Threatened and Recently Extinct Vertebrates of the World: A Biogeographic Approach 1108495869, 9781108495868

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Threatened and Recently Extinct Vertebrates of the World A Biogeographic Approach Habitat loss and degradation are currently the main anthropogenic drivers of species extinctions. The main driver of this worldwide is agriculture, with urban sprawl, logging, mining, and some fishing practices close behind. The physical loss of habitat, such as deforestation for land development and the burning of fossil fuels, are examples of this. The root cause is human overpopulation. This unique volume provides, for the very first time, a comprehensive overview of all threatened and recently extinct mammals, birds, reptiles, amphibians, and fishes within the context of their locations and habitats. The approach takes a systematic examination of each biogeographic realm and region of the world, both terrestrial and marine, but with a particular emphasis on geographic features such as mountains, islands, and coral reefs. It reveals patterns useful in biodiversity conservation, helps to put it all into perspective, and ultimately serves as both a baseline from which to compare subsequent developments as well as a standardization of the way threatened species are studied. Matthew Richardson is a Canadian author known for such awardwinning books as Lemurs of Madagascar (3rd edition, Conservation International, 2010). He has worked alongside some of the most high-profile conservationists in the world, and served as an active member of both the IUCN/SSC Primate Specialist Group and Conservation Planning Specialist Group. Passionate about both human and natural history, he has drawn extensively from these experiences to create what will surely become the foundational text on endangered and extinct species for decades to come.

Published online by Cambridge University Press

Published online by Cambridge University Press

Threatened and Recently Extinct Vertebrates of the World A Biogeographic Approach Matthew Richardson

Published online by Cambridge University Press

Shaftesbury Road, Cambridge CB2 8EA, United Kingdom One Liberty Plaza, 20th Floor, New York, NY 10006, USA 477 Williamstown Road, Port Melbourne, VIC 3207, Australia 314–321, 3rd Floor, Plot 3, Splendor Forum, Jasola District Centre, New Delhi – 110025, India 103 Penang Road, #05-06/07, Visioncrest Commercial, Singapore 238467 Cambridge University Press is part of Cambridge University Press & Assessment, a department of the University of Cambridge. We share the University’s mission to contribute to society through the pursuit of education, learning and research at the highest international levels of excellence. www.cambridge.org Information on this title: www.cambridge.org/9781108495868 DOI: 10.1017/9781108863308 © Matthew Richardson 2023 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press & Assessment. First published 2023 Printed in the United Kingdom by TJ Books Limited, Padstow Cornwall A catalogue record for this publication is available from the British Library. Library of Congress Cataloging-in-Publication Data Names: Richardson, Matt, 1966– author. Title: Threatened and recently-extinct vertebrates of the world : a biogeographic approach / Matthew Richardson. Description: Cambridge, United Kingdom ; New York, NY : Cambridge University Press, 2023. | Includes bibliographical references and index. Identifiers: LCCN 2022023457 (print) | LCCN 2022023458 (ebook) | ISBN 9781108495868 (hardback) | ISBN 9781108863308 (epub) Subjects: LCSH: Rare vertebrates. | Extinct vertebrates. | Habitat conservation. Classification: LCC QL88 .R525 2023 (print) | LCC QL88 (ebook) | DDC 599–dc23/eng/20220729 LC record available at https://lccn.loc.gov/2022023457 LC ebook record available at https://lccn.loc.gov/2022023458 ISBN 978-1-108-49586-8 Hardback Cambridge University Press & Assessment has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

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This book is dedicated to my father Charles Cruse Richardson 1928–2015 my mother Dorothy May Richardson née Lihou 1932–2017 and to my dearest friend Douglas Arthur Sweetman 1964–2019

Published online by Cambridge University Press

Published online by Cambridge University Press

Contents Foreword Stuart L. Pimm xix Preface xxi Acknowledgements xxviii 1

The Arctic Realm 1 The Polar Ice Cap 1 Tundra 1 Coasts and Satellite Islands 3 The Arctic Archipelago 4 Greenland 4 Svalbard 4 Novaya Zemlya 4 Wrangel Island 4 Balance for the Arctic Realm 4

2

The Palearctic Realm 6 The Eurasian Region 11 Mountains and Highlands 13

The Alps 15 Mountains of Central Asia 15 The Caucasus Mountains 16 The Armenian Plateau 17 The Pontic Mountains 17 The Taurus Mountains 17 The Apennine Mountains 17 The Pyrenees Mountains 17 The Cantabrian Mountains 17 The Carpathian Mountains 17 Miscellaneous Mountains and Highlands

18

Lowland Boreal Forests 18 Lowland Broadleaf and Mixed Forests 19 Lowland Mediterranean Forests, Woodlands, and Scrub 19 Eurasian Steppe 20 Western Steppe 20 Central Steppe 21 Eastern Steppe 21

Deserts and Semi-Deserts The Gobi Desert 23 Miscellaneous Deserts

22

23

Isolated Caves, Springs and Pools The Sultan Sazligi Marshes

23

Lakes, Rivers, and Marshes The Caspian Sea 28 The Aral Sea 28

23

Lake Urmia 29 Lake Baikal 29 Lake Saimaa 29 Lake Geneva 29 The Prespa Lakes 29 Lake Ohrid 30 Lake Van 30 Lake Neuchâtel 30 Lake Beysehir 30 Lake Egirdir 30 Lake Burdur 30 Lake Tuz 30 Lake Vistonis 31 Lake Trichonida 31 Lake Skadar 31 Lake Ammer 31 Lake Ladoga 31 The Tagus River 31 The Guadiana River 31 The Guadalquiver River 31 The Danube River 31 The Rhine River 32 The Rhône River 32 The Neretva River 32 The Pinios River 32 The Krka River 32 The Cetina River 33 The Ceyhan River 33 The Buyuk Menderes River 33 The Upper Euphrates River 33 The Upper Tigris River 33 The Chornaya River 34 The Amur River 34 Miscellaneous Lakes, Rivers, and Marshes

Coasts and Satellite Islands 23

The Azores 36 The British Isles 37 Sakhalin 37 Bering Island 38 Sicily 38 Corsica and Sardinia 38 The Aegean Islands 39

36

34

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Contents The Balearic Islands 39 The Aeolian Islands 40 Miscellaneous Islands 40

Mountains and Highlands The Atlas Mountains 76 Levantine Mountains 76 The Al Hajar Mountains 76 The Sarawat Mountains 77 The Kopet Dag Mountains 77 The Alborz Mountains 77 The Zagros Mountains 77 The Hindu Kush 77 The Marrah Mountains 77 The Nur Mountains 78

Balance for the Eurasian Region 40 The Sino-Himalayan Region 42 Mountains and Highlands 43 The Tibetan Plateau 47 The Himalayan Mountains 48 The Purvanchal Range 50 The Loess Plateau 51 The Qinling Mountains 51 The Wuling Mountains 51 The Qionglai Mountains 51 The Min Mountains 51 The Daba Mountains 52 The Tianmu Mountains 52 The Dabie Mountains 52 The Luoxiao Mountains 52 The Huangshan Mountains 52 The Gaoligong Mountains 52 Mount Emei 52 The Hengduan Mountains 52 The Yunnan-Guizhou Plateau 52 The Dayao Mountains 53 The Miaoling Mountains 53 The Hoang Lien Son Range 53 Miscellaneous Mountains and Highlands

Lowland Mediterranean Forests, Woodlands, and Scrub 78 Deserts and Semi-Deserts 78 The Sahara Desert 79 The Arabian Desert 81 The Levantine Desert 83 The Syrian Desert 83 The Dasht-e Kavir Desert 83 Miscellaneous Deserts 83

Isolated Caves, Springs, and Pools The Haditha Aquifer System 84 Wadi Hadhramaut 84

Lakes, Rivers, and Marshes

Coasts and Satellite Islands The Japanese Archipelago The Ryuku Islands 67 Taiwan 69 Hong Kong 70 Cat Ba Island 70

65

65

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84

Coasts and Satellite Islands The Madeira Islands 87 The Canary Islands 87 Cyprus 89 The Socotra Archipelago 89 Miscellaneous Islands 89

3

The Afrotropical Realm 92 The Guineo-Congolian Region Mountains and Highlands 95

89

93

The Albertine Rift Mountains 96 The Cameroon Volcanic Line 98 The Guinea Highlands 101 Miscellaneous Mountains and Highlands

64

85

86

Balance for the Saharo-Arabian Region Balance for the Palearctic Realm 91

Balance for the Sino-Himalayan Region The Saharo-Arabian Region 71

viii

83

The Dead Sea 84 Lake Tiberias 85 The Oum Erbiah River 85 The Lower Tigris/Lower Euphrates River Drainage The Lower Indus River 85 The Orontes River 85 The Jordan River Drainage 86 Miscellaneous Lakes, Rivers, and Marshes 86

Lowland Subtropical Moist Forests 54 Lowland Temperate Deciduous Forests 55 Isolated Caves, Springs, and Pools 55 Lakes, Rivers, and Marshes 55 Lake Dianchi 57 Lake Fuxian 58 Lake Qilu 58 Lake Yangzong 58 Lake Erhai 59 The Upper Mekong River 59 The Upper Irrawaddy River 59 The Upper Salween River 59 The Upper Ganges River 60 The Upper Brahmaputra River 60 The Yangtze River 61 The Pearl River 62 The Yellow River 63 The Red River System 63 Miscellaneous Lakes, Rivers, and Marshes

75

102

Lowland Tropical Rainforests and Swamp Forests Lowland Guinean Rainforests and Swamp Forests 102 Lowland Congolian Rainforests and Swamp Forests 106

Forest–Savanna Mosaic 70

108

Guinean Forest–Savanna Mosaic 108 Congolian Forest–Savanna Mosaic 108

102

Contents

Lakes, Rivers, and Marshes

108

The Upemba Wetlands 112 Lake Tanganyika 113 Lake Albert 113 Lake Rukwa 113 Lake George 114 Lake Barombi-Mbo 114 Lake Bermin 114 Lake Ejagham 114 The Congo River Drainage 114 The Kagera River 119 The Volta River Drainage 120 The Niger River 120 The Ogooué River 121 The Konkouré River 121 The Pra River 122 The Cavalla River 122 The Cross River 122 The Nyong River 123 The Sanaga River 123 The Kouilou-Niari River 123 The Campo River 123 The Cuanza River 123 The Cuvo River 124 The Nyanga River 124 The Chari River 124 The Saint Paul River 124 The Mano River 124 Miscellaneous Lakes, Rivers, and Marshes

Coasts and Satellite Islands Bioko 126 São Tomé 127 Príncipe 128 Annobón 128 Miscellaneous Islands

125

Lowland Mediterranean Forest (Fynbos) 145 Lowland Tropical and Subtropical Savannas 146 Lowland Tropical Savannas 147 Lowland Subtropical Grasslands and Savannas

Steppe The Sahel

Lakes, Rivers, and Marshes

153

Lake Afrera 154 Lake Magadi 154 Lake Natron 154 Lake Victoria 155 Lake Malawi 156 The Upper Nile River Drainage 158 The Omo River 159 The Shabelle–Jubba River Drainage 159 The Athi-Galana River Drainage 159 The Tana River 159 The Pangani River 160 The Malagarasi River 160 The Wami River 160 The Ruvu River 160 The Rufiji River 160 The Zambezi River Drainage 161 The Okavango River 161 The Komati River 161 The Limpopo River 161 The Olifants River 161 Miscellaneous Lakes, Rivers, and Marshes 161

124

Coasts and Satellite Islands

144

The Zanzibar Archipelago 163 The Cape Verde Islands 164 Ascension 165 Saint Helena 165 Tristan da Cunha 165

128

162

Balance for the African Region 166 Balance for the Afrotropical Realm 168 4

The Madagascan Realm 170 Madagascar 170 Mountains and Highlands 177 The Northern Highlands 177 The Central Highlands 179

Lowland Rainforests 143

182

Littoral Forests and Scrub

184

Lowland Dry Deciduous Forests 144

Coastal Moist Forests

144

East African Coastal Forests 144 Southeast African Coastal Forests 145

150

152

Karoo 152 The Danakil Desert 152 The Namib Desert 152 The Kaokoveld Desert 153

128

The Ethiopian Highlands 135 The Northern Somalia Highlands 136 The Eastern Rift Mountains 136 The Eastern Arc Mountains 138 The Southern Highlands 141 The Mulanje Massif 141 The Eastern Highlands 142 The Central Plateau 142 The Cape Fold Mountains 143 The Angolan Highlands 143 Miscellaneous Mountains and Highlands The Tana River Forest

151

Deserts and Semi-Deserts

Balance for the Guineo-Congolian Region The African Region 129 Mountains and Highlands 133

Gallery Forests

151

Tsingy Forest

186

184

Semi-Deserts and Arid Shrublands Succulent Spiny Woodlands Arid Spiny Bush 188

187

187

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Contents

Lakes, Rivers, and Marshes

188

Isolated Caves, Springs and Wells 224

Lake Alaotra 190 The Mananara du Nord River 191 The Lokoho River 191 The Sofia River 191 The Betsiboka River 191 The Onilahy River 192

Coasts and Satellite Islands Nosy Hara 193 Nosy Mangabe 193 Nosy Boraha 193 Nosy Be and Nosy Komba 193 The Glorioso Islands 193

The Comoros Islands Grand Comore 194 Mount Karthala

The Pang Mapha Karst Formation 225

Lakes, Rivers, and Marshes

192

193

194

Anjouan 194 Mayotte 194 Mohéli 194 The Seychelles 194 Mahé 196 Praslin 196 Silhouette 196 Marianne 196 The Aldabra Islands 196 Aldabra Atoll 196 The Amirante Islands

196

The Mascarene Islands Mauritius 197

196

Round Island 199 Serpent Island 199 Gunner’s Quoin, Pigeon Rock, Ile aux Vacoas, and Flat Island

Réunion 199 Rodrigues 200 Balance for the Madagascan Realm 5

201

The Indo-Malaysian Realm 203 The Oriental Region 206 Mountains and Highlands 213

The Western Ghats 214 The Eastern Ghats 217 The Shillong Plateau 218 The Shan Hills 218 The Arakan Mountains 218 The Pegu Range 219 The Cardamom Mountains 219 The Annamite Mountains 219 Miscellaneous Mountains and Highlands

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199

Sri Lanka 242 Lakes, Rivers, and Marshes 245 The Maldives 245 The Andaman Islands 246 The Nicobar Islands 246 Hainan 246 Islands of Rach Gia Bay 247 Miscellaneous Islands 247

242

247

The Tenasserim Range 251 Lowland Tropical Rainforests and Swamp Forests 252 Lakes, Rivers, and Marshes 253 Coasts and Satellite Islands 254

Sumatra 221

Seasonally Flooded and Riverine Grasslands Dry Grasslands 224

224

Coasts and Satellite Islands

Balance for the Oriental Region The Sundaic Region 249 The Malay Peninsula 251

Lowland Tropical Moist Forests and Swamp Forests 221 Lowland Tropical Dry Forests and Shrublands Tropical Savannas and Grasslands 222

Deserts and Xeric Shrublands

225

Lake Inlé 231 Lake Indawgyi 232 The Kaveri River 232 The Periyar River 232 The Panniyar River 232 The Chalakudy River 233 The Sharavati River 233 The Krishna River 233 The Seetha River 233 The Tunga River 233 The Godavari River 233 The Namdapha River 233 The Lower Ganges River 233 The Lower Brahmaputra River 234 The Surma–Meghna River Drainage 234 The Lower Irrawaddy River 235 The Lower Salween River 235 The Great Tenasserim River 236 The Chao Phraya River Drainage 236 The Mae Klong River 237 The Lower Mekong River 237 The Thu Bon River 240 The Phong Nha River 240 The Lam River 240 The Ma River 240 Miscellaneous Lakes, Rivers, and Marshes 241

223

222

254

The Barisan Mountains 255 Lowland Rainforests and Swamp Forests Lakes, Rivers, and Marshes 257 Weh 258 Simeulue 258 Lasia 258 Nias 258 The Mentawai Islands 258 Enggano 259

257

Contents Bangka 259 Belitung 259

259

The Natuna Islands Borneo 259

The Central Highlands 261 Lowland Rainforests and Swamp Forests Isolated Caves, Springs, and Pools 263 Lakes, Rivers, and Marshes 263 The Karimata Islands 264 Miang 264 Maratua 264 Bunyu 264

Java

Buton 287 The Tukangbesi Islands 287 The Selayar Islands 287 Kalaotoa 287

The Moluccas Archipelago 262

Morotai 288 Halmahera 288 The Sula Islands 288 Obira 288 Buru 288 Seram 289 The Banda Islands 289 The Kai Islands 289 The Barat Daya Islands 289 The Tanimbar Islands 289 The Aru Islands 289

265

The Javan Volcanic Chain 266 Lowland Rainforests 267 Lakes, Rivers, and Marshes 267 Bali 267 Bawean 267 The Karimunjawa Islands 268 The Kangean Islands 268 The Masalembu Islands 268

The Lesser Sunda Islands Lombok 290 Sumbawa 290 Flores 290 The Solor Archipelago 291 The Alor Archipelago 291 Sumba 291 Timor 291

Christmas Island 268 The Cocos (Keeling) Islands 268 Balance for the Sundaic Region 268 The Wallacean Region 270 The Philippines 270 The Batanes Islands 274 Luzon 274 The Romblon Islands 276 Mindoro 276 Samar 277 Leyte 277 Panay 277 Palawan 277 Negros 278 Ticao 278 Cebu 278 Bohol 279 Siquijor 279 Mindanao 279 The Sulu Archipelago 280

The Sangir Islands Great Sangir Island Siau 282

The Talaud Islands Karakelang 282 Salibabu 282 Miangas 282

Sulawesi

281

281

282

282

Mountains 284 Lowland Rainforests 285 Lakes, Rivers, and Marshes 285 The Togean Islands 286 The Banggai Archipelago 286

287

290

Balance for the Wallacean Region 291 Balance for the Indo-Malaysian Realm 292 6

The Papua-Melanesian Realm New Guinea 295

294

The New Guinea Highlands 296 Lowland Rainforests and Swamp Forests Trans-Fly Savanna and Grasslands 302 Lakes, Rivers, and Marshes 302 Coasts and Satellite Islands 305

The Bismarck Archipelago New Britain 308 New Ireland 308 The Admiralty Islands 308 The St. Matthias Islands 308

The Solomon Islands

307

309

Bougainville 309 Guadalcanal 310 Malaita 310 Santa Isabel 310 Makira 310 Choiseul 310 The New Georgia Islands 310 The Santa Cruz Islands 311 Rennell Island 311 Ontong Java Atoll 311

Vanuatu (New Hebrides) Espiritu Santo 312 Tanna 312 Aneityum 312

301

311

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Contents Erromango 312 The Banks Islands 312

The Fiji Islands

Lakes, Rivers, and Marshes Pedra Branca Rock 337

312

Kangaroo Island 337 Flinders Island 337 King Island 337 Lord Howe Island 337 Norfolk Island 338 Miscellaneous Islands 338 Balance for the Australian Realm

Viti Levu 313 Vanua Levu 313 The Kadavu Islands 314 Taveuni 314 Rotuma 314 Gau Island 314 The Lau Islands 314

New Caledonia

314

8

Grande Terre 315 The Isle of Pines 317 The Loyalty Islands 317

Balance for the Papua-Melanesian Realm 7

317

The Australian Realm 319 Mountains and Highlands 325 The Great Dividing Range 325 The The The The The The The The The

McIlraith Range 326 Melville Range 326 Clarke Range 326 Mount Carbine Tableland Paluma Range 326 Blue Mountains 326 Australian Alps 326 Central Highlands 326 Baw Baw Plateau 327

330

334

Miscellaneous Lakes, Rivers, and Marshes Coasts and Satellite Islands 334 Tasmania 335

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Wake Island

346

344

346

Nauru 346 The Gilbert Islands 346 Balance for the Micronesian Region The Novozelandic Region 347 The Kermadec Islands 349 New Zealand Main Islands 349

The Snares 358 The Bounty Islands 358 The Antipodes Islands 358 The Auckland Islands 358 The Campbell Island Group 358 Macquarie Island 358

334

xii

The Marshall Islands

342

The Chatham Islands 356 Novozelandic Sub-Antarctic Islands

333

The Fitzroy River 334 The Mary River 334 The Murray–Darling River Drainage

Pohnpei 344 Kosrae 344 Yap Island 345 The Palau Islands 345 The Chuuk Islands 345 The Nomoi Islands 346

342

346

North Island 353 South Island 354 Stewart Island 356

The Dalhousie Springs 332 The Elizabeth Springs 332 The Edgbaston Springs 333

334

342

Pagan 343 Saipan 343 Aguigan 343 Tinian 343 Rota 343 Guam 343

Temperate Savannas and Grasslands 330 Deserts and Shrublands 331 Isolated Caves, Springs, and Pools 332 The Great Artesian Basin 332

The Goulburn River

The Volcano Islands

The Caroline Islands

Tropical and Subtropical Wooded Savannas

Cooper Creek

Chichi Jima 342 Haha Jima 342

The Mariana Islands

326

339

The Polynesian Realm 341 The Micronesian Region 341 The Bonin Islands 342

Iwo Jima

The MacDonnell Ranges 327 Lowland Tropical and Subtropical Rainforests 327 Mediterranean Forests, Woodlands, and Scrub 327 Subtropical Mediterranean Forests and Woodlands 328 Mallee Scrub 328 Coastal Heathlands 328 Savannas and Grasslands 328 Tropical and Subtropical Savannas and Grasslands 329

Lakes, Rivers, and Marshes Lake Eyre 334

336

334

Balance for the Novozelandic Region The Polynesian Region 360 The Hawaiian Islands 361 Hawaii 363 Maui 364

357

359

Contents Oahu 365 Kauai 365 Lanai 365 Molokai 365 Niihau 365 Nihoa 365 Laysan 365

The Line Islands Teraina 366 Kiritimati 366

The Cascade Range 380 The Klamath Mountains 380

The California Coast Ranges 380 The Santa Cruz Mountains 381 The Santa Lucia Mountains 381

The Transverse Ranges 366

The Peninsular Ranges 366

The Marquesas Islands Nuku Hiva 367 Ua Huka 367 Hiva Oa 367 Fatu Hiva 367 Eiao 367 Ua Pou 367 Mohotani 367 Hatutu 367

The Colorado Plateau 381 The Appalachian Mountains

The Appalachian Plateau 382 The Ridge-and-Valley Appalachians The Blue Ridge Moutains 382

368

368

The Society Islands Tahiti 368 Raiatea 369 Moorea 369 Maupiti 369 Huahine 370 Niau 370 Makatea 370

370

The Gambier Islands 370 The Tonga Islands 370 370

The Cook Islands Rarotonga 370 Mauke 370 Atiu 370 Mangaia 371

370

The Austral Islands Tubuai 371 Rimatara 371 Rurutu 371 Rapa Iti 371

The Pitcairn Islands Pitcairn Island 371 Henderson Island 371

382

The Ozark Mountains 382 The Ouachita Mountains 382 Miscellaneous Mountains and Highlands

382

Lowland Forests 383 Lowland Boreal Forests 385 Lowland Broadleaf and Mixed Forests Lowland Coniferous Forests 386

386

Death Valley

389

The Sonoran Desert

371

387

389

389

The Chihuahuan Desert

389

The Guzmán Endorheic Basin 390 The Cuatro Ciénegas Endorheic Basin 390

The Baja California Desert 390 Isolated Caves, Springs, and Pools The Amargosa River 391

390

The Ash Meadows Complex 391

371

The Nearctic Realm 374 Mountains and Highlands 379 The Sierra Nevada Mountains 380 The Pacific Coast Ranges 380 The Olympic Mountains

California Coastal Sage and Chaparral 387 Lowland Grasslands, Savannas, and Shrublands The Great Plains 387 Western Gulf Coastal Grasslands 388 Lowland Deserts and Semi-Deserts 388 The Mojave Desert 389

The Colorado Desert

Easter Island 371 Balance for the Polynesian Region 372 Balance for the Polynesian Realm 373 9

382

South-eastern Conifer Forests 386 Florida Sand Pine Scrub 387

The Tuamotu Archipelago

Eua

381

The Santa Rosa Mountains 381 The San Pedro Mártir Mountains 381 The Laguna Mountains 381

Wallis and Futuna Islands 368 The Samoan Islands 368 Savai’i

381

The San Berardino Mountains 381 The San Gabriel Mountains 381 The Tehachapi Mountains 381

380

The Hediondilla Endorheic Basin 392 Lakes, Rivers, and Marshes 392 The Great Lakes 395 The Klamath Lakes 395 Lake Bonneville 395 Utah Lake 395

Lake Lahontan 396 Lake Waccamaw 396 The Mississippi River Drainage The Missouri River 396 The Ohio River 396 The Arkansas River 398

396

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Contents The Red River 398

The Coosa/Alabama/Mobile River Drainage The Coosa River 398 The Alabama River 399 The Mobile River 399

The San Marcos River

399

399

The San Marcos Springs

The The The The

398

Pascagoula River 399 Pearl River 399 Apalachicola River 399 Ouachita River 399

The Caddo River

The Turks Islands 415 The Caicos Islands 415

The Greater Antilles Cuba 415

400

The Pecos River 400 Devil’s River 400 The Conchos River 400 The Salado River 400 The San Juan River 400

The Colorado River

The Yaqui River

401

402

Miscellaneous Lakes, Rivers, and Marshes Coasts and Satellite Islands 403 Salt Marshes 405 The Pribilof Islands 405 The Aleutian Islands 405 The Alexander Archipelago 406 Vancouver Island 406 The Channel Islands 406 Santa Cruz Island 406 San Clemente Island 406

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Jamaica

423

Mountains 424 Lowland Moist Forests 424 Lowland Dry Forests 425 Isolated Caves, Springs, and Pools The Black River Morass 425

