinghai 24 Habitat 7. Tibetan Plateau Wetlands; Qinghai 25 Habitat 8. Upper Yangtze River; Qinghai 25 Habitat 9. Juniper Forest, Tibetan Plateau; Qinghai 26 Habitat 10. Bamboo Forest, Dujiangyan Longqi-Hongkou Reserve; Sichuan 26 Habitat 11. Coniferous Forest, Juizhaigou; Sichuan 27 Habitat 12. Forest Understory, Dahonggou Creek; Sichuan 27 Habitat 13. Wolong nature Reserve; Sichuan 28 Habitat 14. Karst Forest; Maolan Protected Area; Guizhou 29 Habitat 15. Xishuanbanna national nature Reserve; Yunnan 29 Habitat 16. Mengla Forest; Yunnan 30 Habitat 17. Damingshan nature Reserve; Guangxi 30 Habitat 18. Hainan Forest; Hainan 31 Habitat 19. Mangrove Habitat; Hainan 31 Habitat 20. Mixed Forest, Changbaishan; Jilin 32 Habitat 21. Mixed Conifer Forest, Changbaishan; Jilin 32 Habitat 22. northern Tundra, Changbaishan; Jilin 33 Habitat 23. northern Coniferous Forest, Changbaishan; Jilin 33 Habitat 24. Grassland and Forest Edge, Changbaishan; Jilin 34 Habitat 25. Forest Understory, Mudangjiang; Heilongjiang 34 Contributors — 7 Preface This Princeton Pocket Guide to the mammals of China is designed to give ready access to those encountering the diverse mammal fauna of China. This book is an outgrowth of the earlier comprehensive A Guide to the Mammals of China (Smith and Xie 2008). The full guide contains additional information on the identification of Chinese mammals, including comprehensive keys and descriptions of skulls and osteological features, as well as thorough reviews of systematic controversies concerning Chinese mammals and a history of Chinese mammalogy. it also contains a lengthy bibliography to lead readers to pursue additional background information concerning Chinese mammals. Those interested in these details of Chinese mammals are encouraged to consult this longer treatment. Here, however, we have presented material that can serve directly to identify mammals encountered in the field—concentrating on range maps and descriptions of the external appearance of species and their natural history. increasingly, people are becoming interested in ecotourism and natural history. We hope that this book engenders enthusiasm for the study of Chinese mammals, especially in their natural and wild state. Andrew Smith, Tempe, and Xie Yan, Beijing Contributors FEDERiCo GEMMA Species Plates viale Marconi 19 Rome 00146 italy [email protected] www.federicogemma.it RoBERT S. HoFFMAnn* Sciuridae, Cricetinae, Gerbillinae, Erinaceomorpha, Soricomorpha Division of Mammals Department of vertebrate Zoology national Museum of natural History Smithsonian institution—MRC-108 Washington, DC 20013 USA DARRin LUnDE Arvicolinae, Murinae, Erinaceomorpha, Soricomorpha Division of Mammals Department of vertebrate Zoology national Museum of natural History Smithsonian institution—MRC-108 Washington, DC 20013 USA [email protected] JoHn R. MACKinnon Proboscidea, Primates, Artiodactyla 11 Leycroft Close Canterbury CT2 7LD United Kingdom [email protected] 8 — introduCtion AnDREW T. SMiTH Introduction, Proboscidea, Sirenia, Scandentia, Sciuridae, Gliridae, Castoridae, Dipodidae, Platacanthomyidae, Spalacidae, Cricetinae, Gerbillinae, Hystricidae, Lagomorpha, Pholidota, Cetacea, Maps, Appendixes School of Life Sciences Box 874501 Arizona State University Tempe, AZ 85287-4501 USA [email protected] WAnG SUnG Honorary Editor institute of Zoology Chinese Academy of Sciences 1-5 Beichenxilu, Chaoyang District Beijing 100101 People’s Republic of China [email protected] Don E. WiLSon Chiroptera Division of Mammals Department of vertebrate Zoology national Museum of natural History Smithsonian institution—MRC-108 Washington, DC 20013 USA [email protected] W. CHRiS WoZEnCRAFT* Carnivora Division of natural Sciences Bethel College 1001 West McKinley Avenue Mishawaka, in 46545 USA XiE YAn Introduction; Maps institute of Zoology Chinese Academy of Sciences 1-5 Beichenxilu, Chaoyang District Beijing 