425

425

The Cordillero Central 426 Mona Island 426 Culebra 427 Vieques 427

The Cayman Islands

427

Grand Cayman 427 Little Cayman and Cayman Brac 427

The Lesser Antilles 427 The Virgin Islands 428

Socorro Island 407 Clarión Island 408 San Benedicto 408

The Andros Islands 414 The Exuman Islands 414

402

Anegada 428 Saint Croix 429 Virgin Gorda 429 Peter Island 429

Anguilla

429

Little Scrub Island 429 Sombrero Island 429

410

417

Mountains 420 Lowland Moist Forests 421 Lowland Tropical Dry Forests and Xeric Shrublands Lakes, Rivers, and Marshes 423

Puerto Rico

Guadelupe Island 406 The Coronado Islands 407 The Todos Santos Islands 407 Cedros Island 407 San Pedro Nolasco Island 407 San Marcos Island 407 Santa Catalina Island 407 The Revillagigedo Islands 407

10 The Caribbean Realm 412 The Lucayan Archipelago 413 The Bahamas 413

415

418

Hispaniola

402

Newfoundland 408 Bermuda 408 The Florida Keys 409 Miscellaneous Islands 409 Balance for the Nearctic Realm

415

The Sierra Maestra 416 Lowland Moist Forests 417 Lowland Dry Forests and Xeric Shrublands Lakes, Rivers, and Marshes 417 Coasts and Satellite Islands 417 The Zapata Swamp 418 The Canarreos Archipelago 418 The Sabana-Camagüey Archipelago 418

401

The Little Colorado River The Virgin River 401 The Gila River 402 The Bavispe River

The Turks and Caicos Islands

399

The Rio Grande

Grand Bahama Island 414 San Salvador Island 414 The Abaco Islands 414 The Acklins Bight 414 The Bimini Islands 414 The Inagua Islands 414 New Providence Island 415 The Plana Cays 415

Saint Martin 429 Saint Barthélemy 429 Saint Kitts and Nevis 429 Barbuda 429 Antigua 429 Redonda

430

Montserrat

430

422

Contents

The Guadeloupe Archipelago Grand-Terre and Basse-Terre 430 Marie-Galente 430 The Islands of the Saints 430 The Petite Terre Islands 430

430

The Ameca River 469 The Pánuco River 469 The San Pedro Mezquital River 469 The Grijalva River 470 The San Juan River 470 The Acandí River 470 The Atrato River 470 The Esmeraldas River 471 The Mira River 471 Miscellaneous Lakes, Rivers, and Marshes

Dominica 430 Martinique 431 Mount Pelée

Saint Lucia

431

432

The Maria Islands

432

Saint Vincent 432 The Grenadines

433

11 The Neotropical Realm 435 The Mesoamerican Region 436 Mountains and Highlands 440 Mexican Plateau 442 Sierra Madre Occidental 443 Sierra Madre Oriental 443 Trans-Mexican Volcanic Belt 445 Sierra Madre de Oaxaca 446 Mixteca Range 448 Sierra Madre del Sur 448 Tuxtlas Range 450 Sierra Madre de Chiapas 451 Central American Highlands 452 Isthmian Highlands 457 Tabasará Cordillera 460 San Blas Cordillera 460 Darién Range 460 Pirre Range 461 Baudó Range 461

Lowland Rainforests

471

Mangroves 472 Cozumel 472 Cocos Island 472 Coiba Island 472 Gorgona Island 472 Tumaco and Bocagrande Islands 472 The Marías Islands 473 The Bahía Islands 473 The Corn Islands 473 Escudo de Veraguas Island 474 Miscellaneous Islands 474

432

Barbados 432 Grenada 433 Balance for the Caribbean Realm

The The The The The The The The The The The The The The The The

471

Coasts and Satellite Islands

474

Balance for the Mesoamerican Region The Amazonian Region 475 The Northern Andes 484 The Cordillera Occidental 487 The Cordillera Central 494 The Cordillera Oriental 496 The Mérida Cordillera 508 The Venezuelan Coastal Ranges 509 The Sierra Nevada de Santa Marta 511 Inter-Andean Valleys 511

512

The Guiana Highlands

The Pakaraima Mountains 512 The Parima Mountains 515 The Iwokrama Mountains 515

461

515

Northern Mesoamerican Lowland Rainforests 461 Caribbean Lowland Rainforests 462 Isthmian–Pacific Lowland Rainforests 462 Southern Mesoamerican Lowland Rainforests 463

The Brazilian Highlands

Lowland Tropical Dry Forests

Lowland Tropical Rainforests and Swamp Forests

464

Central American Lowland Dry Forests 464 Pacific Equatorial Lowland Dry Forests 465

Lowland Deserts and Xeric Shrublands The Motagua Valley

465

Isolated Caves, Springs, and Pools

465

The Yucatán Cave and Cenote System 465 The Charco la Palma Spring System 465

Lakes, Rivers, and Marshes

466

Valley of Mexico Wetlands 467 Lake Chichancanab 468 Lake Pátzcuaro 468 Lake Atitlán 468 Lake Alchichica 468 The Lerma–Santiago River Drainage

465

The Atlantic Plateau 515 The Central Plateau 519 The Southern Plateau 519

Lowland Amazonian Rainforests 519 Lowland Atlantic Rainforests 524

Lowland Dry Forests

528

Coastal Caribbean Lowland Dry Forests 528 Lowland Atlantic Dry Forests 528

Tropical Grasslands and Savannas

528

Flooded Savannas and Grasslands 528 Lowland Cerrado Woodlands and Savannas

Caatinga Scrub 529 Isolated Caves, Springs, and Pools The São Domingos Karst Region

Lakes, Rivers, and Marshes 468

519

Lake Junín 532 The Amazon River Drainage

530

529

530

530

532

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Contents The Orinoco River Drainage 536 The Tocantins River 536 The Magdalena River 536 The Essequibo River 538 The São Francisco River 538 The Paraná River 538 The Uruguay River 538 Miscellaneous Lakes, Rivers, and Marshes

Coasts and Satellite Islands

Restinga 539 The Paraguaná Peninsula 539 Trinidad and Tobago 539 Margarita Island 540 Marajó 540 The Fernando de Noronha Islands 540 Trindade and Martim Vaz Archipelago 540 Queimada Grande 540 The Alcatrazes Islands 540 Miscellaneous Islands 541

The The The The The The The The

Andean Plateau 548 Atacama Plateau 548 Aconquija Range 548 Ambato Range 548 Fatima Range 548 Famatina Range 548 Uspallata Range 548 Baguales Range 548

560

Amsterdam Island 564 Saint Paul Island 564

South Georgia Island 564 Balance for the Antarctic Realm

The Cantillana Mountains 549 The Nahuelbuta Range 549 The Pelada Range 549

The Córdoba Ranges 549 The Somuncurá Plateau 549 Lowland Subtropical Dry Forests 549 Lowland Moist Forests 549 Lowland Valdivian Temperate Rainforests 549 Lowland Magellanic Subpolar Forests 549 The Gran Chaco 550 The Pampas 550 Lowland Deserts and Semi-Deserts 551 The Sechura Desert 551 The Atacama Desert 551 Argentine Salt Flats 551 The Pipanaco Salt Pans 551 The Great Salt Flats 551

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558

13 The Antarctic Realm 562 Antarctica 563 Isolated Sub-Antarctic Islands 563 The Kerguelen Islands 563 The Crozet Islands 563 Heard and McDonald Islands 563 Amsterdam and Saint Paul Islands 564

The Chilean Coastal Cordillera 548

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558

Mocha Island 559 The Falkland Islands 559 The Chonos Archipelago 560 Balance for the Patagonian Realm

546

Patagonian Desert 551 Lakes, Rivers, and Marshes 552 Lake Titicaca 552 Miscellaneous Lakes, Rivers, and Marshes

The Juan Fernández Islands Alejandro Selkirk Island 558 Robinson Crusoe Island 559

Balance for the Amazonian Region 541 Balance for the Neotropical Realm 542 12 The Patagonian Realm 544 Mountains and Highlands 546 The Central and Southern Andes

553

Isabela 556 Santa Cruz 556 Fernandina 556 Santiago 557 San Cristóbal 557 Floreana 557 Española 557 Pinta 558 Marchena 558 Genovesa 558 Pinzón 558 Rábida 558 Santa Fé 558 Darwin, Wolf, and Roco Redonda Islands

538

539

Coasts and Satellite Islands The Galápagos Islands 554

553

564

14 The Oceanic Realm 566 The Temperate Northern Atlantic Region The North Atlantic Ridge 578

576

The Seawarte Seamounts 578 The Azores Plateau 578 The Madeira Abyssal Plain 579

The North Sea 579 The Baltic Sea 579 The Bay of Biscay 579 The Mediterranean Sea 579 The Black Sea 579 Balance for the Temperate Northern Atlantic Region 580 The Temperate Northern Pacific Region The Emperor Seamount Chain 583 The Bering Sea 584 The Sea of Okhotsk 584 The Sea of Japan 585 The East China Sea 585

580

Contents

The Yellow Sea 585 The Gulf of California 585 Balance for the Temperate Northern Pacific Region The Tropical Atlantic Region 586 The Bahama Banks 591 The Cay Sal Bank

591

The Cape Verde Rise 591 Saint Peter and Paul Rocks 591 The South Atlantic Ridge 591 The Saint Helena Seamount Chain

592

The Gulf of Mexico The Campeche Bank

595

The Gulf of Guinea 595 Balance for the Tropical Atlantic Region The Western Indo-Pacific Region 596 The Mascarene Plateau 598 The Seychelles Bank

599

The Chagos–Laccadive Ridge The Red Sea 599

595

599

The Gulf of Aqaba 599 The Gulf of Suez 599

The The The The The

Arabian Sea 599 Persian Gulf 600 Gulf of Aden 600 Andaman Sea 600 Bay of Bengal 601

The Hooghly Estuary

606

The Saumarez Plateau 606 The South China Sea 606 The The The The The

Macclesfield Bank 607 Gulf of Thailand 607 Lingayen Gulf 607 Taiwan Strait 607 Verde Island Passage 607

The Philippine Sea

607

The Kyushu–Palau Ridge

The The The The The The The The The

Visayan Sea 608 Samar Sea 608 Bohol Sea 608 Sulu Sea 608 Celebes Sea 608 Molucca Sea 608 Java Sea 608 Bali Sea 608 Flores Sea 609

608

Banda Sea 609 Arafura Sea 609 Bismarck Sea 609 Solomon Sea 609 Coral Sea 609

Chesterfield Bank 610 Great Barrier Reef 610 New Caledonia Barrier Reef Holmes Reefs 610

610

The Bay of San Miguel 618

Balance for the Tropical Eastern Pacific Region The Temperate South America Region 618 The Sala y Gómez Ridge 619 The Scotia Sea 620

601

The Mozambique Channel 600 Balance for the Western Indo-Pacific Region The Central Indo-Pacific Region 601 The Norfolk Ridge 606 The Wanganella Bank

The The The The The

Shark Bay 610 Balance for the Central Indo-Pacific Region 610 The Eastern Indo-Pacific Region 611 The Tonga-Kermadec Ridge 612 The Hawaiian Ridge 612 Balance for the Eastern Indo-Pacific Region 612 The Tropical Eastern Pacific Region 613 The Galápagos Rise 615 The Mathematicians Ridge 617 The Cocos Ridge 617 The Malpelo Submarine Ridge 617 The Nazca Ridge 617 The Gulf of Panama 617

593

Lake Maracaibo 594 The Mesoamerican Barrier Reef

586

609

Dillon Shoals 609 Ashmore Reef 609

The The The The

592

593

The Caribbean Sea

The Timor Sea

601

The Scotia Arc

618

620

The Strait of Magellan 620 Balance for the Temperate South America Region 620 The Temperate Southern Africa Region 621 The Kerguelan Plateau 622 Balance for the Temperate Southern Africa Region 622 The Temperate Australasia Region 622 The Naturaliste Plateau 624 The Tasman Sea 624 The Great Australian Bight 624 Balance for the Temperate Australasia Region 624 Balance for the Oceanic Realm 625 15 Global Balance 626 Global Balance by Vertebrate Class 626

Bibliography Index 713

629

Colour plates can be found between pages 356 and 357.

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Published online by Cambridge University Press

Foreword

Location, location, location: the geography of endangerment. Everyone has images of endangered species – the iconic giant panda, or the tiger, or the blue whale. For those like me who are fanatic birdwatchers, one can easily add less familiar species. I so want to see a giant panda – the closest I’ve been is finding its fresh droppings of chewed bamboo. I also want to see a Chinese monal, a gorgeous pheasant that makes the same mountains its home. It, too, is endangered. Because bird tourism is such a major industry, I can quickly find a list of the bird species I must not miss while I’m there. Tour guides are there to help if needed. Birding teaches me that there are unique places where hard-to-find species concentrate. They are typically those with small geographic ranges. About 90 per cent of threatened species have small ranges. (The rest, like tigers, have large ranges, but for which their ferocity is a threat to us and our livestock.) Simply, geography matters. Matthew Richardson has taken this to heart. Because geography matters, we should look at endangered species from that perspective. No one has done this before so comprehensively, and his insights are vital. First, iconic species excepted, I often do not know what species of mammals, reptiles, amphibians, and fish live in the places where I seek my rare birds. There aren’t many tour guides for these species. It’s not that I don’t care; it’s just that such knowledge is often hard to find. There may not be a guide to all the threatened species in an area. Even if there is, do I want to carry even more books into the field than I already do? This book tells me what else is there.

Second, the book is replete with examples of species that haven’t been seen in an age – the Turkestanian salamander, the Balkan streber (a fish), the Sichuan dormouse, and the Sichuan pit viper. There is so much we don’t know about many species – where to look for them is paramount. Some areas shout that they are poorly explored. This comes at an exciting time. Crowd-sourced, citizen-science efforts have passed a billion bird observations on eBird. Other taxa lag, but iNaturalist logs a million observations in a good month. We are gaining knowledge of where species are at an unprecedented rate, and knowing where to look and what species to seek makes exploration ever more exciting. Parts of our own planet are the ‘final frontier’ when it comes to biodiversity. Above all, this book is a guide for practical conservation. I direct a conservation NGO, Saving Nature (www .savingnature.org), which helps local conservation groups acquire land, restore their natural habitats, and reconnect the fragmented habitats that remain. Where do we choose to invest our limited funds? We care about all species, not just the charismatic large-bodied species. Saving tigers is important, certainly, but many thousands of species are equally interesting and desperately in need of our protection. I now know where to see if a river, a marsh, a cave, some special forest or desert house species for which our actions might help. Much of conservation is about protecting the right locations, and now we have an excellent guide as to where those are. Stuart L. Pimm, Doris Duke Chair of Conservation, Duke University, and President, Saving Nature

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https://doi.org/10.1017/9781108863308.001 Published online by Cambridge University Press

Preface

This book is intended to provide a complete overview of all threatened and recently extinct mammals, birds, reptiles, amphibians, and fishes within the context of where they live in the world. My approach has been a systematic examination of each biogeographic realm and region, both terrestrial and marine, with a particular emphasis upon geographic features such as mountains and islands. All of the information within can more or less be found elsewhere. There is a myriad of sources on the subject, of course, not least of which being the IUCN Red List of Threatened Species upon which I primarily rely. It is this very wealth of data, however, that makes it so difficult for layman and expert alike to see, at a glance, the ‘whole picture’. My goal has been to provide that picture, in the hope that it will reveal patterns useful in the preservation of both species and habitats, and thereby help to put it all into perspective. Towards that end, I have included summaries for each section and a general reckoning at the end of the book, which together provide the bare numbers needed to show the priority areas. I should perhaps add that the omission of invertebrates and plants is not meant to indicate that they are somehow of lesser importance, but one can only do so much and these, I’m afraid, will need to await further volumes, and perhaps different authors. In terms of my own background, and my personal reasons for writing this book, they too are many and complicated. Simply put, I am an author and historian by training with a lifelong interest in wildlife conservation. Growing up in the town of Markham, in Ontario, Canada, I had the extraordinary privilege of living near one of the world’s greatest zoos. I spent much of my childhood at the Toronto Zoo learning about animals. It was the ideal sort of education for a budding naturalist, particularly in its ambitious, zoogeographically arranged layout. Best of all, for me, were the series of sprawling pavilions, each dedicated to a particular region: Indo-Malaya, North America, Eurasia, and of course the incomparable African Pavilion, at the time the largest building in Canada. Beneath its vast glass and steel framework lay an entire rainforest, complete with all the sights and smells and wonders, along with areas representing caves, deserts, and rivers. Here I would dream

of exploring these worlds for real one day, and vowed that at the very least I would one day work there as a zookeeper. Sadly, for whatever reason, I never did realize my ambition to work at the zoo that I loved, but it remains for me a magical place. Another enormous advantage that I had growing up in Markham was the still-prevalent wilderness to be found there. The endless ravines and fields, now sadly fallen victim to rapacious property developers, gave me both scope for exploration as well as a sense of the growing reality of environmental destruction. So too the cottage on Lake Muskoka that my family visited each summer. Siamese cats were always there as well (and continue to be so today), and later, rather more exotic pets. But most of all it was books that gave me the escape I craved, especially those of the British author and zoologist Gerald Durrell, Bernhard Grzimek’s Animal Life Encyclopaedia, James Fisher’s Zoos of the World, and Life Before Man by Z. V. Spinar, with its extraordinarily evocative illustrations by Zdenek Burian. One particular book from my youth had an impact that, in some ways, led directly to the present volume. It was entitled Let Them Live, by the Swedish author Kai CurryLindahl. The latter was a sort of ‘world tour’ of rare and extinct animals, and I was immediately fascinated by the stories that it contained and the exotic places it evoked, so much so that I actually stole it from my local library.* Oddly enough, Prof. Curry-Lindahl is almost completely forgotten today, despite having been one of the early pioneers of conservation. Little did I know, incidentally, that at the time that I first found this book during the 1970s the author himself was actually teaching at Guelph University, a little more than an hour’s drive from where I lived. I never had the honour to meet him, unfortunately, and until recently had never even managed to find a photo of him. He died in 1990.

*

I make no apologies for this: if I hadn’t done so, it would have long since been discarded, and my illicit copy appears to be the only one left in existence. I still treasure it to this day.

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Preface Figure A Kai Curry-Lindahl. (Credit: Kai Curry-Lindahl)

All of these early influences, along with my first serious travels, proved to be the ideal training for the writing career that I was about to fall into. My first published book, oddly enough, was not about animals, but rather, a history of the British royal family. While not an earth-shaking success, it did at least open some doors for me. And so it was that, at the beginning of the twenty-first century, I turned at last to natural history. I had already written a biography of my childhood hero Gerald Durrell, which despite gaining notice among agents and publishers never did see the light of day. My next project was an ambitious, four-volume book series entitled Living Primates. It was as a result of this that I had two very important breaks. The first was in making contact with the late Professor Colin Groves of the Australian National University, one of the most important taxonomists that ever lived. He advised and encouraged me, and never made me feel inferior despite the obvious fact that I was a complete amateur without so much as a degree to my name. He also introduced me to many other primatologists, most notably Dr Russell Mittermeiers, then President of Conservation International and himself one of the most respected scientific leaders in the world. Russ liked what I was doing and generously provided me with funding that enabled me to continue the work. Later, he brought me on board his own projects, which then included the second and third editions of Lemurs of Madagascar as well as the primate volume to the massive Handbook of the Mammals of the World series. It was a heady time, full of travel and excitement, and I was able to meet many other legends from the world of science and conservation, most notably the great wildlife artist Stephen Nash who remains a good friend to this day. It was during this period as well that I worked on a couple of occasions at Gerald Durrell’s extraordinary zoo in Jersey, in the Channel Islands off the coast of France. There I got to know many of the people who had featured so prominently in making the Durrell Wildlife Conservation Trust so pivotally important. Living there, in an old French manor house on the zoo grounds, was eerily like stepping inside some of my favourite books. I will always be grateful for such opportunities.

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So it was that, with the benefit of all these experiences and connections, I decided about 10 years ago that the time had come to take on the one book that I always wanted to write: an updated version of Prof. Curry-Lindahl’s Let Them Live. More to the point, I wanted to copy his basic approach, but on a much more ambitious scale, and to take advantage of the more exacting scientific standards and greater wealth of information that the intervening half-century had provided. Moreover, I wanted the book to serve as a history of wildlife conservation as well as its destruction. The actual writing of it came at a time of deep personal crisis for me, and has been something of a solace over the long years that it has taken to complete it. As mentioned at the beginning of this foreword, the present volume deals with all terrestrial, freshwater, and marine vertebrates that have become extinct during recent historic time (that is to say, since about the year 1500 A . D .), or which are currently threatened with extinction, or are so rare or specialized that even minor restrictions or alterations of their habitats may result in their extermination (i.e. Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List). Also discussed are those forms that were formerly considered to be threatened but which have since been saved through wise conservation policy, as well as those about which little or nothing is known but, it seemed to me at least, are clearly at risk. All taxa are included (i.e. species as well as subspecies), along with any notable populations. While such an approach of necessity tends to focus on ‘the rarest of the rare’, it is important to keep in mind that it is all really just a matter of degree. We have seen, time and time again, how apparently common species can very quickly be wiped out for any number of reasons. Ultimately, all life on this planet is at risk. So it is that, in recent years, the tendency has been to shift the emphasis away from saving individual species to saving entire ecosystems. For my part I believe that both approaches are equally necessary and valid, and indeed it is often that, by putting an emphasis on a single ‘flagship’ species, an entire habitat may be saved. Before proceeding, it would be helpful perhaps to take a closer look at the history of the IUCN and its Red List of Threatened Species. The International Union for the Conservation of Nature (and Natural Resources) was established in 1948, and is headquartered in Gland, Switzerland. As its name suggests, it is an international organization dedicated to both conservation and sustainability. As such, it gathers data and analysis, promotes research and field projects, and generally provides advocacy and education. It is best known, however, for compiling and publishing the Red List, which assesses the conservation status of fauna and flora the world over. The original Red List began in the 1950s as a simple card index of animals deemed threatened with extinction. This gradually evolved, over time, into the Red Data Books published during the 1960s and 1970s, by then covering all animals (as well as plants) regardless of their threat status. With this ever-growing number of species, however, it soon became difficult to keep up with the latest developments in a timely fashion, resulting

Preface

in a more committee-based approach to assessments and an emphasis on special reports. Eventually, with the advent of the Internet in the late 1990s, the Red List at last found a home online as Redlist.org, which had the advantages of greater access for people around the world as well as the ability, at least theoretically, to make updates as needed. It remains by far the most comprehensive and objective global approach for evaluating the extinction risk of plants and animals, providing conservation information for over 105,000 species of plants, animals, and fungi. In short, the IUCN Red List is regarded as the most influential source of information for species conservation in the world, and is used to inform and guide key national and international policy as well as on-the-ground conservation decisions. Over 15,000 scientists worldwide, most of them volunteers, currently contribute to the work, which is funded from a wide variety of donors. Thus, the Red List has served as my primary source of conservation assessment information. For my purposes as an author as well as a researcher, however, there have been many occasions when I have disagreed with certain assessments, and have not been afraid to do so. Having worked on a few of them myself for the Global Mammal Assessment, I know first-hand how difficult the process can be, particularly when one has little or no information to work with. In some rare cases this has meant actually omitting a species or subspecies from the book that is accepted on the IUCN Red List and listed as threatened, but which my own research has convinced me is dubious. In most cases, however, I have erred on the side of caution, including taxa that were once common, perhaps, but have lately become threatened and have yet to have their status updated. So too with the countless forms listed as ‘Data Deficient’. While I tend to agree that the majority in this category are either junior synonyms or at least valid forms although not in any immediate danger, all too often truly threatened species are simply dumped here and forgotten, usually due to a lack of follow-up research. For my part, if I find a valid species or subspecies known only, say, by a single museum specimen collected from a small patch of forest during the nineteenth century that has since been replaced with a large city, I am willing to make the leap that it is probably extinct, or at least extremely threatened. In dealing with species and subspecies it should be noted that mere ‘rarity’ is not always the best criterion for inclusion within a threat category. As noted, just as a common species may be wiped out very quickly for various reasons, so too a rare one may not necessary be threatened if sufficiently isolated or protected. Many types of deep-sea fishes, for instance, are naturally very rare, but are nevertheless stable and out of danger, at least for now. So too, many species or subspecies may have become extinct from a particular part of their former range (extirpated), or become regionally threatened by extinction, yet may still be relatively safe elsewhere. Very often, such threatened populations are vitally important to the health of its ecosystem. Apart from the IUCN Red List, of course, I have also drawn from literally thousands of books and academic papers. The sheer size of this volume has made it impossible, for lack of

space, to include every reference upon which the entries are based. I have, however, included a thorough bibliography and have always been careful to cite direct references. Modern technology has also made it very easy to rely upon the knowledge and criticism of experts around the world, many of whom have taken the time to provide personal reviews and corrections. It may be useful to compare the present volume as well with its other primary influence, Kai Curry-Lindahl’s Let Them Live. When the latter was writing his book during the late 1960s and early 1970s his approach had been almost entirely geographic, set within the biogeographic arrangement first formulated by Alfred Russel Wallace in the late nineteenth century. Wallace was a British naturalist, explorer, geographer, and biologist, best known for independently conceiving the theory of evolution through natural selection along with Charles Darwin. He undertook extensive fieldwork, first within the Amazon basin and then throughout the Malay Archipelago, where he identified the faunal divide now known as the Wallace Line which separates animals of largely Asian origin from those of what he termed ‘Australasia’. The nineteenth century’s leading expert on the geographical distribution of animal species, he is sometimes called the ‘father of biogeography’, and his 1876 two-volume work, The Geographical Distribution of Animals, would serve as the definitive text on the subject for the next 80 years. In it he laid out the basis for the same zoogeographic regions used by the Toronto Zoo in its layout of pavilions. Under his system, Earth’s terrestrial regions were divided up into six biogeographical regions (i.e. the Nearctic, Neotropical, Palearctic, Ethiopian, Oriental, and Australian). The oceans were not divided at all, and while certain isolated islands were singled out as distinct areas they were not treated separately. Wallace’s system has been amended slightly in the years since, but has remained essentially unchanged to this day. Professor Curry-Lindahl naturally followed this basic arrangement in his own book, utilizing a more or less continental or oceanic division to his chapters, but within each continent or ocean the approach had been ‘ecological’ (i.e. coasts, deserts, savannas, etc.), rather than biogeographical. His sections included ‘Africa’, ‘Eurasia’, ‘North America’, ‘Central and South America’, ‘New Guinea, Australia and New Zealand’, ‘Oceania’, ‘Isolated Islands in the Oceans’, ‘The Antarctic’, and ‘The Oceans’. All of his marine areas (e.g. islands, archipelagoes, and oceans) were, however, dealt with on a geographical basis alone. By this arrangement he endeavoured to connect environmental factors like man-made changes in various habitats to the disastrous effects that have often struck the animals living there. Then as now there were usually a number of factors behind the decrease of each species besides man’s direct predation and habitat alterations. In terms of species and subspecies, Prof. Curry-Lindahl’s book also reflected a much simpler period of scientific knowledge. This was a time long before the present mania for taxonomic splitting, whereby a widespread species may be

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Preface

reassessed as two or perhaps even several distinct forms, creating a new wealth of unexpected diversity along with considerable confusion. Oftentimes the split is an obvious one or involves some quite distinct subspecies being raised to species level, as for instance in the case of gorillas and orangutans. More recently, however, species are being ‘created’ based mainly upon tiny differences in their DNA, the animals themselves often looking very similar or even identical in appearance to one another. Many of these new taxa have very limited geographic ranges, and are as a result threatened. All of this has resulted in considerable disagreement among taxonomic experts, who fall broadly into two main schools: the ‘lumpers’ and the ‘splitters’. It is not my place to make definitive statements regarding taxonomy. Indeed, as with Prof. CurryLindahl, I do not really care much whether a taxon is labelled a species, a subspecies, or merely a ‘population of biological importance’, so long as the full diversity of a given region is represented. Thus, Prof. Curry-Lindahl had far fewer species and subspecies to deal with (just under 1400 in total), some of which have since proven to be invalid. But he also had far fewer data with which to work. That said, his book was an honest effort to include all species and subspecies of mammals, birds, reptiles, amphibians, and fishes that, in his day, were threatened with extinction, as well as those that had already become extinct during historic time (by his definition, around 1600 A . D .). His method of assessing them was logical and efficient. Threatened species, by definition, are usually quite localized in their distributions, and assigning them is usually relatively easy. Those occurring in various habitats or regions within a continent or marine area were mentioned only in connection with their characteristic habitat or primary range, and in general, geographical subspecies occurring on the same continent but frequenting different habitats received separate textual treatments. Those that occurred on multiple continents were, in general, treated where their main distribution range was located, although they might be mentioned more than once in the text. Where the range of such an animal was equally divided into more than one section, it was usually dealt with in connection with the section that appeared first in the text. Taking into account the latest scientific thinking, my own volume has been arranged into 14 distinct biogeographic realms, six of which are further divided into two or more biogeographic regions. The terrestrial realms are as follows: the Arctic Realm, the Palearctic Realm (divided between the Eurasian, Sino-Himalayan, and Saharo-Arabian regions), the Afrotropical Realm (divided between the Guineo-Congolian and African regions), the Madagascan Realm, the IndoMalaysian Realm (divided between the Oriental, Sundaic, and Wallacean regions), the Australian Realm, the PapuaMelanesian Realm, the Polynesian Realm (divided between the Micronesian, Novozelandic, and Polynesian regions), the Nearctic Realm, the Caribbean Realm, the Neotropical Realm (divided between the Mesoamerican and Amazonian regions), the Patagonian Realm, and the Antarctic Realm. Finally, the

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world’s oceans are here considered to be a single interconnected whole, the Oceanic Realm, divided between the Temperate Northern Atlantic, Temperate Northern Pacific, Tropical Atlantic, Western Indo-Pacific, Central Indo-Pacific, Eastern Indo-Pacific, Tropical Eastern Pacific, Temperate South America, Temperate Southern Africa, and Temperate Australasia regions. It should be noted that the Arctic and Antarctic realms are really sort of transition zones between the terrestrial realms on the one hand and the Oceanic Realm on the other, having elements of both. As with biogeography on a global scale, there has also been much work since Prof. Curry-Lindahl’s day on how to delineate various types of habitat within them, as for instance rainforests and deserts. In recent years a number of different systems have been developed by various non-governmental organizations (NGOs). The best known of these is perhaps the World Wildlife Fund’s system of biomes, ecoregions, and ecozones that, while comprehensive, remain incomplete and increasingly out of date. Conservation International favours a ‘Hot Spots’ approach; that is to say, one based on priority areas for biodiversity; while the Durrell Wildlife Conservation Trust uses one based on ‘Islands and Highlands’. All are quite valid, even heroic. I myself experimented with all of these biogeographic arrangements, along with one based entirely on geography. As an author, however, I ultimately found that for the purposes of organization neither a purely geographic nor purely biogeographic system was entirely satisfactory. Bioregions (or ecoregions, depending on what you want to call them) work very well on a limited basis, particularly for lowland areas. Unfortunately, when combined together on a global scale the individual parts simply do not add up to a unified whole. Through many years of trial and error (mostly error) I gradually evolved a hybrid approach instead; one based largely upon geographical features such as mountains, islands and lakes, but combined with more generalized categories for the lowlands. The main advantage of this method is that it shows, at a glance, all the threatened and recently extinct taxa from a certain place. Where this system breaks down is with the marine regions. Unlike the terrestrial regions of the world, the global ocean cannot be so easily divided. Many highly threatened species within the latter are to be found worldwide, which is why I have opted to treat the oceans as a single realm, broken down more or less biogeographically and very broadly. The common element in all of this of course is elevation, with the oceans merely being the inverse of the terrestrial. Set within this intricate framework are approximately 14,000 vertebrate taxa (i.e. species and subspecies). While it has been impossible to provide more than the most basic treatment for the vast majority owing to the limitations of space, a few are nevertheless dealt with at length and most receive at least a short comment. That said, biogeography is not an exact science, and it consequently hasn’t always been clear where to assign certain forms within the text. Freshwater fishes are perhaps the most difficult of all. A type of fish confined to a mountain stream, a desert lake, or to seasonal

Preface

pools in the middle of a rainforest may reasonably be placed in any of half a dozen subsections. So, too, one living within a coastal mangrove in both brackish and fresh water. Amphibians can also be incredibly problematic and for much the same reason, particularly those living in isolated caves or rivers. In such cases I have been forced to make judgement calls based on what seems most appropriate. Within each section and subsection of the book I have always endeavoured to work from general to specific, assigning animals to their most appropriate place in the text in order to highlight endemism wherever it exists. Each chapter therefore begins with those species that cannot easily be assigned within one single habitat. From there the organization takes on the additional aspect of elevation, starting with a general ‘Mountains and Highlands’ section and moving down to lowland forests of various sorts, grasslands, and deserts. Wetland areas are similarly treated with a general introductory section followed by lakes and rivers, the latter broken down by their tributaries and discussed from their upper reaches to their lower ones. This is followed by a section on coasts and major islands or islands groups, in which we normally navigate from north to south. Precise geographical terms are obviously of the highest importance within such a system. With very few exceptions, all names for places and physical features in this book have been given in English, in an effort to standardize and thereby avoid a confusing Babel of languages. Wherever possible, however, I have also provided the local names as well. So too with species and subspecies names: English names and punctuation are used as opposed to local names, although exceptions are of course made in cases where the local name is widely known. I have always tried to provide both a common name as well as the scientific (Latin) one, even where it has meant having to coin one of the former myself. Common names for those species that are threatened or extinct are given in bold where we first encounter them in the text. The common names for subspecies, however, as well as for species that have already been discussed earlier or those that are not deemed threatened do not appear in bold. Finally, in terms of illustrations, I have decided to include wherever possible actual photographs of now-extinct species and subspecies as are available, in the belief that they have immense historical importance as well as a powerful impact upon the reader. Sadly, far too many creatures have gone extinct without anyone ever having taken the trouble to photograph them. In such cases I have relied instead upon drawings and paintings. As with photographs I feel that older ones, rendered by those who actually saw the animal in life, are more compelling even if they are less scientifically accurate. In a few instances I have opted to also include images of notable, still extant forms. What would Prof. Curry-Lindahl think about the current environmental situation, compared to his own era? Certainly, from his point of view, things were dire enough. In the midtwentieth century threats to wildlife came mainly in the form

of overhunting and overfishing, with loss of habitat and pollution usually only contributary causes. Sailors from Russia, Norway, and Japan were whaling indiscriminately, the American military was napalming forests in South East Asia, and the Chinese and others had begun to lay the foundations of the horrific mass trade in wildlife for use in what has since become known as ‘traditional medicine’ (despite being neither traditional, nor medicine). Several nations had already begun to tame great rivers with enormous hydroelectric projects. Most ominously of all, the mass destruction of the still vast tropical forests of South East Asia, Equatorial Africa, and South America had already started to have an impact. But it was past extinctions, rather than potential ones, that seemed to have preoccupied him and his contemporaries the most. To quote the very first lines from Let Them Live, ‘Never in the realm of nature have so many been exterminated by so few in so short a time.’ Prof. Curry-Lindahl knew better than anyone the damage wrought by hunting, but like most people of his day, still saw an important economic and even recreational role for it, and certainly didn’t regard it as an evil in itself. Animals continued to be seen as little more than an unlimited, free resource to be exploited, and even the IUCN itself had been created mainly to ensure that this was done wisely, rather than for the preservation of biodiversity for its own sake. Indeed, the world’s wilderness areas were still, by and large, pristine and indeed relatively unexplored. The growing human population, although already becoming a concern, was still half of what it is today. Moreover, mass awareness of human impact on the planet was only just beginning. Most people by then were familiar with the stories of the dodo and the American bison, and could see that overpopulation was a potential problem, but sustainability and their own role in that didn’t enter into it. Nevertheless, an international environmental movement had begun, and Western nations were already enacting legislations to protect both species and habitats. Yet from the standpoint of the early twenty-first century things appear very different indeed. In the half a century since Prof. Curry-Lindahl’s book was published, many species and subspecies have been lost, although not as many as had been anticipated. Some have even made remarkable comebacks due to conservation action, while a few others, long feared lost, have been rediscovered. In terms of total biomass, however, the scale has been frightening. By one recent estimate some 60 per cent of all vertebrate populations have been destroyed since 1970, and vast areas of habitat are gone forever. Still worse, the core problem underpinning environmental destruction, that of human overpopulation, is arguably even more apocalyptic than had been predicted and continues to exponentiate far beyond what anyone could have imagined during the 1960s. Our numbers are increasing by over 200,000 each day, and we are now building the equivalent of a New York City every month. Earth has already far exceeded its maximum capacity. There are simply too many people. We don’t just need to halt this; we must somehow find a way get back down to a truly sustainable level, which is to say a global population

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of under two billion (roughly the level of the late nineteenth century). This is of course unlikely, particularly given the fact that the issue is no longer even talked about for the most part, either because of a feeling that there is nothing that can be done about it or that to do so is somehow politically incorrect. And yet it can be done, if only we all limit ourselves to having two children or less. In what particular ways does human overpopulation manifest itself in terms of the environment? First and foremost, of course, is habitat destruction and degradation. The unprecedented and unrestrained growth of cities and suburbs, along with farming, mining operations, water extraction, logging, hydroelectric dams and the like, as well as pollution in all its forms, are leaving the planet’s remaining wilderness areas fragmented and isolated. As people continue to push wildlife and wild places out often what remains must be strictly managed, like enormous, open-air zoos. Indeed, in the future, when speaking of a particular species’ range, it will likely consist of a grim list of protected areas, of varying effectiveness. In recent years the concept of the national park has evolved, and we now have ones of considerable size, both terrestrial and marine, with some even crossing international borders. Sadly, it has been shown that protected areas, even in supposedly developed countries, are only protected until they aren’t. Another consequence brought about by humans has been the introduction of invasive species wherever they have settled. Such fellow travellers, in particular rats, mongooses, goats, feral cats, and dogs, either prey upon species directly or compete with them in such a way as to make their survival difficult or impossible. We shall see time and time again in this book how introduced species have devastated native fauna, often also resulting in a severe economic impact. It is a record that emphasizes as well how islands are particularly vulnerable. Hunting continues to take its toll. Fortunately, thanks to improved legal protections, so-called sport hunting is less a threat to species as a whole. Gone are the days when individuals could kill rhinos and tigers indiscriminately, although a certain amount of ‘trophy’ hunting continues. It has been suggested by some that such activity can actually be an indirect aid to conservation, in that it brings in tourist dollars and gives an incentive to the local people to preserve certain species. Such ideas are of course spurious. Hunting simply has no role to play in conservation whatsoever, nor, indeed, any place in civilized society. But while sport hunting has diminished, socalled ‘bushmeat hunting’ and trapping have become pervasive throughout the tropical areas of the world, and are directly driving species into extinction. The overcollection of wildlife for use as food and other reasons also remains a serious threat to many species. In Asia, reptile and amphibian populations in particular have been devastated by a culture that consumes anything that moves. Sea turtles were once highly endangered by the collection of their eggs by the millions annually from their vulnerable nesting grounds. Such practices are perhaps, on some level, morally excusable. The greedy and nonsensical

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harvesting of wild animals for use in ‘traditional medicine’ certainly is not. As we have all seen in recent years, apart from the impact on species themselves, this practice is also putting human beings the world over at risk by means of devastating pandemics. Other forms of animal collection such as fishing are quite understandable, if undertaken sustainably. However, many areas of the oceans, not to mention certain freshwater lakes and rivers, have basically been emptied by intensive or industrialized practices. The global depletion of fish stocks such as cod, halibut, and salmon were an entirely preventable tragedy. The horrifying practice of ‘shark finning’ for shark fin soup, and the environmental toll of fisheries bycatch are corollaries. Pollution in all its forms continues to plague both humans and animals. Plastics are a global blight. Sea turtles and other animals commonly confuse plastic bags with jellyfish and are killed, while microplastics fill our oceans and have created vast garbage patches in the middle of each ocean. Oil spills imperil our coasts. In some areas people are now forced to wear masks in order to endure high levels of air pollution. City lights have robbed us of the night sky, and the sounds of industry are omnipresent. We have literally trashed our world. Apart from all of this there are also natural threats – stochastic events such as volcanic eruptions and hurricanes – that are particularly dangerous to those species living in vulnerable areas such as islands. Fires, both natural and manmade, are becoming more frequent and more devastating. Pathogens such as Ebola are affecting both humans and animals: indeed, the fungal disease Chytridiomycosis is literally wiping out scores of amphibians globally. Above all is the terrifying new threat of man-made climate change brought about by carbon emissions from fossil fuels. Sadly, our focus on voluntary carbon reductions was doomed from the start. Desertification and drought continue apace over wide areas and is just one example of its effects. The increased duration and frequency of El Niño warming events within the Eastern Tropical Pacific, which can cause severe and rapid declines for restricted-range, shallow water species, is another, as is the loss of coral reefs due to ‘bleaching’. In the very near future rising sea levels due to melting polar ice will inundate coastlines globally and actually submerge certain low-lying Pacific islands. One of the most important breeding islands for green sea turtles (Chelonia mydas), located off the coast of Queensland in Australia, is a case in point. Once it’s sunk, the turtles will nevertheless continue to be drawn there by some mysterious internal GPS. The planet is warming, and the root cause is overpopulation. Unless and until we reverse that, there really can be no hope. Yet, in the face of all this negativity, it must be said that there have been some positive changes in the last half a century as well. First and foremost, people in general are much more aware about their impact on the environment, and more motivated to do something about it. Scientific knowledge and technology have also vastly improved. Action is being taken on national and international levels, with effective coordination in

Preface

the fight against wildlife smuggling for instance, and there is now a vast array of non-governmental conservation organizations. Zoos and aquaria, once places of mere entertainment, have become powerful weapons. Gerald Durrell and other pioneers showed just how effective captive breeding and reintroduction efforts can be in saving species from extinction, and the list of such species continues to grow. What lessons have I myself learned in writing this book? I suppose the first thing that comes to mind is that I now see biodiversity conservation as not just one of many problems facing the world today, but as nothing less than a global war of annihilation. Moreover, it is a war that we are very close to losing. The enemy is numerous and varied: poachers, bushmeat and trophy hunters, developers, certain politicians, and above all practitioners of the illegal animal trade. Our allies are zoos, NGOs, and above all the people on the ground, all over the world, who have dedicated their lives to protecting and studying wildlife. Sadly, rangers and activists are all too often killed in the front lines. We must treat this war for what it is and begin to think in strategic terms. We need to understand as well that the aforementioned frontlines have shifted. Conservation is no longer simply about saving giant pandas or rhinos. Rather, it is about preserving those areas of the world that feature the highest biodiversity and which are most at threat. Islands and highlands are vital, of course, but it is overwhelmingly the tropical areas of the world that must be our priority. Half of Siberia, in terms of biodiversity, isn’t worth a single province in Costa Rica or Sumatra. What I have tried to do with my emphasis on geography and maps is to highlight these farflung battlefields. So, too, we must be smarter about where we

allocate our all-too-limited conservation funds. Above all, we need to be very careful about the dissemination of information. The enemy reads academic papers as well, and watches with interest camera trap photos of endangered species posted online. We must not continue to provide them with precise locality data, however much that flies in the face of academic tradition. So it is that I never reveal such information in this book unless it is already widely known. Sadly, we have now reached a day where certain types of information must only be disseminated on a ‘need to know’ basis. I believe that we are now quite literally living in that terrifying dystopia that Prof. Curry-Lindahl and others so feared, one of mass extinction and global environmental peril. For the first time in history people are seeing the actual effects of their own presence. The world is a much smaller place than it was just a few short decades ago, and we have gone beyond the fear of losing individual species – the ‘rarest of the rare’ – to that of losing whole ecosystems and perhaps, if we aren’t careful, all life on Earth as well. Faced with all these problems it is easy to feel helpless, but everyone can at least do their bit. So it is that this book serves as my own small contribution. It is my fondest hope that it will at least make people aware of the sheer scope of the global environmental crisis, and perhaps help to set priorities in its preservation. In the end, perhaps the book’s greatest value will be as an artefact for future generations. Another author, 50 years from now, will be able to use it in the same way that I have been able to use Prof. CurryLindahl’s book; as a point of comparison and an inspiration. Maybe by then we will have all come to our senses at last, and there will be no need for books of this sort at all.

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Acknowledgements

I would like to particularly thank the following individuals for their general support over the years, without which this project would not have been possible: the late Professor Colin Groves; Stephen Nash; Dr Lee Durrell; Dr Judith Eger; Dr Julia Sen. I would also like to thank Cambridge University Press and, in particular, my editors Dominic Lewis and Aleksandra Serocka. I could not have asked for a better publisher, and will always consider this one of the greatest honours of my life. The following individuals were kind enough to review sections of the book and to offer comments and corrections: Ramesh ‘Zimbo’ Boonratana; Giorgos Catsadorakis; Patricia Chapple-Wright; Anthony Cheke; Maarten J. M. Chistenhusz; Julian Fennessy; Douglas Gimesy; Steven Goodman; Alex Greewood; Juan Herrero; Heribert Hofer; Branden Holmes;

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Erich Hoyt; Julian Hume; André Koch; Jeffrey Lang; Edward Louis, Jr; Barita Manullang; Darren Naish; Giuseppe Notarbartolo di Sciara; James Perran Ross; Michael Proctor; Arne Rasmusson; Anders Rhodin; Christian Roos; Mariella Superina; Jatna Supriatna; Kenny Travouillon; Jean-Christopeh Vié; Glyn Young; Noam Werner; John Zichy-Woinarski. It goes without saying that all errors and omissions are strictly my own. A number of other individuals, both living and deceased, have also made valuable contributions to this work. Among them are: Bruce Beehler; Quentin Bloxam; Ian Colquhoun; Jamie Copsey; Anneke DeLuycker; Peter Grubb; John Hartley; Sander Hofman; Alison Jolly; Alain Kormann; Olivier Langrand; Trevor Lihou; Jeremy Mallinson; Russell Mittermeier; Anthea Morton Saner; Ira Pastor; Stuart Pimm; Noel Rowe; Simon Stuart.

Chapter

1

The Arctic Realm

The Arctic Realm, as here defined, are those terrestrial areas where the average temperature for the warmest month is below 10℃. It therefore includes all of the Arctic Circle including almost all of Greenland, the northern coast of Siberia and northern Scandinavia, northern Alaska, and northern Canada including the high arctic islands. The focal point is of course the Arctic Ocean, the smallest, shallowest and coldest of the world’s oceans and in many ways little more than an estuary of the North Atlantic. It consists of a roughly circular basin generally taken to include the Barents, Beaufort, Chuckchi, East Siberian, Greenland, Kara, Laptev and White seas, along with Hudson Bay and other tributary bodies of water. The latter is connected to the Pacific Ocean by the Bering Strait and to the Atlantic through the Greenland and Labrador seas. The Arctic Realm is bordered by the Nearctic and Palearctic realms to the south, and has affinities to both in terms of its fauna.

The Polar Ice Cap

The Earth’s northern polar region is covered by floating pack ice (sea ice) over the Arctic Ocean, which for the purposes of this book are considered terrestrial. Portions of the ice that do not melt seasonally can become very thick, up to 3–4 m over large areas, with ridges of up to 20. One-year ice is usually around 1 m thick. With global warming, the extent of arctic ice has decreased about 4.2 per cent per decade since the 1980s. The polar bear (Ursus maritimus) is the largest living bear species, as well as the world’s largest land carnivore. It is distributed in low densities throughout the Arctic Circle. Although some occur in the permanent multi-year pack ice of the central Arctic basin, they are most common in the annual ice over the continental shelf and inter-island archipelagos that surround it. The southern limit of the range extends to the coast of Newfoundland in the north-western Atlantic, while the northernmost record is just 25 km from the geographic North Pole. At least some populations seem to be nomadic, moving on the ice from east to west in a large circumpolar loop that includes Greenland and Baffin Island and in the Old World runs chiefly inside the large islands of arctic Eurasia. Polar bears move south in winter and north in summer, following the food supply – mainly seals – as the ice breaks up and shifts. This movement helps to explain the failure until relatively recently of any country to take

responsibility for the welfare of polar bear populations. The species has probably always been confined to arctic areas, although from time-to-time individuals have strayed to Iceland, the Norwegian mainland, Manchuria, and Japan. Those that have continuous access to sea ice are able to hunt throughout the year. However, those living in areas where the sea ice melts completely each summer are forced to spend several months on land, where they primarily fast on stored fat reserves until freeze-up. This use of land by polar bears during the ice-free season is increasing, at least in some areas. Intensive hunting of the species did not begin until the early seventeenth century, but then increased to such a degree that by 1850 it had been seriously depleted, particularly in the Spitsbergen area and on Novaya Zemlya. In addition, when most of the arctic whales had been exterminated men began to hunt the seals, which increased the hunting pressure on the polar bears as well. It soon became evident that polar bears were declining in number all over their range, including the New World. With increased protection populations began to recover, although ‘subsistence harvesting’ is still allowed in Alaska, Canada, and Greenland (but prohibited in Norway and Russia). Currently, it is the loss of arctic sea ice due to climate change that is the most serious threat.

Tundra The vast arctic tundra is a circumpolar region in the far Northern Hemisphere, north of the boreal forest belt. It is here defined as comprising parts of northern coastal Scandinavia, northern coastal Russia, northern coastal Alaska, the Canadian High Arctic, and the islands of Greenland and Iceland. The word ‘tundra’ usually refers only to areas where the subsoil is permanently frozen (‘permafrost’). The lands within this region are typified by cold winters and cool summers, low precipitation, and an absence of trees. Biodiversity in the tundra is low, with the few plants and animals that manage to survive there being adapted to short growing seasons with long periods of sunlight, as well as to extreme cold, dark, snow and ice-covered winter conditions. Today they are essentially climax communities, with lichens serving as a basis for the existence of the large herds of reindeer or caribou (Rangifer tarandus), which in their turn are the staple winter food of the grey wolf (Canis lupus).

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During prehistoric times the muskox (Ovibos moschatus) occurred throughout the Siberian and North American Arctic from the Urals east to Greenland and south as far as the ice sheets extended. It appears to have died out in Europe around 9000 years ago, and in Siberia around 2000 years ago. In the nineteenth century it was still to be found from Point Barrow, Alaska east across Canada to north-eastern Greenland and south to north-eastern Manitoba. By the early twentieth century it had been wiped out in Alaska by excessive hunting, but has made a considerable recovery in other areas. In the Canadian arctic they now inhabit most large islands (with the exception of Baffin Island) and the mainland tundra of the Northwest Territories and Nunavut from the coast of Hudson Bay west to almost the Mackenzie River, south to the tree line. They still occur naturally over the entire north of Greenland, in addition to several introduced populations further south. The species has also been reintroduced to parts of Alaska (beginning with Nunivak Island as early as 1935), Norway, and the Taimyr Peninsula and Wrangel Island in Russia. Taken together the species appears to be safe. The reindeer (Rangifer tarandus), also known as the caribou in North America, is a type of deer found, as a species, both in the northernmost regions of North America and Greenland as well as in northern Eurasia. Despite this wide range many subspecies are threatened, and a few have already gone extinct. The Barren Ground caribou (R. t. groenlandicus) is found in the Canadian High Arctic islands (Nunavut and Northwest Territories) and western Greenland. The Peary caribou (R. t. pearyi) is found in the Canadian High Arctic islands (Nunavut and Northwest Territories). The Labrador caribou (R. t. caboti) occurs in the tundra regions of Quebec and Labrador. The Porcupine caribou (R. t. granti), so-named for the Porcupine River which runs through much of its range, lives in northern Alaska and adjacent north-western Canada (Yukon). The single herd of around 200,000 animals migrates some 2400 km each year between their winter range and their calving grounds near the Beaufort Sea, the longest land migration route of any mammal. It is highly vulnerable to climatic factors, and the population fluctuates greatly as a result. The Siberian tundra reindeer (R. t. sibiricus) is still found across much of northern Eurasia, but has been supplanted almost everywhere by domesticated reindeer. The largest remaining population of wild reindeer in the Old World lives in the Pyasina River drainage on the Taimyr Peninsula in northcentral Siberia, where the number of individuals exceeds 100,000 as a result of partial protection. The snowy owl (Bubo scandiacus) still has a considerable range across predominantly open tundra regions from western Scandinavia through northern Russia to Alaska, northern Canada, and Greenland. It has also bred occasionally in Iceland and in the United Kingdom. During the winter the birds move further south into the mainland United States, northern Europe, and northern Asia. The species appears to be undergoing a considerable decline in population, however, and is now thought to number less than 30,000.