100101 People’s Republic of China [email protected] *deceased Introduction China is a magnificent country and one of the most diverse on Earth. its size ranks fourth among the world’s nations (9,596,960 km2), and it is home to over 1.3 billion people. The topography of China ranges from the highest elevation on Earth (Mount Everest, or Chomolungma; 8,850 m) to one of the lowest (Turpan Basin; 154 m below sea level). Chinese environments include some of Earth’s introduCtion — 9 most extensive and driest deserts (the Taklimakan and Gobi) and its highest pla- teau (the Tibetan Plateau or “Roof of the World”). Habitats range from tropical to boreal forest, and from extensive grasslands to desert (see the habitat images, map 1, and map 2). This wide variety of habitats has contributed greatly to the richness of China’s mammal fauna. Additionally, the geographic location of China, at the suture zone between the Palaearctic and indo-Malayan biogeo- graphic regions, further contributes to the country’s mammal diversity. overall, more than 10 percent of the world’s species of mammals live in China (556/5,416); an additional 29 species of cetaceans live in offshore waters (see Appendix i). Almost 20 percent (106/556) of China’s mammals are endemic, and one of these is among the most recognizable of the world’s mammals, the Giant Panda. China is considered a “megadiversity” country and has the third highest diversity of mammals among all countries (following Brazil and indonesia). China’s Geography and Mammal Biogeography There have been many attempts to describe China’s diverse landscape. Conven- tionally, China has been divided into three major physical geographic regions: the Tibetan (Qinghai-Xizang) Plateau, northwest arid China, and eastern mon- soon China (map 3). The Tibetan Plateau is one of the highest and most remote landscapes on Earth. The plateau averages between 3,000 and 5,000 m in elevation and encom- passes roughly a quarter of China. The word “plateau” is a misnomer, as this area is crisscrossed by numerous impressive mountain chains, such as the Anyemaqin Shan, Bayan Har Shan, and Tanggula Shan, and many smaller spur ranges. nev- ertheless, approximately 70 percent of the plateau is composed of alpine meadow or semisteppe vegetation. The Qaidam Basin, an interesting area of tectonic col- lapse, is found at the northern extreme of the plateau at an elevation of only 2,600 m. The arid northwest encompasses about 30 percent of China and represents an eastern extension of the great Eurasian deserts and grasslands. one of the world’s most desolate deserts, the Taklimakan (translation: “those who go in do not come out alive”) lies north of the Tibetan Plateau and the Kunlun Mountains. The cooler Dzungarian Basin, China’s second-largest desert, lies in the far north- west. various smaller deserts extend to the east, increasingly interspersed with semidesert and temperate steppe grasslands. Finally, the rocky Gobi Desert occupies the northern part of China and extends into Mongolia. Two of Asia’s major mountain ranges break up this barren expanse in the northwest: the Tian Shan and the Altai. one can stand below sea level in Turpan Basin and clearly see the snow-capped top of Bogda Feng (in a spur of the Tian Shan) at 5,445 m. Eastern monsoon China comprises about 45 percent of the country but is home to roughly 95 percent of China’s human population. This land is crossed by major rivers that originate on the Tibetan Plateau, most notably the Huang He (Yellow River), Yangtze and Mekong. Almost all of the arable land has been con- verted to agriculture, and much of the original forest habitat has been destroyed. Most of this landscape is low in elevation and consists of broad alluvial valleys, coastal plains, and modest ancient mountain ranges. The south is seasonably humid, and the plains are punctuated by dramatic limestone pillars. The climate