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The Siberian crane (Leucogeranus leucogeranus) is a large and spectacular snow-white species historically spread over an immense area from the Ural River in the south-east to the coast of the Arctic Ocean in north-eastern Siberia, where it favours bogs in conifer forests and steppes. Although long protected by law over most of its range, it was much persecuted and disturbed by hunters over its long migration routes and, like the more famous whooping crane (Grus americana) of North America, had difficulty maintaining its numbers. By the mid-twentieth century it was already reduced to two widely separated groups, known as the Eastern Flyway and Western/ Central Flyway populations. The vast majority, the Eastern Flyway population, breed in the Yakutia region of northeastern Arctic Russia between the Kolyma and Yana rivers and south to the Morma Mountains. Younger, non-breeding birds summer in Dauria on the border between Russia, Mongolia, and China, and occasionally in central Mongolia as well. The main wintering sites were formerly in the middle to lower reaches of the Yangtze River, although today almost all winter at or near Lake Poyang in China. The birds rely on a network of important wetlands along their migration route, which follows the Yana, Indigirka, and Kolyma rivers before continuing along the Aldan River and its tributaries and south into China. The other, remnant population of Siberian cranes (less than 20 birds) breeds in West Siberia and is divided into two further subpopulations, the Western Asian Flyway and Central Asian Flyway flocks. The Central Asian flock breeds in the Kunovat River drainage in Russia, and historically wintered in Keladeo National Park, north-western India. None have been seen at Keoladeo since the early 2000s, although unconfirmed but credible reports of passing birds continue from Russia, Uzbekistan, Kazakhstan, Pakistan, and India. The Western Asian flock breeds in the Konda and Alymka river drainages of West Siberia, and winters in Fereydoonkenar in Iran. It uses the Volga River delta as a migration stopover, and passes over Azerbaijan during its migration. Captive-breeding and reintroduction efforts have recently begun in Iran. Currently, the total population for the species as a whole is about 3750, up from less than 2000 in 1965. The red-breasted goose (Branta ruficollis) breeds on the Taimyr, Gydan, and Yamal peninsulas of north-central Russia. Prior to the 1950s much of the population wintered along the western coast of the Caspian Sea, primarily in Azerbaijan, and in Iran and Iraq, although the wintering grounds thereafter rapidly shifted to the western Black Sea coast. The total number is small and prone to dramatic fluctuations for reasons that are not fully understood. The lesser white-fronted goose (Anser erythropus) historically bred across much of subarctic Eurasia but is now confined to four main areas of northern Scandinavia and the northern coast of Siberia, from where it migrates in winter to Europe, the Middle East, and southern Asia. It has undergone a significant decline due to by-catch mortality in gillnets, oil pollution, disease, and hunting.

Coasts and Satellite Islands

The long-tailed duck (Clangula hyemalis) has a circumpolar distribution, breeding along the arctic coasts of North America, Greenland, Iceland, Europe, and Asia, from where it winters in the north-eastern United States, north-western Europe, and in central and western coastal Asia. It is threatened by hunting, fisheries by-catch, pollution, and disease. Steller’s eider (Polysticta stelleri) is a small sea duck that breeds patchily along the northern coasts of Siberia and Alaska, from where it winters in Novaya Zemlya, Norway, south-western Alaska, and northern Japan. It has undergone a significant decline, particularly in Alaska, due to hunting, habitat destruction, and possibly the effects of climate change. The Eskimo curlew (Numenius borealis) was historically abundant in its breeding grounds on the barren tundra of Canada north of the Arctic Circle, roughly between the Bathurst Peninsula and Point Lake and perhaps extending into Alaska. It migrated southward across Hudson Bay to Labrador and New England, whence it started its non-stop flight over the Atlantic Ocean and the Caribbean Sea to winter in southernmost South America. The spring flight followed a more westerly route, because there are records indicating a flyway passing over Yucatán, Texas, and west of the Great Lakes to north-western Canada. Hence, it flew twice a year across both Americas. Excessive shooting during the migration in eastern Canada and New England is thought to account for the tragic decline of this species. In 1863 over 7000 birds were killed in one day on Nantucket Island. As late as the period between 1856 and 1875 immense flocks used to rest in Texas, but by 1905 only three birds were seen there. Hecatombs of these curlews were shot as they migrated across the United States. Hunters sent wagonloads of birds back from the shooting grounds. They were very easy to kill, because they fed close together and were trustful of humans. In 1929 A. C. Bent, in his Life Histories of North American Shore Birds, declared of the curlew: ‘It is now but a memory of the past.’ However, in 1932 an individual was taken in Newfoundland, and a week later four were seen on Long Island. It was last confirmed from its wintering grounds in South America in 1939. Sporadic sight records followed elsewhere at long intervals: 1945, 1946, 1950, 1959, 1960, 1962, 1963, and 1970 on the coast of Texas and Louisiana as well as on the Atlantic coast. Some of these are doubtful, but in 1963 one bird was shot with certainty in Barbados. While there have been several unconfirmed reports in the decades since, with the latest alleged sighting occurring in Barbados as recently as 2012, the species is now almost certainly extinct. A related species, the bristle-thighed curlew (N. tahitiensis), breeds in the tundra on the lower Yukon River and the central Seward Peninsula of western Alaska, wintering on various islands in the South Pacific. The total population is estimated at around 10,000. Two migratory wading birds of the genus Calidris are threatened by coastal development in their wintering grounds. The spoon-billed sandpiper (C. pygmaea) breeds in north-

Figure 1.1 One of four known photos of a living Eskimo curlew, taken on Galveston Island, Texas, in 1962. (Credit: Don Bleitz, courtesy of the archives of the Western Foundation of Vertebrate Zoology, Camarillo, California.)

eastern Russia and winters in South East Asia, where it has declined dramatically since the 1970s. The current total population is believed to be around 500. The great knot (C. tenuirostris) breeds in north-eastern Siberia and winters mainly in Australia, but also patchily throughout the coastal areas of South and South East Asia and the Arabian Peninsula.

Coasts and Satellite Islands This section includes the coastal areas of northern Russia, northern Alaska, and northern Canada (along with the High Arctic islands), as well as Greenland, Iceland, and various smaller islands. The walrus (Odobenus rosmarus) is a very large, flippered marine mammal characterized by its tusks and whiskers, which it uses to help obtain the molluscs upon which it feeds. It is still found across a large area of polar seas, although its distribution was much reduced as a result of the severe exploitation that began in the sixteenth century. It was not until the introduction of modern firearms and arctic transport, however, that the species began to be seriously threatened. It was considered an important natural resource, yielding oil, hides, and ivory. Now somewhat protected from hunting, it still faces an uncertain future due to habitat changes brought on by climate change. The Atlantic walrus (O. r. rosmarus) was historically common along the coasts of the North Atlantic and the Arctic Ocean southward to the Russian and Norwegian mainland and, in the Western Hemisphere, to the coast of Labrador, with vagrants being reported as far south as New England and the Bay of Biscay. Today it is found discontinuously from the eastern Canadian Arctic and Greenland to the western Kara Sea. The Pacific walrus (O. r. divergens) recovered remarkably from a seriously depleted state in the mid-twentieth century, and currently ranges from the Bering and Chukchi seas, which constitutes the core of its range, to

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the Laptev Sea in the west and the Beaufort Sea in the east. Vagrants are occasionally reported in the North Pacific south to Japan and south-central Alaska. The hooded seal (Cystophora cristata) is so-named for an inflatable sac found on the head of adult males. It lives on drifting pack ice in the Arctic Ocean and the central and western North Atlantic, ranging from Svalbard in the east to the Gulf of St. Lawrence. Prior to the 1940s adults were hunted extensively for their leather and oil, and the young for their distinctive blue and black pelts. The animals are also frequently killed for subsistence hunting. Numbers have increased in most areas with better protection in recent decades, which includes an allowable catch limit of 10,000 annually, but the species remains vulnerable to the effects of climate change. The Ungava harbour seal (Phoca vitulina mellonae) is a freshwater species confined to a few lakes and rivers in northern Quebec, where the total population is thought to be less than 100. The harp seal (Pagophilus groenlandicus) occurs in the northernmost parts of the Atlantic and throughout much of the Arctic Ocean. The species is strongly migratory, with the main breeding grounds in the White Sea, on the pack ice in Norwegian waters (particularly off Jan Mayen), off Labrador and northern Newfoundland, and in the Gulf of St. Lawrence. At the beginning of the twentieth century and even as late as the early 1940s the world population was estimated at about 10 million. Owing to reckless hunting this number had fallen to about 3 million within 20 years, where it has remained more or less stable ever since. A particularly cruel and barbaric form of commercial sealing, in which the pups are brutally bludgeoned for their fur, still takes place annually in Canada, Norway, Russia, and Greenland. The destruction is particularly marked in Newfoundland, where the small profit earned by seasonal hunters is far outweighed, economically, by the enormous damage done to Canada’s international reputation.

The Arctic Archipelago The Arctic Archipelago includes all of the high arctic islands lying to the north of the Canadian continental mainland, with the exception of Greenland. Two subspecies of grey wolf (Canis lupus) historically inhabited the Arctic Archipelago. Bernard’s grey wolf (C. l. bernardi) is known only from a few specimens collected from Banks and Victoria islands, where it died out around 1920. The Arctic wolf (C. l. arctos) is confined to the Queen Elizabeth Islands, but is not currently considered to be threatened.

Greenland Greenland (Kalaallit Nunaat in Greenlandic; Grønland in Danish) is the world’s largest island. It is almost entirely covered by a massive ice sheet, the weight of which has depressed the central land area to form a basin lying more than 300 m below sea level. Elevations along the more temperate coasts rise suddenly and steeply.

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The East Greenland caribou (Rangifer tarandus eogroenlandicus) appears to have been confined to the tundra regions of eastern Greenland, where it went extinct around 1900.

Svalbard Svalbard is an archipelago located about midway between Norway and the North Pole. Originally used as a whaling station during the seventeenth and eighteenth centuries, today the only permanently inhabited island is Spitsbergen. The Svalbard reindeer (Rangifer tarandus platyrhynchus), the smallest of all the reindeer, is confined to the Svalbard Archipelago.

Novaya Zemlya Novaya Zemlya is an archipelago in the Arctic Ocean of northern Russia and extreme north-eastern Europe. It is composed of two main islands, the northern Severny Island and the southern Yuzhny Island. The Novaya Zemlya reindeer (Rangifer tarandus pearsoni) is confined to the archipelago. At the end of the nineteenth century there were about 20,000 reindeer on Novaya Zemlya. Heavily hunted both for local consumption as well as export, they would be reduced, only a few decades later, to just a handful of survivors on the northeastern part of Severny Island. Fortunately, a prohibition on hunting was put in place in time to save the subspecies, and it has since recovered.

Wrangel Island Wrangel Island is located in the Arctic Ocean between the Chukchi and East Siberian seas. The Wrangel lemming (Lemmus portenkoi) and Wrangel collared lemming (Dicrostonyx vinogradovi) are both confined to Wrangel Island, where they are considered intrinsically vulnerable due to their small range and marked population fluctuations.

Balance for the Arctic Realm The Arctic Realm was among the last places on Earth to be permanently settled by humans. The first to live there arrived in Siberia around 20,000 years ago, from where they slowly migrated eastward across the Bering Strait land bridge to North America and, finally, Greenland. These prehistoric peoples were, and largely remain, nomadic hunter-gatherers, entirely dependent on the reindeer herds and marine mammals for sustenance. They ultimately developed into the Inuit, a group of culturally similar indigenous people that live throughout the region today. Modern European exploration of the Arctic Realm also began relatively late. Greenland was first reached (and partially mapped) as early as 1499 by the Portuguese explorers Gaspar and Miguel Corte-Real. After that, exploration was undertaken either by land east from Russia, or by western Europeans seeking a Northwest Passage to the Old World. The latter

Balance for the Arctic Realm

would result in the mapping of what is now the Canadian High Arctic, Alaska, and the islands of the northern Pacific. By the early twentieth century the focus was the North Pole. In 1908–09 the Americans Frederick Cook and Robert Peary each claimed to have reached it, although both are now widely doubted. In 1926 Roald Amundsen, Lincoln Ellsworth, and Umberto Nobile in the airship Norge became the first definitely known to have sighted the North Pole. With the coming of Europeans to the Arctic during the nineteenth and twentieth centuries, pollution and hunting began to take more of a toll, particularly on large animal populations. However, the latter were also responsible for the creation of the first national parks and environmental protections. The northern polar region remained relatively pristine up until very recently. Indeed, it still contains some of the last, and most extensive, wilderness areas remaining in the world. Today, however, the Arctic is being opened up at an increasing pace for exploitation of its vast wealth of natural resources, which include oil, natural gas, minerals, fish,

and, to some extent, forests. Settlement and tourism will only increase as human populations continue to expand, perhaps encouraged by the Arctic’s extraordinary abundance of freshwater (about one-fifth of the world’s total). All of this will be to the detriment of its sensitive environment, fragmenting habitats, eroding ground cover and disturbing important breeding grounds. The primary threat, however, is now global warming, with the consequent shrinkage (and ultimately perhaps complete loss) of arctic sea ice and the Greenland ice sheet, as well as the thawing of permafrost. In recent historical time (i.e. since A . D . 1500), the Arctic Realm has lost three subspecies of mammal and one species of bird. In addition, there are 14 species/9 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 5 species/9 subspecies are mammals, and 9 species are birds.

Vertebrate Class

Extinct

Possibly extinct

Extinct in the Wild

Threatened

Mammals

~ species

~ species

~ species

5 species

3 subspecies

~ subspecies

~ subspecies

9 subspecies

3 taxa

~ taxa

~ taxa

14 taxa

1 species

~ species

~ species

9 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

1 taxon

~ taxa

~ taxa

9 taxa

1 species

~ species

~ species

14 species

3 subspecies

~ subspecies

~ subspecies

9 subspecies

4 taxa

~ taxa

~ taxa

23 taxa

Birds

Total vertebrates

Note: ~, not applicable.

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Chapter

2

The Palearctic Realm

The Palearctic Realm, as here defined, is divided into three zoogeographic regions (Eurasian, Saharo-Arabian, and SinoHimalayan) that together comprise all the Old World terrestrial areas as far south as northern Africa, the Middle East, the mountains of Central Asia, and into southern China and the islands of Japan. The tiger (Panthera tigris) is the largest cat species and, with its striped fur pattern, the most instantly recognizable. A number of subspecies once ranged collectively across much of Asia from the Black Sea in the west to the Indian Ocean in the south, and from the Russian Far East to Indonesia. Over the past century they have lost at least 93 per cent of their historic range and have been extirpated from Western and Central Asia, the islands of Java and Bali, and from large areas of south-eastern, southern, and eastern Asia due to habitat destruction and hunting (both for trophies as well as use in ‘traditional medicine’). The snow leopard (Panthera uncia) has a wide but scattered distribution in the high mountains of Central and South Asia, with core areas including the Altai, Tian Shan, Kun Lun, Pamir, and Karakorum ranges. The species has declined everywhere owing to persistent illegal hunting for its beautiful and valuable fur and for its bones, which are used in ‘traditional medicine’. The leopard (P. pardus) has, as a species, the largest distribution of all wild cats, occurring widely if patchily across most of Africa as well as eastern and southern Asia. Nevertheless, a number of subspecies have been largely or wholly wiped out. Prehistoric fossil remains of the Asiatic black bear (Ursus thibetanus) have been discovered at various localities in Europe as far north as the Ural Mountains and in Germany and France, although in historic times the species has been limited to Asia, where it remains widespread, if patchily distributed, in a variety of forest types. Habitat loss combined with hunting for skins, paws and, increasingly, for gall bladders used in ‘traditional medicine’ have all contributed to declines, and several subspecies are considered threatened. The Indochinese black bear (U. t. mupinensis) is still found over a wide area of the Himalayas and Indochina. The Ussuri black bear (U. t. ussuricus) lives in southern Siberia, northeastern China, and on the Korean Peninsula. The grey or timber wolf (Canis lupus) was, historically, the world’s most widely distributed animal species, being found

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across much of the Old and New Worlds. While the nominate form, the Eurasian grey wolf (C. l. lupus), is not considered threatened, a few other generally recognized subspecies do have limited distributions and will be discussed below. Another, the Ezo grey wolf (C. l. hattai), once ranged across Hokkaido, Sakhalin, the Kamchatka Peninsula, and the Kuril Islands, but appears to have gone extinct sometime during the late nineteenth century. The Asiatic wild dog or dhole (Cuon alpinus) was historically found throughout much of central, eastern and southern Asia but has disappeared from most of these areas. Surviving populations are fragmented and continue to decline due to habitat destruction and depletion of their prey base. The Ussuri wild dog (C. a. alpinus) remains widespread on the Indian subcontinent and Indochina, but is most likely extirpated from China, Mongolia, and the Russian Far East. The aurochs (Bos primigenius) was an impressive species of wild cattle that once inhabited the forests and grasslands of Europe, Asia, and North Africa. A large animal (modern bulls weighed around 700 kg), it was the ancestor of European domestic cattle and, it has been suggested, possibly of the European bison (Bison bonasus) as well. There were three subspecies. The North African aurochs (B. p. africanus) and the Indian aurochs (B. p. namadicus) were both extinct before about A . D . 1500 owing to the destruction of forests, competition from domestic animals, and hunting. The Eurasian aurochs (B. p. primigenius) survived somewhat longer. Historically it was found from the British Isles and Scandinavia in the north-west to the Mediterranean countries in the south and Siberia and Syria in the east; by the thirteenth century it was restricted to Russia, Poland, Lithuania, Moldova, Transylvania, and East Prussia. By 1409 only Poland and perhaps Russia seem to have possessed surviving populations. Efforts to preserve these magnificent animals were made in the former country, and a small herd persisted in the Jaktorów Forest near Warsaw until the beginning of the 1620s. In 1627 the last individual there, a female, died. Beginning in the 1920s, attempts were made to create look-alikes by means of selective breeding. More recently, a project to ‘breed back’ cattle that not only resemble aurochs but can fill their ecological role, through eventual reintroductions to the wild, have been explored. The Asiatic wild ass (Equus hemionus) was originally found throughout the steppes and deserts of central Asia and

The Palearctic Realm

the Middle East, but has been almost entirely eliminated everywhere due to competition with livestock and overhunting. Five subspecies survive in isolated pockets, which will be discussed below. The wild horse (E. ferus), which once ranged across the steppes and grasslands of Europe and Asia, had three subspecies that survived into modern times: the domesticated horse (E. f. caballus), the undomesticated Eurasian wild horse or tarpan (E. f. ferus), now extinct, and the still-extant Mongolian wild horse (E. f. przewalskii). The latter two will be discussed below. The term ‘wild horse’, incidentally, is also used colloquially in reference to free-roaming herds of feral horses such as the mustang in the United States and the brumby in Australia, but these are all untamed members of the domestic horse subspecies, and not to be confused with true wild horse subspecies. The wild goat (Capra aegagrus) is the ancestor of the domestic goat (C. hircus). Nominally widespread in the high rocky and mountainous areas of Europe, Asia Minor, the Middle East and Central Asia, it is everywhere rare and often absent in many parts of its former range. Threats include hunting (particularly for its majestic curved horns) and loss of habitat. The bezoar wild goat (C. a. aegagrus) is found sporadically in central Afghanistan, Armenia, Azerbaijan, Georgia, Turkmenistan, Iran, Turkey (Anatolia), and possibly extreme northern Iraq. It was extirpated from Syria and Lebanon in the early twentieth century. The marbled polecat (Vormela peregusna) inhabits deserts, semi-deserts and steppe habitats from south-eastern Europe through central Asia to northern China and south to the Middle East. Although widespread, it has declined everywhere due primarily to loss of habitat. The Japanese sea lion (Zalophus japonicus) was historically confined to the region of the Sea of Japan, more than 8000 km from the nearest colonies of its closest relative, the California sea lion (Z. californianus). It was known for certain from Kyushu, Shikoku, and on islands around Honshu. The last credible reports were from 1951, when 50 or 60 of these animals lived around Takeshima, a rocky islet in the open sea between Japan and Korea, but these soon disappeared after the island was occupied by soldiers. There have been no documented reports whatsoever since the late 1950s despite extensive search efforts. Individual sightings in 1974 and 1975 cannot be verified, nor can confusion with escaped California sea lions be ruled out. The Mediterranean monk seal (Monachus monachus) was historically found continuously on the coasts and islands of the Black Sea and along those of the Mediterranean to North Africa and the major islands of the North Atlantic. Hunting, pollution, and disturbances in the caves where it breeds led to a severe decline during the twentieth century. It has been extinct in the Black Sea since the 1990s (although a few may still survive in the Sea of Marmara), and today only a few small, isolated colonies exist in the Mediterranean itself, mainly in the Ionian and Aegean seas, the coast of mainland Greece, Cyprus, and western and southern Turkey. An unknown

number may still survive as well on the Mediterranean coasts of eastern Morocco and perhaps Algeria. It was formerly to be found in North Atlantic waters as well from Morocco to Cabo Blanco, including the Canary Islands, Madeira Islands, and the Azores. Vagrants have been reported as far south as Senegal, the Gambia and the Cape Verde Islands, and as far north as Portugal and the Atlantic coast of France. Today, only two Atlantic subpopulations are known to exist: one at Cabo Blanco on the Mauritania/Western Sahara border, and the other in the Madeira Islands. The total population is thought to be under 500. The Eurasian otter (Lutra lutra) remains extremely widespread in the lakes, rivers, and coastal waters of Europe, Asia, and northern Africa. Nevertheless, the species has disappeared from many areas due mainly to chemical pollution in the water which is then absorbed by fish. The otters, which prey chiefly on fish, rapidly build up a fatal dosage of poisonous compounds. The species has recovered in many areas where water quality has improved, but continues to be threatened in others. The sable (Martes zibellina) is a type of mustelid that historically ranged throughout the forests of Eurasia. Long hunted for its highly valued fur, the species has been extirpated from Europe but can still be found from the Urals to northern Japan, and is not currently considered to be threatened. The long-fingered mouse-eared bat (Myotis capaccinii) is widespread across the Mediterranean region and the Middle East, but highly dependent upon wetlands for hunting and caves for roosting. While loss of habitat is the main threat, the species is additionally collected in northern Africa for use in ‘traditional medicine’. Felten’s mouse-eared bat (M. punicus) is found from Morocco to western Libya, with isolated populations on Corsica and Sardinia. It is threatened in northwestern Africa by destruction of its cave roosts by fire and vandalism, and by overcollection for ‘traditional medicine’. Mehely’s horseshoe bat (Rhinolophus mehelyi) is found discontinuously around the Mediterranean Sea from northwestern Africa and the Iberian Peninsula through the Balearic Islands, southern France, Sardinia, Sicily, the Balkan Peninsula, and Asia Minor. A cave-roosting species, it is vulnerable to disturbance and destruction of its large colonies. Steller’s sea eagle (Haliaeetus pelagicus) breeds on the Kamchatka Peninsula and in the coastal area around the Sea of Okhotsk, as well as on the lower reaches of the Amur River and on northern Sakhalin. The majority winter in the southern Kuril Islands and on Hokkaido. In 2012 the total population was estimated at between 4600 and 5100 and continues to decline due to habitat degradation, pollution, poisoning, and overfishing. Pallas’ fish-eagle (H. leucoryphus) has a peculiar, essentially land-locked distribution for a sea eagle, being found sporadically in wetland areas throughout central and southern Asia. The total population, estimated at less than 2500, is threatened by human persecution and loss of habitat. The eastern imperial eagle (Aquila heliaca) has an extensive distribution, breeding in south-eastern Europe and through western and central Asia, from where many

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The Palearctic Realm

populations migrate in winter to north-eastern Africa, the Middle East, and southern Asia. The total population is small, however, and sensitive to human influence. The steppe eagle (A. nipalensis) has a similarly large distribution, breeding across much of central Asia and migrating at other times over much of Africa and southern Asia. It has undergone particularly rapid declines in its European range and has been extirpated from many areas. The greater spotted eagle (Clanga clanga) is an extremely widespread if nevertheless rare Palearctic species that favours boreal forests near wetlands for breeding, at other times migrating as far south as North Africa and South East Asia. It is threatened mainly by hybridization with lesser spotted eagles (C. pomarina) and habitat destruction. The Egyptian vulture (Neophron percnopterus) is a small species still found over a wide area of southern Eurasia, Africa, and south-western Asia, where it is divided into three subspecies. The nominate form (N. p. percnopterus) is found from southern Europe and northern Africa to north-western India, where it has suffered significant declines due to poisoning, habitat destruction and disturbance, and collisions with wind turbines. The osprey (Pandion haliaetus) is a large, fish-eating hawk with an unusual, nearly global distribution. The Palearctic osprey (P. h. haliatus) is still found over much of Europe, Africa, and Asia. During the late nineteenth and early twentieth centuries it was threatened by egg collectors and hunting. Later, during the 1950s and 1960s, it underwent a precipitous decline due to the toxic effects of insecticides such as DDT on its reproduction. It recovered quickly in many countries after the banning of DDT in the early 1970s, and is no longer considered threatened. The saker falcon (Falco cherrug) breeds from central Europe east to Manchuria, migrating from there to Ethiopia, the Arabian Peninsula, northern Pakistan, and western China. It has declined rapidly in recent years, particularly in central Asia, mainly due to habitat destruction and illegal capture for use in Arab falconry. The peregrine falcon (F. peregrinus) is notable for being the fastest bird in the world and, indeed, the fastest member of the animal kingdom. Like the osprey, it too is found nearly globally, but was at one time threatened by pesticide spraying. It is no longer considered threatened. The great bustard (Otis tarda), one of Eurasia’s most spectacular birds, was originally a species of the steppes but has adapted somewhat to agricultural landscapes. Nevertheless, it has it has suffered serious declines owing to habitat fragmentation and hunting. The Asian houbara bustard (Chlamydotis macqueenii) is found in desert and steppe regions from east of the Sinai Peninsula to Mongolia. Ruthlessly hunted across its wide range, it was considered great sport in colonial India in particular. However, it was not until the introduction of modern firearms and jeeps that the major population declines really got underway. In 1971, for example, one hunting party alone killed 2000 of the birds in Pakistan. The species was very nearly

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driven to extinction in the Middle East and other areas, and although better protected now both legal hunting as well as poaching continues in the Arab world, where the meat is unfortunately considered to be an aphrodisiac. The red-crowned crane (Grus japonensis) is divided into two separate populations. The first breeds in south-eastern Russia, north-eastern China, and Mongolia, and winters mainly in the Yellow River delta, coastal China, and in the Korean demilitarized zone. The second lives and breeds in eastern Hokkaido, Japan, and is non-migratory. The latter population is stable or slightly increasing, while the continental one is severely declining owing to loss and degradation of wetlands through conversion to agriculture and industrial development. The total world population is estimated at around 3000. The black-necked crane (G. nigricollis) is a medium-sized species that breeds among the alpine bog meadows of the Tibetan Plateau and remote parts of Ladakh, India, with some populations wintering in Bhutan. The total population, around 10,000, is threatened mainly by loss of habitat. The white-naped crane (Antigone vipio) breeds in far south-eastern Russia, Mongolia, and northern China, from where it migrates to eastern China, the Korean Peninsula, and southern Japan. It has undergone a considerable decline due to loss of wetlands. The oriental white stork (Ciconia boyciana) was once common in eastern Siberia, China, the Korean Peninsula, and Japan, but declined to the point of extinction by the mid-twentieth century due to loss of habitat and heavy hunting. Today it breeds mainly in the Amur and Ussuri river drainages along the Russia/China border, with smaller numbers in the lower reaches of the Wuyuerhe River in Heilongjiangg province. The main wintering grounds are in the lower Yangtze drainage and in southern China as far south as Taiwan and Hong Kong. Small numbers are still found in the Koreas and Japan and irregularly in the Philippines, northeastern India, Myanmar, and Bangladesh. The total population is thought to be around 3000. The northern bald ibis or waldrapp (Geronticus eremita) is a large, distinctive species that was historically widespread across the Middle East, northern Africa, and southern and central Europe. Fossil material has been found dating as far back as 1.8 million years, and the ancient Egyptians were sufficiently familiar with the bird to use it as a hieroglyph, so it must have been common in Egypt some thousands of years ago. Long persecuted by humans in their cliff-side breeding colonies, the species has also declined owing to a number of other factors including loss of habitat and pesticide poisoning. It disappeared from Europe over three centuries ago, and since the beginning of the twentieth century has been confined to disjunct populations that may ultimately be revealed to be distinct subspecies: a western one in North Africa, and an eastern one in the Middle East. In North Africa the species had colonies throughout the Atlas Mountains, from where it formerly migrated each year to Europe. It disappeared from

The Palearctic Realm

Algeria in the 1980s, but still has a stronghold in southern Morocco, with three breeding subcolonies in Souss-Massa National Park and another major one at nearby Tamri, totalling around 500 in all. The eastern population bred in Syria, where it was described as still fairly common as recently as the 1980s. From there it migrated south through Jordan, Saudi Arabia, and Yemen, wintered in central Ethiopia, then migrated back to Syria through Eritrea, Sudan, Saudi Arabia, and Jordan. Long believed to have died out; in 2002, however, a tiny colony consisting of just seven individuals was rediscovered at Palmyra. Unfortunately, as of 2015 it appears that these birds, too, have been extirpated. A semi-wild population numbering around 100 still exists at Birecik, southern Turkey. The species has long been established in captivity, and limited reintroduction programmes have been attempted at sites in Austria, Spain, and Morocco. The Asian crested ibis (Nipponia nippon) historically nested in the Russian Far East, Japan, and mainland China, and was a non-breeding visitor to the Korean peninsula and Taiwan. It is now extinct over virtually all of its former range. There is a fairly successful captive breeding programme in Asia, but the only known remaining wild populations are in central China (Shaanxi), along with a reintroduced one on Sado Island in Japan. The Chinese egret (Egretta eulophotes) breeds on small islands off the coasts of far-eastern Russia, the Korean Peninsula, and mainland China, and winters among the shallow tidal estuaries, mudflats, and bays of South East Asia. The species was almost brought to extinction during the nineteenth century by plume hunters. Despite this the persecution continued, and by the 1960s there were only scattered reports of its existence, mainly on the coast of the Yellow Sea. In recent years numbers have stabilized at between 2600 and 3400, although since the mid-1980s all breeding records have been from small, uninhabited offshore islands. Saunder’s gull (Saundersilarus saundersi) breeds mainly in eastern coastal China and sporadically at various sites on the south-western coast of South Korea, from where it migrates to eastern and southern China, Taiwan, western Japan, and Vietnam. It is threatened by the loss of tidal flats and salt marshes due to coastal development. The black-faced spoonbill (Platalea minor) is a type of wading bird that breeds only on islets off the western coast of the Korean Peninsula and north-eastern China, from where they winter in parts of southern China (Taiwan, Macau, and Hong Kong), Japan, Vietnam, Cambodia, Thailand, and the Philippines. The species has long been rare and declining due to habitat destruction and pollution, reaching an all-time population low in the 1990s, but has since recovered somewhat. One of the most shameful examples of mindless extermination is that of the great auk (Pinguinus impennis). This flightless, penguin-like bird at one time occurred widely in massive breeding colonies on the islands of the North Atlantic as well as on the western European, north-western

African and eastern North American coasts. First discovered in 1534, its persecution began almost immediately. Vast numbers were slaughtered by ship’s crews who drove them into stone pens where they were killed, or directly to their ships, where they were cooked to extract the fat bird’s oily substances. Many ships also used the birds as fuel under pots in which other auks were being cooked. Finally, the nestlings were used as bait in fishing and eggs were collected for food. When the species became rare, museums and private collections hurried to get their share of any remaining specimens and eggs. The last two birds were killed on Eldey Island off Iceland in 1844. Two sailors had found a nesting pair, and proceeded to strangle them before smashing the last egg with a boot. The swan goose (Anser cygnoid) has its key breeding grounds in south-eastern Russia, Mongolia, and northernmost China, with virtually the entire population wintering in the Yangtze floodplain of east-central China. While uncommon in the wild state the species has been domesticated, with introduced and feral populations occurring in many areas outside its natural range.

Figure 2.1 Great auk (mounted specimen). (Credit: Hulton Archive/Stringer/ Getty Images.)

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The Palearctic Realm

The white-headed duck (Oxyura leucocephala) breeds primarily in Russia and Kazakhstan as well as in smaller, more isolated areas of Spain, Algeria, Tunisia, and central Asia, from where it winters in the Middle East and south-central Asia. In the early twentieth century the total population likely exceeded 100,000, although this fell to an estimated 20,000 by 1996. The primary threat is competition and hybridization with introduced ruddy ducks (O. jamaicensis). The velvet scoter (Melanitta fusca) is a type of duck that breeds in Scandinavia and western Siberia, from where it migrates to southern Europe and the Middle East. It has undergone considerable declines in recent decades, the reasons for which remain unclear. The northern pochard (Aythya ferina) is a type of diving duck that still breeds across much of Eurasia, from where it winters in southern Asia and northern Africa. Another species, Baer’s pochard (A. baeri), breeds in south-eastern Russia and northern China, from where it winters in southern Asia. Both are threatened by habitat destruction, hunting, the overcollection of eggs, and other factors. The scaly-sided merganser (Mergus squamatus) is a type of sea duck that inhabits a variety of habitats in extreme southeastern Russia, the Korean Peninsula, and north-eastern China, with most wintering in central and southern China. The species began to decline in the 1960s and 1970s due to deforestation, although in more recent decades illegal hunting has become the primary threat. The marbled teal (Marmaronetta angustirostris) is a partially migrant species still found patchily across much of southern Europe, northern Africa, and western and central Asia, but has been significantly reduced due to loss of habitat and hunting. The crested shelduck (Tadorna cristata) is known only from three museum specimens, one collected in Russia and the other two in South Korea. It was presumably familiar in Japan, as it was drawn by artists there during the nineteenth century. It was thought to be long extinct when a male and two females were sighted on islands south of Vladivostok in 1964. A further sighting of two males and four females was claimed in North Korea in 1971, although this seems unlikely. More recently there have been several unconfirmed reports from north-eastern China, leading to the theory that the species may breed in remote mountainous areas far inland, only travelling to the coasts at other times. In any case, if it still survives the total population must be quite small. The horned grebe (Podiceps auritus) as a species is found over a wide area of the temperate Eurasia and North America. The Eurasian horned grebe (P. a. auritus) is everywhere declining due to the effects of human disturbance, loss of habitat due to deforestation around breeding lakes, and other factors. The slender-billed curlew (Numenius tenuirostris) is only known to breed in a small area of south-central Russia, from where it winters in a few areas of northern Africa and the Arabian Peninsula. Last recorded in 2004, it may possibly be extinct.

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The sociable lapwing (Vanellus gregarius) is a type of wading bird that breeds on the open grasslands of Russia and Kazakhstan, from where it migrates to certain key wintering sites in Israel, Syria, Eritrea, Sudan, and north-western India. It has undergone a serious decline since the mid-nineteenth century from hunting pressure and other, as yet poorly understood reasons. The European turtledove (Streptopelia turtur) breeds over a wide area of the south-western Palearctic region, from where it migrates to sub-Saharan Africa to winter. Unfortunately, it has everywhere undergone a serious decline owing to a number of factors, including loss of foraging and nesting sites, disease, and hunting along its migration routes. Four subspecies are recognized. The northern turtledove (S. t. turtur) occurs from Europe (including the Madeira and Canary Islands) to western Siberia. The yellow-breasted bunting (Emberiza aureola) was, historically, one of the most abundant passerine birds in Eurasia, breeding from northern and eastern Europe through Siberia, Kazakhstan, northern China and Mongolia to Far Eastern Russia, Korea, and northern Japan. During the autumn the birds would stop over in large numbers in the Yangtze Valley before continuing on to their wintering grounds in South and South East Asia. The species began to undergo a drastic decline during the early 1990s, most likely due to overcollection for food by mist net during migration, and has since disappeared from most or all of Finland, Belarus, Ukraine, and large areas of western Russia. The large-billed reed warbler (Acrocephalus orinus), described as ‘the world’s least known bird’, was long known only from a single specimen collected in north-western India (Himachal Pradesh) in 1867. The species was rediscovered in Thailand in 2006, and has since also been found in northeastern Afghanistan, Tajikistan, and Bangladesh. It appears to be a rare long-distance migrant that breeds within the Palearctic Realm and winters in southern Asia. The aquatic warbler (A. paludicola) has a highly fragmented breeding range in the mires and marshes of Poland, Belarus, Ukraine, Germany, Lithuania, and western Russia, from where it migrates to western Europe and north-western Africa. The species suffered major declines in the second half of the nineteenth century due to habitat destruction, and further extirpations continue. Pleske’s grasshopper-warbler (Locustella pleskei) is a rare species that breeds on small islands in Peter the Great Bay of far-eastern Russia, the Izu Islands, and islands off Kyushu, Japan, the Korean Peninsula, and eastern China, from where it winters in southern China and Vietnam. It is threatened by habitat destruction. Tristram’s white-bellied woodpecker (Dryocopus javensis richardsi) is a large and spectacular subspecies historically found across the Korean Peninsula and on the Japanese island of Tsushima. It was extirpated from the latter due to intensive hunting and collection by museums, and became rare in Korea owing to deforestation. Despite being legally protected since

The Eurasian Region

1952 it disappeared from South Korea by 1978, and today fewer than 50 still survive in a few areas of montane forest in North Korea. The spur-thighed tortoise (Testudo graeca) occurs in a wide area of southern Europe, North Africa, and southwestern Asia, in a varying number of subspecies. It is everywhere threatened by collection for use as pets. Lataste’s viper (Vipera latastei) is found patchily in both the Iberian Peninsula and north-western Africa, in a wide range of habitats and altitudes. Sticklebacks (Pungitius) are freshwater, brackish or marine fish related to seahorses. The Amur stickleback (P. sinensis) remains relatively widespread through eastern Asia, but has disappeared from many areas due to pollution and the construction of flood control and dams, which can change the velocity of rivers and render them unsuitable for the species. The short-spined ninespine stickleback (P. tymensis) is confined to Sakhalin, the Kuril Islands, and Hokkaido (Japan), where it is threatened by loss of habitat and invasive species.

The Eurasian Region The Eurasian Region spans half the Earth, including as it does virtually all of Scandinavia and Siberia (including the Kamchatka Peninsula, the Korean Peninsula, and islands of the northern Pacific), all of Europe (including most of the Mediterranean islands and the Anatolian and Balkan peninsulas), and much of central and western Asia. Although usually divided into two continents, Europe and Asia, this immense area is in fact a single physical complex that formed between 375 and 325 million years ago with the merging of three main landmasses. This in turn would be joined for a time to Laurentia (now North America) to form Euroamerica. Eurasia features a tremendous variety of topography, from Lake Baikal (the world’s deepest lake) to some of the world’s highest mountains. Its zones of climate range from subarctic through temperate to subtropical, and its vegetation from coniferous forest in the north through deciduous woodland, steppe, desert, and the Mediterranean maquis scrub in the central and south-western parts, to subtropical rainforests in the south-eastern parts. The great taiga is the world’s largest forest, extending from the Scandinavian Mountains in the west to the Pacific Ocean in the east. Also, the steppes of Eurasia are larger than any other similar region. This extraordinary range of habitats within one continental block makes Eurasia very rich in animal species evolved over eons. The fact that this largest of continents was connected with Africa and America for long periods has also enriched the Eurasian fauna through animal migrations. The history of the European wood bison or wisent (Bison bonasus) is almost as dramatic as that of its more famous American counterpart. A huge forest-dwelling animal, its disappearance is intimately connected with the retreat of the Eurasian deciduous forests. In prehistoric times it occurred

over almost the whole of Europe east to the Lena and possibly existed in ancient Assyria, Mesopotamia, and Persia as well, which at that time were forested. Three subspecies are recognized, only one of which, the European lowland wood bison (B. b. bonasus), survives today. By the early twentieth century the latter had been reduced to a single herd of about 700 animals in the Bialowieza Forest of Poland, where they had long been protected. Because many of the animals there were tame, they became easy prey for invading troops during the First World War, who reduced the population to about 150. The last free individuals were shot in Bialowieza in 1921. Fortunately, some were preserved in zoos and their numbers were gradually built up again. But after World War II only 16 had survived in captivity at Bialowieza. They increased and in 1952 some were released back into the wild, the first to roam freely in one of Europe’s last virgin forests for over 30 years. By the close of the 1960s there were over 1000 pure European bison, of which more than 215 lived in freedom in both Bialowieza and in the Zverevskoye Forest of north-western Ukraine. Since then they have been introduced into protected areas throughout western and eastern Europe with varying degrees of success, where today the total population is around 1500. There is a similar number in captivity worldwide. The elk or wapiti (Cervus canadensis) is one of the largest species of deer. A number of subspecies are found across North America and north-eastern Asia, a few of which are threatened. The reindeer (Rangifer tarandus), previously discussed in this volume, is a type of deer with a circumpolar distribution that has adapted itself to a number of different environments. While not threatened as a species a number of Eurasian subspecies are, and will be discussed below. The Caspian tiger (Panthera tigris virgata) had at one time a wide range in the sparse forests south and east of the Black and Caspian seas, although the Pontic-Caspian steppe and into the deserts of central Asia. By the late 1960s intensive hunting and habitat destruction had wiped them out in most of these areas, and only a handful survived. It was last recorded in the early 1970s, and is now considered extinct. The Siberian or Amur tiger (P. t. altaica), the largest of all the big cats, was formerly found throughout eastern Siberia, Mongolia, Manchuria, and the wooded parts of northern China, where it was adapted to the deep winter snows. Today it is confined to the Amur-Ussuri region in far-eastern Siberia, with the exception of a small population in Hunchun National Siberian Tiger Nature Reserve in north-eastern China, near the border with North Korea. In 2015 there was an estimated population of 480–540 in the Russian Far East. The Amur leopard (Panthera pardus orientalis) is confined to cold regions of the Russian Far East and north-eastern China, having disappeared from its former range on the Korean Peninsula. Among the world’s rarest animals, the total population is thought to be less than 100. The Eurasian lynx (Lynx lynx) is found across most of Europe and Asia and is not considered threatened as a species,

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The Palearctic Realm Figure 2.2 Caspian tiger in the Berlin Zoo, 1899. (Credit: Berlin Zoo.)

although many populations, particularly in southern areas, are isolated and at risk. The European wildcat (Felis silvestris silvestris) inhabits forests across continental Europe from the Iberian Peninsula to Turkey and the Caucasus, but has become uncommon everywhere. A small number remain in the Scottish Highlands, but the species has otherwise been extirpated from the British Isles. While legally protected, they are frequently mistaken for feral cats and shot, and interbreeding with domestic cats is an additional threat. The brown bear (Ursus arctos) is, as a species, found across much of North America and northern Eurasia, where it has manifested into a number of subspecies. A few will be discussed below. The Mongolian marmot (Marmota sibirica), as a species, is found in a variety of habitats in east-central Asia. The steppe Mongolian marmot (M. s. sibirica) occurs in southern Russia, Mongolia, and northern China, while the montane Mongolian marmot (M. s. caliginosus) occurs in northern, western, and central Mongolia. Both are threatened by loss of habitat and hunting for their meat and pelts. The Spanish imperial eagle (Aquila adalberti) is, at least in terms of breeding, entirely endemic to montane and lowland woodlands in the Iberian Peninsula of central and southwestern Spain and adjacent parts of Portugal. Resident, nonbreeding populations were at least historically to be found in Morocco and perhaps elsewhere in North Africa, although the species now appears to be largely a vagrant in these areas. It has recovered from a low of only 30 breeding pairs in the 1960s to an estimated 486 pairs by 2016, but remains particularly

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vulnerable to electrocution by poorly insulated powerlines, as well as by loss of habitat and persecution. Blakiston’s eagle-owl (Bubo blakistoni) is the largest living owl. There are two recognized subspecies, both of which are threatened by loss of their preferred riverine forest habitat. The mainland subspecies (B. b. doerriesi) is found in the forests and coastal mountain ranges of eastern Siberia, eastern China, and the Korean border area, including Sakhalin Island (although there has not been a verified record from there since 1974). The hooded crane (Grus monacha) breeds in south-central and south-eastern Siberia, a small area of north-eastern China (Heilongjiang), and perhaps Mongolia. The majority of the population, estimated in 2015 at roughly 15,000, winters in southern Japan, with smaller numbers in south-eastern China and South Korea. The yellow-eyed pigeon (Columba eversmanni) breeds in southern Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, Kyrgyzstan, Afghanistan, north-eastern Iran, and extreme north-western China, from where it migrates to Pakistan and north-western India. During the nineteenth and early twentieth centuries huge flocks were reported in its wintering grounds, particularly in the Punjab. The species declined rapidly thereafter due to loss of habitat and intensive hunting. The rustic bunting (Emberiza rustica) has a massive distribution, breeding across the Palearctic from Norway to Kamchatka, from where it migrates south in winter to central and eastern Asia. Nevertheless, in recent decades the global population has declined dramatically, most likely due to increased logging in its breeding range and to large-scale

The Eurasian Region

trapping combined with increasing agriculture in its nonbreeding range. The streaked reed warbler (Acrocephalus sorghophilus) is known from a few records originating within a small area of north-eastern China, and Taiwan, from where the species (at least historically) wintered on Luzon in the Philippines. Its breeding range remains unknown. The white-browed reed warbler (A. tangorum) breeds in south-eastern Russia and north-eastern China, from where it winters in southern Laos, Cambodia, Thailand, and Peninsular Malaysia. The species is everywhere threatened by loss of its swampy grassland habitat. Carbonell’s wall lizard (Podarcis carbonelli) is a rare species whose nominate subspecies (P. c. carbonelli) occurs in highly fragmented populations in western and central Portugal, extending slightly into west-central and southwestern Spain. The Iberian Lataste’s viper (Vipera latastei latastei) occurs in both Spain and Portugal south of the Pyrenees, where it is threatened due to loss of habitat and direct persecution by humans.

Mountains and Highlands With few exceptions, the great mountain chains of Eurasia run from west to east across the two continents. Many of them form massive barriers between well-distinguished biogeographic regions. They also serve as enormous refuges for plants and animals that have been exterminated elsewhere, or have been driven by advancing cultivation into remote montane areas. The Balkan lynx (Lynx lynx balcanicus) is confined to the mountainous regions of eastern Albania and western North Macedonia, with smaller populations in Kosovo and Montenegro. Long threatened by illegal hunting, the total number is thought to be less than 50. The mouflon (Ovis gmelini) and its many subspecies are a group wild sheep and the ancestors of all domestic sheep (O. aries). Today it inhabits steep mountainous woodlands of the Near and Middle East, although historically the range extended further to the Crimean Peninsula and the Balkans. Populations were long ago introduced to a number of islands in the Mediterranean, perhaps as feral domesticated animals, where they have naturalized to the hilly interiors and given rise to arguably distinct forms. Those from the islands of Corsica and Sardinia are commonly known as the European mouflon (O. g. musimon), having been subsequently introduced to parts of mainland Europe as well. Like all mouflon this subspecies has been heavily depleted by hunting for its meat and horns. The Transcaspian mouflon (O. g. arkal) is found in western Kazakhstan, Turkmenistan, Uzbekistan, and north-eastern Iran. In the mid-1970s the total population was estimated at over 20,000, of which around 15,000 lived in Golestan National Park in Iran. The Bukhara mouflon (O. g. bocharensis) is found in Tajikistan, Turkmenistan, and Uzbekistan. The Afghan mouflon (O. g. cycloceros) is found in southern

Turkmenistan, eastern Iran, Afghanistan, and northern Pakistan. The Armenian mouflon (O. g. gmelini) is found in north-western Iran, Armenia, and Azerbaijan. Another type of wild sheep is the argali (Ovis ammon). Several subspecies are found in the mountains and deserts of central Asia, where they are heavily hunted. The North China argali (O. a. jubata) is confined to a few pockets in northern China. Severtzov’s argali (O. a. severtzovi) is known only from a small area of southern Uzbekistan. The Karaganda argali (O. a. collium) is confined to the low hills of central and eastern Kazakhstan. The Gobi argali (O. a. darwini) is confined to a few areas of northern China and Mongolia. The markhor (Capra falconeri) is a large, typically screwhorned wild goat found patchily in the mountains of westcentral Asia. Several subspecies are threatened by loss of habitat, domestic grazing herds and intensive hunting. The Astore markhor (C. f. falconeri) is confined to parts of northeastern Afghanistan (Kunar and Nuristan), northern Pakistan, and northern India (Jammu and Kashmir). The Bukharan markhor (C. f. heptneri) occurs in parts of north-eastern Afghanistan, southern Tajikistan, Turkmenistan, and southern Uzbekistan. The straight-horned markhor (C. f. megaceros) survives in parts of north-eastern Afghanistan (Kabul, Parwan, and Paktia provinces) and central Pakistan (Baluchistan and Punjab provinces). The Iberian ibex (Capra pyrenaica) is another species of wild goat that historically occurred throughout the mountainous areas of the Iberian Peninsula and south-western France. Of the four described subspecies, two are now extinct. The Portuguese ibex (C. p. lusitanica) was still abundant at the end of the eighteenth century, ranging in all the north-western mountains of Portugal, Galicia, Asturias, and western Cantabria. Thereafter its decline was rapid as hunting pressure increased for its meat, hides, horns, and bezoar stones, which local people erroneously believed to have medicinal properties. By 1870 it was rare. The last herd, about a dozen animals, was reported in 1886. An old female was captured alive in 1889, but only survived three days. Two more were found dead the following year in Galicia, victims of an avalanche. The last known individual in Spain died in 1890, and the last sighting was in the Serra do Gerês, Portugal, in 1892. A mounted specimen was on display in the Bocage Museum, Lisbon, until destroyed by fire in 1978. The Pyrenean ibex (C. p. pyrenaica) was once abundant on both sides of the Pyrenees as well as in the Cantabrian Mountains of southern France. It too was severely persecuted by hunters and decreased rapidly during the nineteenth and twentieth centuries. Competition with domesticated sheep, goats, cattle, and horses also contributed to the decline. It was long thought to have gone extinct in the 1910s, until a small population of around 20 was discovered in Ordesa y Monte Perdido National Park, in the Spanish central Pyrenees. By 1989 only a dozen or so were left. The last individual, a female named ‘Celia’, was bizarrely killed by a fallen tree on 6 January 2000. In recent years there have been attempts made

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The Palearctic Realm

Figure 2.3 A nineteenth-century photo of a Portuguese ibex (Capra pyrenaica lusitanica). (Credit: unknown.)

to clone this individual and thereby make the Pyrenean ibex ‘unextinct’, although it would seem that modern scientific expertise cannot overcome the lack of a male cell donor. Two species of chamois (Rupicapra), a kind of goatantelope, inhabit the mountains of Eurasia. The southern chamois (R. pyrenaica) is divided into three subspecies inhabiting south-western Europe. The Pyrenean chamois (R. p. pyrenaica) and Cantabrian chamois (R. p. parva) were both nearly hunted to extinction by the 1940s, but have since recovered their numbers and are no longer considered threatened. A third remains highly endangered and is discussed elsewhere in this book. The northern chamois (R. rupicapra) is, as a species, common and widespread in the mountains of central and southern Europe and Asia Minor. However, several subspecies are considered to be highly threatened. The Balkan chamois (R. r. balcanica) inhabits most of the mountainous regions of Albania, as well as Bulgaria’s four main massifs. In Greece it is confined to a few widely scattered populations. The gorals (Naemorhedus) are a group of small goatantelopes confined to the mountainous regions of Asia, where they have been much reduced by hunting. The long-tailed goral (N. caudatus) is found patchily in eastern Russia (Primorsky and Khabarovsk territories) and north-eastern China, with a small population living in the Demilitarized Zone on the Korean Peninsula. The Eurasian musk deer (Moschus moschiferus) is still found over a wide area of north-eastern Asia in suitable montane taiga habitat, but has been much reduced by hunting. The

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Siberian musk deer (M. m. moschiferus) occurs in the Russian Far East, Mongolia, and northern China (Xinjiang, Heilongjiang, and Inner Mongolia). Turov’s musk deer (M. m. turovi) is confined to an area of the Russian Far East. The Verkhoyansk Ridge musk deer (M. m. arcticus) is confined to the Verkhoyansk Range in the East Siberian Mountains. The Korean musk deer (M. m. parvipes) is found in north-western China and the Korean Peninsula. Pikas (Ochotona) are small, mountain-dwelling relatives of rabbits. Hoffmann’s pika (O. hoffmanni) is known only from two, widely separated localities (the Bayan-Ulan Ridge in northern Mongolia and the Erman Mountains of south-eastern Russia). The Korean pika (O. coreana) is confined to the mountains of north-eastern China (Jilin) and north-eastern North Korea. The woolly dormouse (Dryomys laniger) is largely confined to the Taurus Mountains of southern Anatolia, Turkey, along with a few isolated localities in north-eastern Anatolia. The Balkan snow vole (Dinaromys bogdanovi) is a ‘living fossil’ from isolated, mountainous regions of the Balkans, where it may be threatened by competition with the European snow vole (Chionomys nivalis). Schaub’s mouse-eared bat (Myotis schaubi) is a rare species known only from a few localities within the southern Caucasus Mountains of Armenia, and from the north-western Zagros Mountains of Iran. Fossils indicate that it historically ranged in Russia and Hungary as well. The Iberian rock lizard (Iberolacerta monticola) is found patchily in the highlands of north-western Spain and central Portugal. The Spanish keeled lizard (Algyroides marchi) is confined to a few isolated localities in the Alcaraz, Cazorla, and Segura ranges of south-eastern Spain. Darevsky’s viper (Vipera darevskii) is known only from Armenia and possibly two localities in eastern Turkey. The Kurdistan newt (Neurergus derjugini) is known only from a small area of the Avroman Mountains in western Iran (Kermanshah province), but may occur in adjacent northeastern Iraq as well. Its breeding streams have been seriously impacted by drought, water extraction and pollution, and the species is still collected for the international pet trade. The Urmia newt (N. crocatus) is confined to the mountains west of Lake Urmia in northern Iraq, south-eastern Turkey, and (at least historically) north-western Iran. It is threatened mainly by dam construction. The Anatolian newt (N. strauchii) is divided into two subspecies found in the mountains of eastern Turkey. Strauch’s newt (N. s. strauchii) is confined to streams south and west of Lake Van, while Baran’s newt (N. s. barani) is known from a few areas near the town of Pütürge. The golden-striped salamander (Chioglossa lusitanica) is confined to the highlands of north-western Spain and northern and central Portugal, where it is dependent upon streams. Two subspecies have been described, the Lusitanian golden-striped salamander (C. l. lusitanica) and the long-footed goldenstriped salamander (C. l. longipes).

The Eurasian Region

The Caucasian salamander (Mertensiella caucasica) is found in north-western Anatolia (Turkey) and western Georgia, where it is threatened by loss of habitat. Bejara’s fire salamander (Salamandra salamandra bejarae) is confined to the mountains of north-central Spain.

The Alps The Alps are the highest and most extensive mountain range lying entirely within Europe, stretching approximately 1200 km across eight countries (France, Switzerland, Italy, Monaco, Liechtenstein, Austria, Germany, and Slovenia). The mountains were formed over tens of millions of years as the African and Eurasian tectonic plates collided. At 4810 m Mont Blanc, which spans the French–Italian border, is the highest, but the alpine region contains around a hundred peaks higher than 4000 m. This extreme altitude and size greatly affect the climate of Europe. Generally speaking, they can be divided into two main subranges, the Eastern and Western Alps, separated in eastern Switzerland near the Splügen Pass. A series of lower ranges run parallel to the main chain of the Alps, including the French Prealps and the Jura Mountains. The alpine ibex (Capra ibex) had been driven to the point of extinction in the early nineteenth century, but was saved thanks to conservation efforts chiefly by the Italians. The species is now considered safe in Italy, with populations reintroduced to Austria, France, Germany, and Switzerland and new ones established in Bulgaria and Slovenia. Two subspecies of northern chamois (Rupicapra rupicapra), previously mentioned in the general section, are found within in this region. The alpine chamois (R. r. rupicapra) occurs in the Alps of Austria, Germany, and eastern France, where it is threatened mainly by poaching and overhunting. The Chartreuse chamois (R. r. cartusiana) is confined to the Chartreuse limestone massif near Grenoble, on the western edge of the French Alps. The Bavarian pine vole (Microtus bavaricus) is known only from Rofan Mountain, in the northern Tyrol of Austria. Lanza’s alpine salamander (Salamandra lanzai) is confined to a small area of the Western Alps on the border between Italy and France.

Mountains of Central Asia The mountains of Central Asia, as here defined, are those ranges bordering the Tibetan Plateau to the north and west, spanning parts of Afghanistan, western China, Kazakhstan, Kyrgyzstan, Tajikistan, and Uzbekistan. Together they provide a diversity of habitats including montane grasslands, meadows, temperate coniferous forests, and alpine tundra. The northern wild dog (Cuon alpinus hesperius) is known from the Altai and Tian Shan mountains, possibly extending to the Pamir Mountains and Kashmir. It was historically much more widespread. Two subspecies of Tarbagan marmot (Marmota sibirica) are found in high-elevation steppe and alpine meadow areas of Central Asia. The Khentii marmot (M. s. sibirica) is confined

to the Khentii Mountains of north-eastern Mongolia (Tuv and Khentii provinces). The Mongolian marmot (S. s. caliginosus) is still relatively widespread in south-western Russia (southwestern Siberia, Tuva, and Transbaikalia), north-eastern China (Inner Mongolia and Heilongiiang), and western Mongolia, but is everywhere suffering massive declines. The white-throated bushchat (Saxicola insignis) is a type of passerine bird that breeds very locally in the mountains of Mongolia and adjacent areas of south-central Russia, from where it migrates to the grasslands of northern India and Nepal. Formerly common, it is now rare in its wintering range due to loss of habitat. The Turkestanian salamander (Hynobius turkestanicus) is known only from a few specimens collected in 1909 from an imprecise locality in central Asia, but thought to be somewhere in eastern Uzbekistan or southern Kyrgyzstan. The Tian Shan Mountains The Tian Shan Mountains is a large system of central Asian mountain ranges located in north-western China (Xinjiang), Kazakhstan, Kyrgyzstan, and Uzbekistan. The Tian Shan argali (Ovis ammon karelini) is a type of wild sheep that, as here defined, is confined to the Tian Shan Mountains. Menzbier’s marmot (Marmota menzbieri) inhabits highelevation meadows and steppe within the western Tian Shan Mountains, where its range is divided into two separate pockets separated by more than 100 km. It is threatened by habitat loss due to agriculture. The Ili pika (Ochotona iliensis) is confined to a few highelevation areas on two spurs of the Tian Shan Mountains in China, where the total population is thought to be less than 1000. The Central Asian salamander (Ranodon sibiricus) is a rare species confined to the Dzungarian Alatau Ridge on the border between China and Kazakhstan. The range is extremely fragmented due to the scarcity of suitable habitat (i.e. flat, high-elevation plateaus with a dense network of permanent streams). The Karakoram Range The Karakoram is a large mountain range spanning the borders of northern Pakistan, western China, and northeastern India, with the north-western extremity extending into north-eastern Afghanistan and Tajikistan. It includes K2, the second highest mountain in the world. The woolly flying squirrel (Eupetaurus cinereus) was originally known from a few skins collected in extreme northern Pakistan, in the portion of Kashmir under Pakistani control, and from northern Sikkim (India). Thought to be extinct for 70 years, it was rediscovered in small numbers in 2004. The Pamir Mountains The Pamir Mountains are located at the junction of the Himalayas with the Tian Shan, Kunlun, Hindu Kush,

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The Palearctic Realm Figure 2.4 Caucasian wood bison photographed in 1889. (Credit: E. Demidoff.)

Suleman, and Hindu Raj ranges. They are among the world’s highest. The Marco Polo argali (Ovis ammon polii) is a type of wild sheep found mainly in the Pamir Mountains. It is threatened mainly by trophy hunters, who prize its large, spiralling horns. The Alay Mountains Also known as the Alai Mountains, this range runs roughly east to west in Kyrgyzstan and Tajikistan. The Alay mole vole (Ellobius alaicus) is known only from a small area of high-elevation meadow steppe in the Alay Mountains, south Kyrgyzstan. The Altai Mountains The Altai Mountains are located in central and east East Asia where Russia, China, Mongolia, and Kazakhstan come together. The Altai argali (Ovis ammon ammon) is the largest of all wild sheep and possesses the heaviest horns. It is confined to the higher elevations of the Altai Mountains, where not surprisingly it is heavily hunted.

The Caucasus Mountains Located between the Caspian and Black seas, the Caucasus Mountains straddle south-eastern Europe and central Asia. They are comprised of the Greater Caucasus in the north and the Lesser Caucasus in the south. The Caucasian wood bison (Bison bonasus caucasicus) was an inhabitant of the wooded slopes and valley meadows of the Caucasus. It will perhaps never be known how long this subspecies had been isolated from its relative, the European lowland wood bison (B. b. bonasus), which once roamed the forest steppes and woods of lowland Europe. What we know of the

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Caucasian wood bison goes back only as far as 150 years ago. In 1914 there were about 500 such animals in the Caucasus, but persecution drove them higher and higher up into unsuitable habitats, and by 1925 they were reported to be extinct in the wild. However, in 1940, some captive specimens that were at least partly of Caucasian ancestry were brought to a large enclosure on the northern, forested slopes of the Caucasus. In 1954 a small herd was set free and has been strictly protected since then. In the scientific sense, however, the pure Caucasian bison must be considered extinct. The Caucasian lynx (Lynx lynx dinniki) was formerly distributed throughout much of south-western Asia, but is now confined to the northern Caucasus Mountains. The Greater Caucasus The Greater Caucasus Mountains are located in south-western Russia, northern Georgia, and northern Azerbaijan. The West Caucasian tur (Capra caucasica) is a large, heavy-set goat confined to a few localities in the western part of the Greater Caucasus Mountains. The Caucasian chamois (Rupicapra rupicapra caucasica) is confined to the Greater Caucasus. Birch mice (Sicista) are a group of small, jumping rodents. The Caucasian birch mouse (S. caucasica) and the Kazbeg birch mouse (S. kazbegica) are both confined to the Greater Caucasus of Georgia and Russia. The subalpine meadow wall lizard (Darevskia alpina) is confined to a narrow belt of subalpine meadows in the Greater Caucasus of Georgia and Russia. Two species of viper (Vipera) are endemic to the Greater Caucasus, where they are threatened by habitat destruction and degradation, overcollection for the international pet trade,

The Eurasian Region

and human persecution. The magnificent viper (V. magnifica) is confined to a small area of south-western Russia. The Caucasus subalpine viper (V. dinniki) is known from southwestern Russia, northern Georgia and northern Azerbaijan. The Lesser Caucasus The Lesser Caucasus Mountains are located in southern Georgia, Armenia and Azerbaijan. The Armenian birch mouse (Sicista armenica) is known only from a few subalpine meadows in north-central Armenia. The Armenian mouse-eared bat (Myotis hajastanicus) is a rare and possibly extinct species known only from the Lake Sevan basin in Armenia. Rostombekov’s wall lizard (Darevskia rostombekovi) is known from a few isolated subpopulations in northern Armenia and western Azerbaijan.

The Armenian Plateau The Armenian Plateau (also known as the Armenian Highlands) is located to the west of the Anatolian Plateau in Armenia, Azerbaijan, north-western Iran, and eastern Turkey (eastern Anatolia). Uzzell’s wall lizard (Darevskia uzzelli) and the Bendimahi wall lizard (D. bendimahiensis) are both confined to small areas of north-eastern Turkey. Pleske’s racerunner (Eremias pleskei) is a type of lizard confined to sandy, semi-desert areas of the Armenian Plateau. It is seriously threatened by habitat destruction. The Armenian steppe viper (Vipera eriwanensis) is found patchily in north-eastern Turkey, Armenia, and Azerbaijan.

The Pontic Mountains The Pontic Mountains are located in north-eastern Turkey and south-western Georgia. Clarks’ wall lizard (Darevskia clarkorum) is confined to the Pontic Mountains. The Black Sea viper (Vipera pontica) is known only from the Coruh River valley in north-eastern Turkey (Artvin province) and from an additional specimen found in a tea plantation near the Turkish–Georgian border.

The Taurus Mountains The Taurus Mountains are located in southern Turkey, where they separate the Mediterranean coastal region from the central Anatolian Plateau. The Anatolian meadow viper (Vipera anatolica) is known only from a single locality in south-western Turkey. The Taurus frog (Rana holtzi) is confined to a small area of high montane lakes in the Bolkar Range, where it is highly threatened by habitat disturbance and introduced carp.

The Apennine Mountains The Apennines are a range consisting of smaller parallel chains extending some 1200 km along the length of peninsular Italy.

Its system of national parks contains some of the bestpreserved montane forests and grasslands on the continent. The Apennine chamois (Rupicapra pyrenaica ornata) has probably been rare for centuries, its numbers only starting to increase in 1920s as a result of increased protection. They plummeted again to just a few dozen in Abruzzo National Park during World War II, but have slowly recovered since then. Today three small populations, numbering perhaps around 1100, survive in the Abruzzo, Majella, and Gran Sasso-Monti della Laga national parks.

The Pyrenees Mountains The Pyrenees are a range of mountains in south-western Europe forming a natural boundary between Spain and France. Two species of rock lizard (Iberolacerta) endemic to the Pyrenees are threatened by loss of habitat and collection for the international pet trade. The Aran rock lizard (I. aranica) was long thought to be confined to the Mauberme Massif in the central Pyrenees, but in 2006 a new population was discovered in Mont Valier (France). Aurelio’s rock lizard (I. aurelioi) is restricted to a small part of the border area of Andorra, France, and Spain, with the majority of the population living in the latter country. The Pyrenean frog (Rana pyrenaica) is largely confined to the southern slopes of the west-central Pyrenees, where it is threatened by loss of habitat and introduced species.

The Cantabrian Mountains The Cantabrian Mountains stretch east to west for over 300 km across northern Spain. The broom hare (Lepus castroviejoi) is confined to an area of the Cantabrian Mountains, where it occupies a small elevational range of between 1000 and 1900 m. The Cantabrian capercaillie (Tetrao urogallus cantabricus) is a type of grouse that formerly ranged the length of the Cantabrian Mountains, but is now confined to a few areas of north-western Spain. In 2006 the total population was estimated at around 625.

The Carpathian Mountains The second longest mountain range in Europe, the Carpathians form an arc roughly 1500 km in length across central and eastern Europe. The Carpathian wood bison (Bison bonasus hungarorum) historically occurred in the Carpathian Mountains of Moldova and Romania, and perhaps Hungary and Ukraine as well. It began to die out in the eighteenth century due to overhunting, with the last known specimen being shot in 1852. There are currently plans to introduce European wood bison from Poland or the Caucasus into Transylvania. The Carpathian lynx (Lynx lynx carpathicus) is found in the Carpathian Mountains of Romania, Slovakia, and Hungary, where it is threatened by poaching and loss of habitat. A further population was introduced into Croatia, Slovenia, and Bosnia during the 1970s.

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The Palearctic Realm

The Carpathian chamois (Rupicapra rupicapra carpatica) occurs in many populations throughout the Carpathian Mountains and Transylvanian Alps, where there have been a number of successful reintroductions. The Tatra Mountains The Tatra Mountains are a subrange of the Carpathian Mountains forming a natural border between Poland and Slovakia. The Tatra Mountains chamois (Rupicapra rupicapra tatrica) is confined to the Tatra Mountains.

Miscellaneous Mountains and Highlands The Scandinavian Mountains, as their name suggests, are a range that run through the Scandinavian Peninsula. The mountains are ancient and not very high but steep in places, with the western slopes dropping precipitously into the North and Norwegian seas, forming fjords. By contrast, in the northeast they curve gradually towards Finland. The combination of northerly location and ocean moisture has resulted in numerous ice fields and glaciers. Vegetation is typically montane birch forests and grasslands. The mountain reindeer (Rangifer tarandus tarandus) is found only in the montane tundra regions of the Fennoscandian Peninsula, Norway. The Dinaric Alps are located in southern Croatia, northern Albania, Bosnia-Herzegovina, and Montenegro. The Mosor rock lizard (Dinarolacerta mosorensis) is confined to a few isolated populations in the south-western Dinaric Mountains. The Pindus Mountains are located in western Greece and southern Albania. The Greek meadow viper (Vipera graeca) is confined to the subalpine regions of the Hellenides mountain system of southern Albania and central Greece. The Catalan Pre-Coastal Range is located in north-eastern Spain. The Montseny brook newt (Calotriton arnoldi) is confined to the El Montseny Massif, where it is only known in seven mountains streams within the boundaries of El Montseny Natural Park. The Central Cordillera (Sistema Central in Spanish and Portuguese) is located in west-central Spain and east-central Portugal. Cyrn’s rock lizard (Iberolacerta cyreni) and Martinez-Rica’s rock lizard (I. martinezricai) are both confined to the Central Cordillera. The Sierra Nevada Mountains are located in south-eastern Spain. The Betic midwife toad (Alytes dickhilleni) is confined to a few localities within the Sierra Nevada Mountains. The Penibaetic Mountains are located in southern coastal Spain. The long-snouted fire salamander (Salamandra longirostris) is confined to a small area of southern Spain. The Karatau Mountains are located in southern Kazakhstan. The Kara Tau argali (Ovis ammon nigrimontana) is a type of wild sheep confined to the Karatau Mountains. The Helan Mountains (Helan Shan in Chinese, and formerly known as the Alashan Mountains) are an isolated, desert mountain range located in south-eastern Inner Mongolia. The silver pika (Ochotona argentata) is confined to a small area of the Helan Mountains.

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The Anatolian Plateau rises from the lowland coast of the Aegean Sea and eventually converges with the Armenian Highlands to the east of Cappadocia. Its highest point is the dormant stratovolcano Mount Ararat. Wagner’s mountain viper (Montivipera wagneri) is confined to eastern Turkey (Kars, Erzurum, and Agr provinces). The central Turkish mountain viper (M. albizona) is confined to two localities in central Anatolia (Sivas and Kahramanmaras provinces). Both are threatened by overcollection for the international exotic animal trade.

Lowland Boreal Forests Boreal forests, also known as taiga, exist as a nearly continuous belt of mainly coniferous trees interspersed with lakes, bogs, and heaths stretching across both Eurasia and North America. In Eurasia it covers most of Sweden and Finland, much of Norway, some of the Scottish Highlands and the coastal lowlands of Iceland, much of Russia from Karelia in the west to the Pacific (including most of Siberia), and parts of northern Kazakhstan, northern Mongolia, and northern Japan (Hokkaido). Up until relatively recently these immense forests were largely untouched by humans, although in recent decades the southern areas in particular have begun to be conquered. Many animal species have been driven away as a result, retreating to montane regions or more remote areas of forest. On the other hand, some species have been favoured by the opening up of the coniferous forests and the replacement of parts of it by a secondary growth of mixed forests or cultivated areas. The Kamchatka brown bear (Ursus arctos beringianus), the largest bear in Eurasia, is found in the Kamchatka region and a few offshore islands. While the population remains fairly intact the animals are heavily hunted, both legally and otherwise, for their fur and gallbladder as well as for ‘sport’. The Finnish forest reindeer (Rangifer tarandus fennicus) is today confined to north-western Russia (Karelia) and adjacent parts of Finland, where it migrates seasonally between the two countries. As late as 1750 it ranged over the greater part of the Finnish taiga eastward to the Ural Mountains. It was exterminated in Finland due to hunting and forest loss between 1880 and 1910, but returned spontaneously in small numbers during the mid-twentieth century after habitat disturbances in the Russian part of its range. In 1970 the total population was thought to number only a few hundred, although numbers have since increased to several thousand. Other Eurasian reindeer subspecies facing similar threats are the Busk reindeer (R. t. buskensis) from northern Russia, the Kamchatkan reindeer (R. t. phylarchus) from the Kamchatka Peninsula and regions bordering the Sea of Okhotsk and the Siberian forest reindeer (R. t. valentinae) from the forests of the Ural and Altai Mountains. The long-tailed birch mouse (Sicista caudata) is a relatively widespread but naturally rare species from north-eastern China (Heilongjiang and Jilin), the Ussuri region of Far Eastern Russia and Sakhalin Island.

The Eurasian Region

Lowland Broadleaf and Mixed Forests Three or four thousand years ago, temperate deciduous and mixed forests covered a vast area from the British Isles and the northern Iberian Peninsula in the west across central Europe and eastward in a narrowing strip far into Siberia and the Korean Peninsula. Almost the same type of forest existed in China, Manchuria, and Japan. Stone Age men had burned and cut here and there, but it was not until farming spread in central Europe and Asia that the forests began to disappear. Industrialization combined with growing human populations caused them to retreat at an ever-accelerating rate. With the forests went many of the larger carnivores and also several hoofed animals. Several mammals belonging to forest regions of Eurasia have vanished forever. Two highly threatened subspecies of Central Asian red deer (Cervus hanglu) inhabit forested river valleys in Central Asia. The Bactrian deer (C. h. bactrianus) is found in Kazakhstan, Uzbekistan, Tajikistan, Turkmenistan, and northern Afghanistan. Its range and numbers were much reduced during the 1960s due to persecution and the destruction of riverine vegetation, but conservation activities have succeeded in restoring it to much of its former range. The Yarkand deer (C. h. yarkandensis) shares a similar type of habitat in the valleys of the Tarim, Konqi, and Qarqan rivers of northwestern China (Xinjiang). In the 1960s it was described as very rare, but precise information about its status continues to be lacking. The water deer (Hydropotes inermis), so-named for its preference for riverine areas, is small with notably prominent canines and an absence of antlers. There are two subspecies. The Korean water deer (H. i. argyropus) was historically found along the western coast of the Korean Peninsula and into north-eastern China (Jilin and Liaoning). It has since been extirpated from the latter country. The Muisk vole (Microtus mujanensis) is known only from the Muya Valley in east-central Siberia. The greater noctule bat (Nyctalus lasiopterus) is found in scattered populations across central and southern Europe from the Iberian Peninsula to the Urals, where it favours deciduous forests. A tree-roosting species, it is threatened mainly by deforestation. The Charnali wall lizard (Darevskia dryada) is confined to lowland forests of the Black Sea coastal region (north-eastern Turkey and possibly south-western Georgia), where it is threatened by deforestation. Kaznakov’s viper (Vipera kaznakovi) is confined to the Black Sea coast and forested foothills of the Caucasus Mountains, where is it threatened by overcollection for the international pet trade and loss of habitat. Two species of Asian salamander (Hynobius) endemic to the Korean Peninsula are threatened by loss of habitat. The Kori salamander (H. yangi) is confined to forests in southeastern South Korea (Gyeongsang province). The Cheju salamander (H. quelpaertensis) is known from the southern tip of

South Korea, where it occurs on both the mainland as well as on the islands of Chindo and Cheju.

Lowland Mediterranean Forests, Woodlands, and Scrub Areas of lowland Mediterranean forest, woodland, and scrub within the Eurasian region extend from the Iberian Peninsula to the Levant, and include the Mediterranean Islands. The Anatolian leopard (Panthera pardus tulliana) was a little-known form native to south-western Turkey. It was last recorded in the mid-1970s, and is now considered extinct. The Iberian lynx (Lynx pardinus) was historically distributed over large areas of the Iberian Peninsula and southern France, but habitat destruction, particularly deforestation, along with sharp declines in rabbit populations and human persecution reduced it to two separate regions. In 1950 the northern population still extended from the Mediterranean to Galicia and parts of northern Portugal, while the southern included suitable habitat in central and southern Spain. By the end of the twentieth century, however, it had been reduced to just two small subpopulations (in the mountains of southcentral Spain and the coastal plains of the south-west, respectively), numbering perhaps around 100 animals in total. Since then, a range of conservation measures have been implemented in both Spain and Portugal including a captive breeding programme, habitat restorations, and limited translocations and reintroductions. The Italian hare (Lepus corsicanus) occurs in southern Italy and on the islands of Elba, Sicily, and Corsica (where it was introduced by humans but has since been largely extirpated). It has declined due to habitat destruction, hunting, and competition from introduced species. The Mersin spiny mouse (Acomys cilicicus) is known only from a small area of coastal rocky scrubland in southern Anatolia, Turkey. Roach’s mouse-tailed dormouse (Myomimus roachi) is known only from scattered records across western coastal Turkey, south-eastern Bulgaria, and possibly eastern Greece, although fossil material indicates that it was historically much more widespread. Hermann’s tortoise (Testudo hermanni) is confined to southern Europe, where it is divided into two subspecies. The western Hermann’s tortoise (T. h. hermanni) is found in eastern Spain, southern France, central Italy, the Balearic Islands, Corsica, Sardinia, and Sicily. The eastern Hermann’s tortoise (T. h. boettgeri) is found in Bosnia-Herzegovina, Croatia, Montenegro, Serbia, Kosovo, North Macedonia, Romania, Bulgaria, Albania, Turkey, and Greece. Both are threatened by overcollection for the international pet trade. Orlov’s viper (Vipera orlovi) is confined to a small area of the Black Sea coast in south-western Russia, where it is threatened by overcollection for the international pet trade. The bay Lycian salamander (Lyciasalamandra billae), Antalya Lycian salamander (L. antalyana), Marmaris Lycian

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The Palearctic Realm Figure 2.5 Anatolian leopard (Panthera pardus tulliana). (Credit: unknown.)

salamander (L. flavimembris), and Atif’s Lycian salamander (L. atifi) are all confined to small areas of coastal south-western Turkey, where they are threatened by habitat destruction and overcollection. Fazila’s Lycian salamander (L. fazilae) is confined to south-western coastal Turkey and to the islands of Tersane and Domuz. Luschan’s Lycian salamander (L. luschani) is divided into two subspecies, the nominate form of which (L. l. luschani) is confined to a small area of coastal south-western Turkey.

Eurasian Steppe The Eurasian steppe consists of vast temperate plains stretching across much of central Asia from Eastern Europe to Manchuria. To the north they are bounded by boreal forests. There is no clear southern boundary, although the land becomes increasingly dry as one moves south. The steppe narrows at two points, thereby dividing it into three major parts. All are covered by more or less dense herbaceous vegetation, sometimes with bushes but usually with no trees except along rivers. In the past they were inhabited by wild herds of large mammals, mainly horses and asses, adapted to this special habitat, now mostly depleted by hunting. More recently steppes have come under increasing threat due to agricultural development. The Eurasian wild horse or tarpan (Equus ferus ferus) historically ranged from Spain to central Russia. Its humancaused extinction began in southern Europe, possibly in antiquity. Humans had been hunting wild horses continually for meat since the Palaeolithic, just as areas of available habitat for these large herbivores were continually being lost due to

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the advent of civilization. The subspecies held out longest on the steppes of southern Russia and Ukraine, where it was still living as late as the nineteenth century. By then most ‘tarpans’ had become hybrids, having long interbred with domestic horses, and true wild horses were already very rare by 1880. After that only dubious sightings were documented. The last true tarpan mare was accidentally killed during an attempt to capture her, and the last known individual of all died in captivity on an estate near Poltava in 1909. The story did not end there, however. Beginning in the 1930s several attempts were made to develop horses that looked like tarpans by way of selective breeding. These ‘bred back’ animals do have a superficial resemblance to the extinct form, but cannot be said to be true Eurasian wild horses.

Western Steppe The Western or Pontic–Caspian Steppe begins near the mouth of the Danube and extends north-east almost to Kazan and then south-east to the southern tip of the Ural Mountains. Its northern edge was a broad band of forest steppe (a mosaic of deciduous forests and grasslands) that has now been entirely destroyed by conversion to agriculture. Most of the original large animals have also long-since been exterminated. The European ground squirrel (Spermophilus citellus) is found in central and south-eastern Europe where it is dependent upon short turf in order to construct its colonial tunnel systems. The species has adapted somewhat to artificial habitat such as pastures, golf courses, and parks, but has nevertheless been eliminated from many parts of its range.

The Eurasian Region Figure 2.6 Only known photograph of an alleged living tarpan, taken at the Moscow Zoo in 1884. The animal is most likely a hybrid. (Credit: Moscow Zoo.)

The Podolian mole-rat (Spalax zemni) has a patchy distribution in Ukraine where it is threatened by habitat destruction. The Anatolian vole (Microtus anatolicus) is confined to an isolated area of steppe in central-western Turkey (Konya province). Pannonian Steppe The Pannonian Steppe is an enclave separated from the main Eurasian Steppe by the mountains of Transylvania. It is located mainly in Hungary but also includes parts of Austria, Bulgaria, Romania, Serbia, Slovakia, and Croatia. The Pannonian viviparous lizard (Zootoca vivipara pannonica) is confined to a small area of Austria and Hungary. The Romanian meadow viper (Vipera ursinii moldavica) and Orsini’s meadow viper (V. u. rakosiensis) are both rare and threatened by overcollection for the international pet trade.

Central Steppe The greatest grass steppe in the world, the Central or Kazakh Steppe extends from the Urals to Dzungaria (north-western China), where it narrows between the Tian Shan and Altai Mountains. To the south it grades off into semi-desert and desert. The saiga antelope (Saiga tatarica) historically inhabited a vast area of Eurasian steppe and semi-desert from the foothills of the Carpathian and Caucasus Mountains to Mongolia and north-western China. Two subspecies are recognized. The Russian saiga antelope (S. t. tatarica) is by far the more widespread, but its original range has been much reduced due to hunting and habitat destruction. Today only small

pockets survive in Kazakhstan, Mongolia, Russia, Turkmenistan, and Uzbekistan. The Olkhon mountain vole (Alticola olchonensis) is confined to rocky steppe areas on Olkhon and Ogoi islands, Lake Baikal. The Central Asian tortoise (Agrionemys horsfieldii) is threatened by overcollection for the international pet trade.

Eastern Steppe The Eastern Steppe extends from north-western China through Mongolia to Manchuria, where it very nearly reaches the Pacific Ocean. The Mongolian wild horse (Equus ferus przewalski) is the last surviving subspecies of the now extinct wild horse. Until the eighteenth century, when it first became known to Western science, it ranged from the Russian steppes east to the semideserts of Kazakhstan, Mongolia, and northern China. Soon after it appears to have undergone a catastrophic decline. With the introduction of modern firearms Chinese and Mongolian hunters had become a serious threat to the survival of this species. Simultaneously, nomadic tribes with cattle occupied the areas where it lived and particularly the watering places, forcing the horses to retreat to less-favourable habitat in the mountains. It remained rare and little-known until reported again from Central Asia by the explorer, Colonel Nikolai Przewalski, at the end of the nineteenth century. European zoos quickly became interested in adding the animals to their collections, and several ambitious expeditions were mounted to their last known stronghold, the arid steppes in the corner between China, Mongolia, and Russia. In all, some 53 animals were captured and brought back successfully to Europe around

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The Palearctic Realm

the turn of the century, with a few more collected in the 1930s and 1940s. The few remaining wild animals survived until the mid-twentieth century, where they were last seen in 1969 in the Dzungarian Gobi Desert. Fortunately, by then the captive population had grown to around 150. Since the 1990s reintroduction efforts have been undertaken in protected areas of Mongolia and China, with further projects planned for Kazakhstan and Russia. The Evoron vole (Microtus evoronensis) is known only from a small area of far south-eastern Siberia within the lower Amur River floodplain. Jankowski’s bunting (Emberiza jankowskii) breeds in grassland areas of north-eastern China (Manchuria) and far north-eastern North Korea, where it is threatened by loss of habitat. It formerly also occurred in Russia (southern Primorye), but disappeared from there by the early 1970s. The Peking gecko (Gekko swinhonis) still has a relatively wide distribution in the grasslands of north-eastern China, but is threatened by harvesting for ‘traditional medicine’ and habitat destruction. Mongol Steppe The Mongol Steppe includes both Mongolia as well as the Chinese province of Inner Mongolia, with the two being separated by the Gobi Desert. It is bordered on the south by the Tibetan Plateau. The Mongolian saiga antelope (Saiga tatarica mongolica) is nowadays confined to western Mongolia north of the Altai Mountains, having been extirpated in China in the middle of the twentieth century due to hunting for its horns, which are highly valued in ‘traditional medicine’. In recent years mass die-offs of entire herds have occurred in both subspecies as a result of disease.

Deserts and Semi-Deserts Vast areas of central Asia consist of deserts or semi-deserts, the latter sometimes shading off into arid steppe. For the purposes of this book I have treated these habitats as one, because most of the larger mammals in these poor environments were driven into them from grasslands by man and his livestock. A surprising number of the larger animals have survived in the Eurasian deserts until the mid-twentieth century, when man, equipped with motorized vehicles and modern firearms, began to slaughter them recklessly. The two-humped or Bactrian camel actually consists of two closely related species that, although similar in appearance, have nevertheless descended from distinct ancestors. The domesticated Bactrian camel (Camelus bactrianus) remains a common animal of the steppes and deserts of Iran, Afghanistan, Pakistan, Kazakhstan, Mongolia, and China. The wild Bactrian camel (C. ferus) historically ranged from about the great bend of the Yellow River, across the deserts of southern Mongolia and north-western China to central Kazakhstan. By the middle of the nineteenth century it had

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been extirpated from the western part of its range and survived only in remote areas of the Gobi and Taklimakan deserts, where populations are highly fragmented. Whether the latter descend from wild animals or their ancestors merely returned to the wild state from domestication is a matter of dispute, but in either case they have now long since been living as pure wild camels. During the 1920s the species could be met everywhere in the Gobi Desert, but during the following decades it declined rapidly. Heavy persecution and competition with domesticated animals for pasture and water probably caused the decline. Today, only about 1400 survive, mostly in the Lop Nur Reserve in China, with a smaller population in the Gobi Desert of Mongolia. The Turkmenian wild ass (Equus hemionus kulan) was historically found in the semi-deserts of northern Afghanistan, western China, Kazakhstan, Kyrgyzstan, southern Siberia, Tajikistan, Turkmenistan, and Uzbekistan. By the 1940s a combination of hunting, competition with livestock and habitat destruction had reduced it to only one location, the Badkhyz Strictly Protected Area in southern Turkmenistan. Subsequent reintroduction efforts in Turkmenistan, Kazakhstan, and Uzbekistan have proven to be only temporarily effective owing mainly to high poaching pressure, and populations have declined to a current total of around 2500–3000. The goitered gazelle (Gazella subgutturosa) is so-named for an enlargement of the neck and throat that males develop during mating season. Three subspecies are found throughout the arid and semi-arid regions of Central Asia and the Near East, where they have been historically slaughtered in great numbers and are still subject to illegal poaching. The Persian goitered gazelle (G. s. subgutturosa) still survives in Azerbaijan, Syria, Iran, southern Afghanistan, and western Pakistan, but has disappeared from much of its former range. The Turkmenian goitered gazelle (G. s. gracilicornis) lives in Kazakhstan, Uzbekistan, Turkmenistan, and Tajikistan, while the Yarkand goitered gazelle (G. s. yarkandensis) occurs in northern and northwestern China and Mongolia. The pallid pygmy jerboa (Salpingotus pallidus) is confined to a few localities in central and eastern Kazakhstan. Dahl’s jird (Meriones dahli) is confined to Armenia and eastern Turkey (Agri province). The desert dormouse (Selevinia betpakdalaensis) is a rare species known only from a few localities over a relatively wide area of central Kazakhstan. The Bokhara mouse-eared bat (Myotis bucharensis) is known only from a few specimens collected from three localities in Central Asia (Uzbekistan and Tajikistan). Two lizards of the genus Phrynocephalus are threatened by loss of habitat. Strauch’s toad-headed agama (P. strauchi) inhabits the Fergana Valley of Uzbekistan, Kyrgyzstan, and Tajikistan. Horvath’s toad-headed agama (P. horvathi) is confined to a few isolated populations in the Araks River valley of Armenia, Azerbaijan, and eastern Turkey.

The Eurasian Region

The Gobi Desert The Gobi Desert is a large, cold expanse located in northern and north-western China and southern Mongolia. It is the fifth largest desert in the world and the second largest in Asia. Much of it is not actually sandy, but rather, exposed bare rock. The Mongolian wild ass or khulan (Equus hemionus hemionus) is today largely confined to arid and semi-arid regions of the Gobi Desert in northern China and southern Mongolia. It formerly also occurred in eastern Kazakhstan and southern Siberia, but has been extirpated there due to overhunting. While the population remains large, it continues to be threatened by poaching and loss of habitat.

Miscellaneous Deserts The Karakum Desert (‘Black Sand’) is located east of the Caspian Sea and covers much of Turkmenistan. Golubew’s toad-headed agama (Phrynocephalus golubewii) is an extremely rare species confined to a small area of salt flats near Bami Station in southern Turkmenistan. The Kyzylkum Desert (‘Red Sand’) is a large desert located in Central Asia between the rivers Amu Darya and Syr Darya, a region historically known as Transoxania or Sogdiana. Today it is divided between Kazakhstan, Uzbekistan, and Turkmenistan. Heptner’s pygmy jerboa (Salpingotus heptneri) is confined to the Kyzylkum Desert, where it is known only from a few specimens. The Taklamakan Desert is located in north-western China (south-western Xinjiang). Vaurie’s nightjar (Caprimulgus centralasicus) is known only from a single specimen collected in the mid-twentieth century. Subsequent surveys have failed to locate it, and it is thought that the species may be migratory and either breed or winter elsewhere.

Isolated Caves, Springs and Pools Within the Eurasian Region the most notable karstic region occurs in the Balkans. The olm or proteus (Proteus anguinus) is an entirely aquatic salamander that inhabits subterranean caves within the Dinaric Alps. The nominotypical form (P. p. anguinus) ranges from southern Slovenia and adjoining north-eastern Italy through coastal Croatia and the karst regions of BosniaHerzegovina. It likely occurs in western Montenegro as well. The black olm (P. p. parkelj), first discovered in 1986, is endemic to the underground waters of the Doblicica karst spring near Crnomelj, Slovenia. Both subspecies are extremely vulnerable to changes in their habitat, in particular pollution. The Petzea rudd (Scardinius racovitzai) is a type of freshwater fish confined to a single small hot spring in northwestern Romania, where it is threatened by pollution and water extraction. The Almiri toothcarp (Aphanius almiriensis) is known from two springs in western Greece, one near Almiri (where it has not been recorded since 2003 and is probably extirpated), and the other at Meligou in the Peloponnese. The latter

locality was unfortunately transformed into a bathing area, but the species is still thought to survive in surrounding lagoon habitat. Two minnows of the genus Delminichthys are threatened by water extraction and pollution. The Modro Oko minnow (D. krbavensis) is confined to a single karstic spring in western Croatia. Ghetaldi’s minnow (D. ghetaldii) is confined to karstic streams and springs in southern Dalmatia. Three species of spring minnow (Pseudophoxinus) are highly restricted in distribution. The Akstafa spring minnow (P. sojuchbulagi) is known only from springs in Azerbaijan, where it was last recorded in 1948. None have been found in subsequent searches. The Antalya spring minnow (P. antalyae) is confined to two springs and a stream flowing to the Bay of Antalya in south-western Turkey. The Turianchi spring minnow (P. atropatenus) is confined to two small but very deep springs in Azerbaijan. The Timavo Spring sculpin (Cottus scaturigo) is confined to Timavo Spring in north-western Italy, a resurgence of the partially subterranean Reka River. The Greek ninespine stickleback (Pungitius hellenicus) is confined to three localities in the Spercheios Valley of central Greece, specifically the Aghia Paraskevi spring, a number of drainage channels, and three natural wells. Stefanidis’ spined loach (Cobitis stephanidisi) has been extirpated from its original range in the Kefalovriso karstic spring, within the former Lake Karla basin of central Greece. However, it was subsequently found in the nearby Chasambali spring, where it is seriously threatened by water extraction, pollution, and drought. Starostin’s stone loach (Troglocobitis starostini) is confined to a single cave in the Kugitang Hills of eastern Turkmenistan, where it is highly susceptible to human disturbance and pollution. The cave stone loach (Barbatula sp.) is an as-yet undescribed species first discovered by cave divers in southern Germany in 2015.

The Sultan Sazligi Marshes The Sultan Sazligi marshes are located in south-central Turkey. At one time a very important wetland area, they were unfortunately drained and largely dried out in the early twenty-first century. All remaining springs are fed from a single aquifer. The Sultan Sazligi toothcarp (Aphanius danfordii) and the Sultan Sazligi spring minnow (Pseudophoxinus elizavetae) are both confined to two small spring fields about 15 km apart along with a few canals within the Sultan Sazligi marshes.

Lakes, Rivers, and Marshes The Eurasian Region is rich in freshwater wetlands, being crossed by a number of large river systems and dotted by mainly smaller lakes. Most of the latter were produced by the great ice sheets that covered the northern part of this portmanteau continent for long periods. Northern Eurasia is therefore

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The Palearctic Realm

much richer in lakes than is the southern part. A high proportion of the original shallow lakes and marshes of northern Europe have long since been drained for agricultural purposes. Long regarded as a step towards effective land use and prosperity, during the nineteenth and twentieth centuries draining had become a sort of mania. Oftentimes these efforts proved to be less than successful. Economically they failed to produce the expected agricultural harvests, biologically the productivity of the areas was greatly reduced, and hydrologically the surrounding lands suffered owing to the sinking ground-water level. These unanticipated consequences of draining resulted from a complete failure to appreciate the intimate relationship between aquatic resources and the longterm productivity of the soil. In more recent years hydroelectric systems, including reservoirs with continuously changing volumes, as well as serious pollution have also affected lakes and rivers, interfering with natural food chains, biotic productivity, and the reproduction of waterfowl and economically important fishes such as salmon and trout. With the disappearance or poisoning of many marshes, lakes, and deltas, the habitats of aquatic animals have been greatly reduced and many species have gone extinct or become seriously threatened. The Eurasian beaver (Castor fiber) was once widespread in Europe and Asia, but was hunted to near-extinction both for its fur and castoreum (a yellowish secretion used as a tincture in perfumes). By 1900, only 1200 survived in 8 relict populations. Reintroduced throughout much of its former range, it now occurs from Great Britain to Mongolia and China, although it remains absent from southern Europe and the Middle East. The European mink (Mustela lutreola) is a semi-aquatic mustelid that was historically found from Finland to east of the Ural Mountains and south as far as northern Spain and the Caucasus. Heavily hunted for its luxuriant fur, it is now reduced to a few isolated pockets across this vast range. Habitat destruction and the introduction of American mink (Neovison vison) are additional threats. The Russian desman (Desmana moschata) is a small, longsnouted, semi-aquatic insectivore that inhabits wetlands of the Volga, Don, and Ural river drainages in Russia, Ukraine, and Kazakhstan. Long hunted for their rich, thick fur, the main threats now are habitat destruction and degradation and competition with introduced muskrats (Ondatra zibethicus). The Pyrenean desman (Galemys pyrenaicus) occurs in the northern and central parts of Spain and Portugal, the French Pyrenees, and Andorra, but has experienced extreme range contractions across its range. The southern water vole (Arvicola sapidus) is confined to isolated wetland areas of France, Spain, and Portugal, where it is threatened mainly by loss of habitat. The relict gull (Ichthyaetus relictus) breeds in a small number of saline lakes in the arid steppe regions of Mongolia, eastern Kazakhstan, south-central Russia, and northern China, from where it migrates to South Korea and

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perhaps north-eastern China as well. It is threatened mainly by coastal development in its non-breeding range. The Far Eastern curlew (Numenius madagascariensis) breeds in eastern Russia and north-eastern Mongolia, from where it winters as far south as Australia and New Zealand. It is threatened mainly by loss of habitat. Swinhoe’s rail (Coturnicops exquisitus), the world’s smallest rail species, was long known only from two small breeding areas separated by more than 1000 km in northeastern China and south-eastern Siberia. A new breeding population situated between the two was discovered in the Amur region in 2018, and more recently in Japan. The species is everywhere rare and declining due to loss of habitat. The large-headed water snake (Natrix megalocephala) is found in western Transcaucasia in south-western Russia, western Georgia, and far north-eastern Turkey, where it prefers fast-flowing mountain streams as well as lower-elevation woodlands. It is threatened by loss of habitat and introduced racoons (Procyon lotor). The Italian yellow-bellied toad (Bombina pachypus) ranges through much of the Italian Peninsula south of the Po River, where it is nevertheless threatened by the loss of its wetland habitat. The Suweon tree frog (Dryophytes suweonensis) is confined to wetlands areas in the lowlands of the north-western South Korea, where it is threatened by loss of habitat. The Tavas frog (Rana tavasensis) is confined to two areas of south-western Turkey. The Italian agile frog (R. latastei) is confined to a few localities in northern Italy and (marginally) southern Switzerland. Both are threatened by loss of habitat. Three species of Eurasian water frog (Pelophylax) are threatened by loss of habitat, pollution, and overcollection. The Epirus water frog (P. epeiroticus) is confined to small pockets of suitable habitat in southern Albania and western Greece (including the island of Corfu). The Albanian water frog (P. shqipericus) is confined to western Albania and southern Montenegro. The Korean water frog (P. chosenicus) is confined to the western Korean Peninsula. The European sturgeon or beluga (Huso huso) is the largest freshwater fish in the world, with the all-time accepted record being a 7.2-m individual taken from the Volga estuary in 1827 that weighed 1571 kg. It is an anadromous species historically known from the Caspian, Black, Azov, and Adriatic basins, but has been extirpated from the Adriatic and Azov seas due to overfishing and loss of spawning sites due to dams. It has long been in heavy demand for the female’s valuable roe – better known as beluga caviar – and although governments have attempted for decades to restrict the trade, poaching continues. Sturgeons of the genus Acipenser are widespread across Eurasia and North America, where they are anadromous (i.e. able to live in both fresh and salt water). Many species are highly threatened by overfishing, damming of their spawning rivers, and the international trade in caviar. The Atlantic sturgeon (A. sturio) was historically found throughout the

The Eurasian Region

North and Baltic seas, the English Channel, the European and Icelandic coast of the Atlantic, the northern Mediterranean, and parts of the Black Sea. Today only a single small population remains, in the Garonne River of south-western France, where it last spawned in 1994. The starry sturgeon (A. stellatus) was historically found throughout the Caspian, Black, Azov, and Aegean sea basins, but has been extirpated from the latter and it is predicted that the remaining natural population will soon follow. The Siberian sturgeon (A. baerii) is normally divided into two subspecies, the nominate form of which (A. b. baerii) historically inhabited all the Siberian rivers draining into the Kara, Laptev, and East Siberian seas as well as the Irtysh River in north-western China (Xinjiang). The latter wild population was extirpated in the 1950s, although small numbers continue to be artificially restocked. The sterlet (A. ruthenus) inhabits the large rivers flowing into the Black, Azov, and Caspian seas, as well as a few others in Siberia. Populations migrating between fresh and salt water have all been extirpated. The diamond sturgeon (A. gueldenstaedtii) was historically found in the Caspian, Black and Azov sea basins, although aquaculture has resulted in both intentional and accidental introductions throughout Europe. It is currently known only from the Caspian Sea, where it spawns in the Ural and Volga rivers, and from the Black Sea where spawning occurs in the lower Danube and Rioni rivers. There is no native spawning population remaining in the Sea of Azov, only introduced (stocked) individuals. The ship sturgeon (A. nudiventris) was historically abundant in the Black, Aral, and Caspian sea basins, with possible relict populations in the Rioni River of Georgia and the Safid Rud in Iran. The healthiest population is in Lake Balkhash in Kazakhstan, well outside its natural range, where the species was introduced in the 1960s for commercial purposes. It has also been introduced to the upper Illi River in China and to the Syr-Darya River in the Aral Sea basin. The Adriatic sturgeon (A. naccarii) was historically confined to the rivers and lagoons of northern Italy and the eastern coasts of the Adriatic Sea. All wild populations are now most likely extirpated, with the species now totally dependent upon stocking from aquaculture. The lavaret whitefish (Coregonus lavaretus) was historically confined to Lake Bourget in France and to Lake Geneva, with an additional population having been introduced to Lake Aiguebelette in France centuries ago. The Lake Geneva population was extirpated in the early twentieth century, although the reason is unknown. The pfärrit whitefish (C. confusus) was historically known from lakes Morat and Bienne in Switzerland, although it may occur in Lake Neuchâtel as well. The Lake Morat population was extirpated in the 1960s due to eutrophication and water level management. Trybom’s whitefish (C. trybomi) was historically known for certain only from a few lakes in southern Sweden. Only the population in Lake Fegen survives today, the other populations having been extirpated in the 1970s and 1980s due to acid rain and the stocking of introduced species. The Siberian taimen (Hucho taimen) is a salmon-like game fish with a wide geographic range within Eurasia, including

parts of the Caspian and Arctic drainages as well as portions of the Pacific drainages. It is everywhere threatened by overfishing, pollution, and loss of habitat. The Yessey char (Salvelinus tolmachoffi) is confined to lakes Yessey, Siltak, and Bezymannoye in the Khatanga River drainage of north-central Siberia, and to Khantaiskoye Lake in the Yenisey drainage. It may also be present in a few lakes of the Gydanskiy Bay basin on the Taimyr Peninsula. It is threatened by pollution from heavy metal mining and by overfishing. The common carp (Cyprinus carpio) has been domesticated since the Middle Ages, and cultivated stocks are assumed to have been derived from the wild European carp (C. c. carpio) which is native to the Black, Caspian, and Aral sea basins. This subspecies has been introduced to other parts of Europe as well as to the Middle East and north-western Africa, and has also been established worldwide in large quantities for use as food. Unfortunately, hybridization with domesticated introduced stocks has become a seriously threat to genetically pure populations. Another subspecies, the Deniz carp (C. c. yilmaz), is confined to Anatolian Turkey. The Austrian lakes trout (Salmo schiefermuelleri) is known historically from three subalpine lakes (Attersee, Traunsee, and Fuschlersee). It is most likely extinct. The Adriatic trout (S. obtusirostris) is confined to a few rivers in Croatia, BosniaHerzegovina, and Montenegro. The Pelagos trout (S. pelagonicus) is confined to a few rivers in North Macedonia and northern Greece. The flathead trout (S. platycephalus) is known only from three mountain streams in central Turkey. There is some question, however, as to the validity of the species. The Valencia toothcarp (Valencia hispanica) is confined to coastal areas of east-central Spain. The species appears to have formerly ranged as far as southern France. Letourneux’s toothcarp (V. letourneuxi) is confined to Lake Butrint in southern Albania and to a few areas of coastal western Greece. The species is believed to have been extirpated from the islands of Corfu and Lefkas. Kemal’s golden barb (Garra kemali) is confined to central Anatolia, where it is known from the Eregli Marshes, Lake Meyil, and the Lake Beysehir basin. Four barbels of the genus Barbus are threatened by habitat degradation, water extraction, and introduced species. The brook barbel (B. caninus) was historically confined to the Po and Isonzo rivers of northern Italy, southern Austria, and southern Switzerland. It is not clear whether it still survives in the Isonzo River. The Crimean barbel (B. tauricus) is confined to a few streams on the southern tip of the Crimean Peninsula. The Macedonian barbel (B. macedonicus) is confined to a few rivers in northern Greece and North Macedonia. The Catalan barbel (B. haasi) is confined to the Ebro River drainage and a few coastal rivers in north-eastern Spain. The Greek barbel (Luciobarbus graecus) is confined to the Sperchios River drainage and Lake Yliki in eastern Greece. It formerly occurred in Lake Paralimni as well, which has since

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The Palearctic Realm

been drained. The Bulatmai barbel (L. capito) is found in the Caspian and Aral seas along with their inflowing rivers. It has been extirpated over much of its range due to expanding hydropower development, overfishing, and pollution. The Iberian barbel (L. comizo), Valencia barbel (L. guiraonis), and Steindachner’s barbel (L. steindachneri) are all confined to the Tagus and Guadiana river drainages of south-western Spain and southern Portugal, where they have declined considerably since the 1990s. The Dalmatian barbelgudgeon (Aulopyge huegelii) inhabits both above-ground and subterranean karst rivers as well as lakes in Croatia and Bosnia-Herzegovina, but is everywhere threatened by pollution and invasive species. Two species of scraper (Capoeta) are endemic to Turkey, where they are threatened by water extraction, drought, and introduced species. The longsnout scraper (C. mauricii) appears to have been historically widespread in central Turkey, but is two confined to the south-eastern Lake Beysehir basin and few other isolated streams. The Pamphylian scraper (C. antalyensis) is confined to the Aksu and Koprucay river drainages of south-western Turkey. The pike-asp (Aspiolucius esocinus) is found in the Amu Darya and Syr Darya rivers and a few lakes in Tajikistan, Kazakhstan, Turkmenistan, and Uzbekistan. The Antalya bleak (Alburnus baliki) is known only from four streams draining to the Gulf of Antalya in south-central Turkey. The Manyas bleak (A. carinatus) is confined to Lake Kus (Lake Manyas) and Lake Uluabat (Lake Apalyont) in north-western Turkey, from where they ascend a handful of tributaries to spawn. The eastern Aegean bleak (A. demiri) is confined to a few river drainages in south-western Turkey. The Gediz bleak (A. battalgilae) is confined to the lower Gediz and Koca drainages of east-central Turkey, where it has been extirpated from a number of localities. The Italian bleak (A. albidus) is confined to the rivers of southern Italy. Nasreddin’s bleak (A. nasreddini) was historically found in Lake Eber and Lake Aksehir and their tributaries. Massive water extraction and pollution have destroyed most of its habitat, and the species is currently known only from a single tributary of Lake Aksehir, the Ortakoy River. Members of the genus Telestes are small cyprinid fish that live in the karstic rivers and lakes of the Balkan region. Many are threatened by habitat loss due to water extraction and drought. The spring dace (T. fontinalis) is confined to intermittent rivers and karsts in Croatia. The Boeotian dace (T. beoticus) is confined to the Kifissos and Asopos river drainages of south-eastern Greece. It was historically also found in the catchment rivers for lakes Yliki and Paralimni, which are now totally drained. The Croatian dace (T. croaticus) is confined to a few river systems in central Croatia. The striped dace (T. metohiensis) is confined to a few streams in southern Croatia, Bosnia-Herzegovina, and Montenegro. Four species of nase of the genus Chondrostoma are threatened by water extraction, introduced species, and pollution. The eastern Aegean nase (C. holmwoodii) was

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historically known from the Kucuk Menderes, Izmir, Gediz, and Bakir river drainages of western Turkey. It has apparently been extirpated from the Kucuk Menderes. The Tefenni nase (C. fahirae) is historically known from Kirkpinar spring near Tefenni in south-western Turkey, where it was extirpated and had to be reintroduced, with unknown success. Populations are found in Lake Karatash and a stream within the Lake Burdur basin as well. The Italian nase (C. soetta) was historically confined to northern Italy, Slovenia, and southern Switzerland, from where it has been introduced into a few lakes in west-central Italy. It has been extirpated from Slovenia and the Isonzo River drainage. The minnow nase (C. phoxinus) is known only from a few freshwater karst localities in Croatia and Bosnia-Herzegovina. Coelho’s nase (Pseudochondrostoma duriense) is confined to a few river drainages in north-western Spain and northern Portugal. Willkomm’s nase (P. willkommii) is confined to southern Spain and southern Portugal. Both are threatened mainly by habitat destruction. The south-west European nase (Parachondrostoma toxostoma) is confined to the Rhône drainage of France and Switzerland and to coastal rivers in France. It is threatened by dam construction and introduced species. The Turia nase (P. turiense) is confined to the Turia and Mijares river drainages of east-central coastal Spain, where it is threatened by pollution and introduced species. Three species of nase of the genus Iberochondrostoma endemic to the Iberian Peninsula are threatened by habitat destruction, pollution, and predation and competition from introduced fish species. Almaca’s arch-mouthed nase (I. almacai) is confined to the Mira, Arade, and Bensafrim river drainages of southern Portugal. The Lusitanian arch-mouthed nase (I. lusitanicum) is confined to a few rivers in south-western Portugal. Lemming’s arch-mouthed nase (I. lemmingii) is confined to the Tagus, Guadiana, Odiel, and Guadalquivir river drainages of south-western Spain and southern Portugal. The Aegean minnow (Phoxinus strymonicus) is known only from the Aggitis (Strymon) River in northern Greece, although the species may also occur in the Loudias and Filiouris drainages. Strandja’s minnow (P. strandjae) is confined to the Veleka and Resowska river drainages of Bulgaria and Turkey. The Dinaric minnow (Phoxinellus alepidotus) is confined to a few river drainages in southern Croatia and BosniaHerzegovina. A great number of spring minnows of the genus Pseudophoxinus are endemic to Anatolian Turkey, where they are threatened by water extraction and drought. The giant spring minnow (P. anatolicus) is still found in a number of lakes and streams in central Turkey and their tributaries and in the Eregli marshes. It has been extirpated from Lake Beysehir. The Sandikli spring minnow (P. maeandricus) is known from Karadirek stream near Sandikli in west-central Turkey, an isolated basin which flows underground to Isikli spring, and to the Lake Hotamis basin in south-central Turkey where it

The Eurasian Region

was extirpated some time ago due to the draining of the marshes. The Pamphylian spring minnow (P. alii) is known from the Ilica and Kormurculer streams as well as a section of the Aksu River, all of which flow into the Bay of Antalya in south-central Turkey. The Lycian spring minnow (P. evliyae) is known from the Lake Avlan basin, which drains through the Akçay River to the Mediterranean, and to the Lake Söğüt basin, both in south-western Turkey. Lake Avlan has become a seasonal lake, making three springs the only refuge sites for the species, while Lake Sogut was drained many years ago and the species only survives in a few canals fed by a single spring. The central Anatolian spring minnow (P. burduricus) is confined to the Lake Burdur and Lake Salda drainages and to a few other isolated springs and streams. The fat spring minnow (P. crassus) is known from a few lakes and streams in west-central Turkey. The species most likely occurred in Lake Samsam, which was entirely drained in the 1970s. The Apamean spring minnow (P. maeandri) is confined to an undefined locality in Turkey. The Izmir minnow (Ladigesocypris mermere) is known only from a few specimens collected from southwestern Anatolia. The spotted minnow (Delminichthys adspersus) and Ghetaldi’s minnow (D. ghetaldii) are both confined to a few rivers and lakes in Croatia and Bosnia-Herzegovina. The Peloponnese minnow (Pelasgus laconicus) is confined to fragmented populations within the Eurotas and Alfeiós rivers in southern Greece. The Estremadura minnow (Achondrostoma occidentale) is known from the Alcabrichel, Sizandro, and Safarujo drainages of west-central Portugal, although it is believed that the Safarujo population was extirpated when the stream dried up. Arcas’ minnow (A. arcasii) is confined to a few rivers in northern Portugal and northern Spain. Both are threatened by habitat destruction, pollution, and predation and competition from introduced fish species. The Crimean riffle minnow (Alburnoides maculatus) is confined to small streams of the Crimean Peninsula, where it is threatened by water extraction and drought. The European mudminnow (Umbra krameri) has a scattered distribution in the Danube and Dniestr River drainages of south-central Europe, where it is threatened by the disappearance of its backwater habitat. It has been extirpated from many locations. The Spartan minnowroach (Tropidophoxinellus spartiaticus) is confined to a few small rivers and streams in the southern Peloponnese of Greece. Steindachner’s chub (Iberocypris alburnoides) is still fairly widespread in lakes and rivers within the southern Iberian Peninsula of Spain and Portugal, but is everywhere under threat by loss of habitat and introduced species. The Aksehir chub (Squalius recurvirostris) is known from at least three stream catchments, one each flowing to lakes Ilgin, Eber, and Aksehir in west-central Turkey. The species historically occurred in lakes Eber and Aksehi themselves, but

have been extirpated due to desiccation and pollution, although it still survives in Lake Ilgin. The thick-lipped chub (S. cephaloides) is known only from the Tesvikiye and (formerly) Armutlu streams in north-western Turkey. The Toscana chub (S. lucumonis) is confined to a few populations with the Tiber, Arno, Ombrone, and Serchio river drainages of central-west Italy. The Málaga chub (S. malacitanus) is confined to the Guadalmina and Guadiaro rivers of southern coastal Spain. The Valencia chub (S. valentinus) is confined to a few coastal rivers in south-eastern Spain. The Arade chub (S. aradensis) is confined to the Arade, Algibre, and Bordeira river drainages of southern Portugal. The Stymphalia chub (S. moreoticus) is known only from Lake Stymphalia and Vouraikos river drainage of south-central Greece. The Neretva chub (S. svallize) is confined to the Neretva and Trebisnjica river drainages in Bosnia-Herzegovina, southern Croatia, and Montenegro. The Livno chub (S. tenellus) is confined to a few karstic streams in Bosnia-Herzegovina and southern Croatia, and may have been introduced to Lake Blidinje more than a century ago. The Elmali rudd (Scardinius elmaliensis) is confined to a few streams and springs in south-western Turkey. The Tiber rudd (S. scardafa) was formerly found throughout the Tiber River drainage, but is now entirely confined to Lake Scano, a locality outside its natural range, where it was introduced in the late nineteenth century. Meidinger’s roach (Rutilus meidingeri) is currently known from lakes Attersee, Mondsee, and Wolfgangsee in northern Austria, with an additional small population in an Austrian stretch of Danube. The species has been extirpated from Lake Traunsee in Austria and from Lake Chiemsee in south-eastern Germany (although it continues to be stocked there). Panos’ roach (Leucos panosi) is confined to lakes Trichonis and Ambrakia and to the Achelous and Louros river drainages of western Greece. The Yliki roach (L. ylikiensis) was historically known from the Kifissos River drainage, including lakes Yliki and Paralimni (the latter now dry) in central Greece. Duran’s sculpin (Cottus duranii) is only known from a few specimens collected from the upper Loire and Dordogne drainages of south-central France. The Eber gudgeon (Gobio intermedius) historically occurred in lakes Eber and Aksehir in west-central Turkey, but are now confined to a few inflowing streams due to drought and pollution. The Salgir gudgeon (G. krymensis) is confined to a few rivers at the southern tip of the Crimean Peninsula, where it is threatened by water extraction. The Benacus gudgeon (Romanogobio benacensis) was historically confined to the Po River drainage and coastal rivers in northern Italy, where it has disappeared from many areas. The species has been introduced into the Arno and Ombrone rivers, and to the Isonzo and Reka rivers of Slovenia. The Arno goby (Neogobius nigricans) is confined to streams and lakes within the Tyrrhenian catchment of northcentral Italy. It is threatened by habitat degradation and introduced species.

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The Palearctic Realm

Brauner’s goby (Benthophiloides brauneri) is a rarely recorded freshwater and brackish water species from the surrounding rivers, lakes, and estuaries of the Black and Caspian sea basins. It is threatened by loss of habitat due to water extraction and drought. The Iyidere goby (Ponticola rizensis) is confined to three streams in north-eastern Turkey, where it is threatened by dam construction. The Gediz dwarf goby (Knipowitschia mermere) is known from Lake Marmara and the lower Gediz and Madra river drainages of western Turkey. The Croatian dwarf goby (K. croatica) is found in clear karstic springs, lakes, and smaller rivers within the Neretva and Matica drainages of Croatia and Bosnia-Herzegovina. It is threatened by eutrophication, pollution, and water extraction. The Baetic toothcarp (Aphanius baeticus) is confined to the lower Guadalquivir River and a few streams in southwestern Spain. While protected within Doñana National Park it remains threatened by habitat destruction, pollution, repeated drought, and the introduction of exotic species. Aristotle’s catfish (Silurus aristotelis) was historically confined to Lake Trichonida, Lake Lysimachia, and Lake Amvrakia in western Greece. The species is of considerable commercial interest to local fisheries and was introduced to Lake Pahvotis as well as to Lake Volvi in northern Greece. The Kashmir mountain catfish (Glyptothorax kashmirensis) is known from the Jhelum River within the Indus River drainage of Pakistan, as well as from northern India (Jammu and Kashmir). An additional record from the Ganges River drainage in Nepal requires confirmation. It is thought to be seriously threatened by damming. The Carian sportive loach (Oxynoemacheilus germencicus) is known only from the Buyuk Menderes and lower Gediz drainages in western Anatolia. It likely also occurred in the Kucuk Menderes River, which lies between the two known rivers, but is now extirpated there due to pollution and water extraction. Two pond loaches of the genus Seminemacheilus are endemic to west-central Turkey, where they are threatened by water extraction and drought. Lendli’s pond loach (S. lendlii) was historically widespread across central Anatolia but is now known only from a few isolated springs and tributaries of Lake Tuz and Lake Beysehir. The Ispart pond loach (S. ispartensis) is confined to the Egirdir, Aksehir, and Eber lake basins. Stephanidis’ spined loach (Cobitis trichonica) is confined to lakes Trichonis, Lysimachia, Ozeros, and Amvrakia, and to the Acheloos River drainage in western Greece. The brown spined loach (C. puncticulata) is confined to a few rivers and lakes in north-eastern Greece and north-western Turkey. The Phrygian spined loach (C. phrygica) is confined to a few lakes, springs, and streams in south-western Turkey. The Bithynian spined loach (C. splendens) is confined to a single small stream on the Black Sea coast of north-western Turkey. Calderon’s spined loach (C. calderoni) is confined to the Douro, Ebro, and

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Tagus river drainages of northern Portugal and northern Spain. The Illyrian spined loach (C. illyrica) is confined to a small area of southern Croatia. De Buen’s spined loach (C. paludica) is confined to the rivers of the central and southern Iberian Peninsula. Zanandrea’s spined loach (C. zanandreai) was historically confined to the Volturno River drainage of south-central Italy. It is also present in Lake Fondi and a few associated lagoons, where it was most likely introduced. The Turkish brook lamprey (Lampetra lanceolata) is confined to widely separated streams in northern Turkey, one flowing into the Black Sea at Trabzon and another running to Lake Sapanca in Western Anatolia. The Greek brook lamprey (Eudontomyzon hellenicus) is today known only from two brooks within the Struma River drainage of north-eastern Greece. It was historically found in the Louros River drainage in western Greece as well, but appears to have extirpated.

The Caspian Sea The Caspian Sea is the largest enclosed inland body of water on Earth by area, although shrinking rapidly due to climate change and human activity along the Volga River. Its fauna is poor but extremely varied owing to its former connections with other seas from the Arctic to the Mediterranean. The Caspian seal (Pusa caspica) is a small, ice-breeding species endemic to the Caspian Sea. Unsustainable commercial hunting has been responsible for a dramatic decline in population. The Persian sturgeon (Acipenser persicus) is confined to the Caspian Sea, where it is most abundant in the southern part. Historically it ascended all the rivers of the basin to spawn, but is now restricted to the lower courses of a few rivers in Iran. The species is heavily fished for its meat and roe, and continues to be stocked from aquaculture. The Volga shad (Alosa volgensis) is an anadromous species of commercial fishing importance which historically ascending from the Caspian Sea up the Volga, Terek, and Ural rivers to spawn. It has been extirpated from the latter two rivers, and the construction of the Volgograd Dam has severely curtailed its migration in the Volga. The beloribitsa whitefish (Stenodus leucichthys) has lost all of its spawning grounds along the Volga River due to the construction of dams. It is now extinct in the wild, but survives in cultured stocks. The Caspian kutum (Rutilus kutum) was historically common, but appears to have declined dramatically in recent years owing to marine pollution and overfishing. The Khvalyn spined loach (Cobitis amphilekta) is known for certain only from Kyzylagach Bay and the lower Kumbashi River in Azerbaijan.

The Aral Sea The Aral Sea was an endorheic lake (i.e. one with no outflow) lying between Kazakhstan in the north and Uzbekistan in the south. Formerly the fourth largest lake in the world, it began to shrink in the 1960s after the rivers that fed it were diverted by

The Eurasian Region

Soviet irrigation projects. By 1997 it had declined to just 10 per cent of its original size, in the process splitting into four separate lakes. By 2009 the south-eastern lake had disappeared, and the south-western lake had been reduced to a thin strip. Satellite images taken by NASA in 2014 revealed that the eastern basin had completely dried up (it is now known as the Aralkum Desert). The shrinking of the Aral Sea has been called one of the worst environmental disasters in modern history. The region’s once-prosperous fishing industry has been essentially destroyed. The Aral barbel (Luciobarbus brachycephalus) was historically found throughout the Aral Sea basin, along with the Chu River drainage and the western Caspian Sea. Today it survives only in a few reservoirs of the Amu Darya and Syr Darya (tributaries of the Aral Sea) and the Karakum Canal, where it is invasive. In the western Caspian Sea the landlocked population in the Kura River is thought to be stable, as are those in the southern tributaries, but migrating populations have declined due to a lack of spawning sites and poaching. The Amu Darya and Syr Darya Rivers The Amu Darya (historically the Oxus) and Syr Darya (historically the Jaxartes) rivers are the primary tributaries of the Aral Sea. The sharpray (Capoetobrama kuschakewitschi) is a type of cyprinid fish confined to the Aral Sea basin. Kuschakewitsch’s sharpray (C. k. kuschakewitschi) is confined to the Amu Darya and Syr Darya river systems. Fedtschenko’s false shovelnose sturgeon (Pseudoscaphirhynchus fedtschenkoi) was historically found in the Aral Sea as well as the Syr Darya and the middle and lower reaches from the Kara Darya River. Last recorded in 1960, it is most likely extinct. The dwarf false shovelnose sturgeon (P. hermanni) and Kaufmann’s false shovelnose sturgeon (P. kaufmanni) are both confined to a few localities within the Amu Darya River.

is the deepest and oldest lake in the world, and the largest freshwater lake by volume. Famous for its crystal-clear waters and unique wildlife, the lake is nevertheless under threat by pollution, poaching and development. The Baikal seal (Pusa sibirica) is the only entirely landlocked seal species in the world and one of the smallest. Endemic to the lake and its tributary rivers, it remains quite common despite hunting and pollution, but could be at risk in the future owing to climate change. The Baikal sturgeon (Acipenser baerii baikalensis) is confined primarily to the northern end of the lake, migrating up the Selenga River to spawn. Long a target for fishermen due to its enormous size (specimens up to 125 kg were once common), it is now officially protected but remains threatened. The Baikal whitefish or omul (Coregonus migratorius), one of the lake’s most economically important fish, was once overfished to such an extent that by the 1960s its survival, also imperilled by other environmental disturbances, was threatened. It has since recovered.

Lake Saimaa Lake Saimaa is located in south-eastern Finland. One of the largest natural freshwater lakes in Europe, it was formerly connected to the Gulf of Finland and the Baltic Sea, but has been landlocked since the last Ice Age. The Saimaa ringed seal (Pusa hispida saimensis) is endemic to the lake, where the total population is estimated at around 200–250 individuals. It has been fully protected since 1958, but pollution and complaining fishermen constitute a threat to its existence.

Lake Geneva

The Chu River The Chu River is located in northern Kyrgyzstan and southern Kazakhstan. The Chu sharpray (Capoetobrama kuschakewitschi orientalisi) is confined to the Chu River.

Lake Geneva (Lac Léman in French) is located on the northern side of the Alps between France and Switzerland. It is one of the largest lakes in western Europe. Two species of economically important whitefish (Coregonus) endemic to Lake Geneva have been driven to extinction due to a combination of eutrophication and overfishing. The gravenche whitefish (C. hiemalis) appears to have disappeared in the early 1900s, while the fera whitefish (C. fera) was last recorded in 1920.

Lake Urmia

The Prespa Lakes

Lake Urmia (Daryace Orumiye in Persian) is an endorheic salt lake located in north-western Iran. At its greatest extent it was the largest lake in the Middle East, but has shrunk to a fraction of its former size owing to damming of the rivers that flow into it and groundwater pumping in the surrounding area. The Urmia bream (Acanthalburnus urmianus) is confined to southern and western tributaries of the Lake Urmia basin.

The Prespa Lakes are two connected freshwater lakes located high in the Balkan Mountains of south-eastern Europe. Great Prespa Lake is shared between Greece, Albania, and North Macedonia, while Small Prespa Lake is shared only between Greece and Albania. The area contains three national parks. The Prespa bleak (Alburnus belvica) is confined to the Prespa Lakes. The Prespa trout (Salmo peristericus) is known for certain only from Agios Germanos stream in the Prespa Lakes region of north-western Greece, although it may be present as well in one or two streams in North Macedonia.

Lake Baikal Lake Baikal is a rift lake located in southern Siberia near the Mongolian border. So large that it is often mistaken for a sea, it

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The Palearctic Realm

The Prespa minnow (Pelasgus prespensis) is confined to the Prespa Lakes, where it is threatened by water extraction, pollution, disease, and the introduction of exotic fish species. The Prespa riffle minnow (Alburnoides prespensis) is confined to the Prespa Lakes. The Prespa nase (Chondrostoma prespense) is confined to the Prespa Lakes. The Prespa roach (Rutilus prespensis) is confined to shallow nearshore and swampy areas of the Prespa Lakes. The Prespa spined loach (Cobitis meridionalis) is a rare nearshore species from the Prespa Lakes.

Lake Ohrid Lake Ohrid straddles the mountainous border between southwestern Macedonia and eastern Albania. One of Europe’s deepest and oldest lakes, it features a unique aquatic ecosystem of global importance and many endemics. The Belushka trout (Salmo ohridanus), Struga trout (S. balcanicus), Ohrid summer trout (S. aphelios), and Lumi trout (S. lumi) are all endemic to Lake Ohrid, where they are threatened by artificial hybridization with the more common Pestani trout (S. letnica) as well as by overfishing, poaching, and degradation of the lake’s water quality. The Ohrid minnow (Pelasgus minutus) was historically confined to swampy areas of the Lake Ohrid basin. It has not been collected since 1973, and may be extinct. The Ohrid riffle minnow (Alburnoides ohridanus) is confined to the surf zone along the lake’s shoreline. Interbreeding with introduced species is a possible threat. The Ohrid gudgeon (Gobio ohridanus) is naturally endemic to Lake Ohrid, although it has been introduced to France and perhaps other countries.

Lake Van Lake Van (Van Golu in Turkish) is a large endorheic lake located in eastern Turkey. The Karasu Sha bleak (Alburnus timarensis) is confined to the lower part of Karasu stream in the Lake Van basin. The Van scraper (Capoeta kosswigi) is confined to the Lake Van basin. The Van sportive loach (Oxynoemacheilus ercisianus) is confined to a few streams flowing into Lake Van, where it is threatened by sand mining and dam construction.

Lake Neuchâtel Lake Neuchâtel (Lac de Neuchâtel in French/Neuenburgersee in German) is a large lake located in western Switzerland. The Neuchâtel deepwater char (Salvelinus neocomensis) is known only from three specimens collected in 1896, 1902, and 1904. Research undertaken in the 1950s and in 2003 appear to confirm its extinction. The Neuchâtel whitefish (Coregonus candidus) is naturally confined to the lake, where eutrophication caused a population decline in the 1970s. It has since somewhat recovered. An introduced (possibly hybridized) population lives in Lake Maggiore.

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Lake Beysehir Lake Beysehir (Beysehir Golu in Turkish) is a large freshwater lake located in south-central Turkey. The Beysehir bleak (Alburnus akili) was endemic to Lake Beysehir, where it became extinct soon after the introduction of pike-perch (Sander lucioperca) in 1955. The Beysehir nase (Chondrostoma beysehirense) is confined to part of Lake Beysehir and three adjacent streams. Two species of gudgeon (Gobio) are endemic to the Lake Beysehir basin. The Beysehir gudgeon (G. microlepidotus) is known only from a few streams flowing into Lake Beysehir, although it may occur in the lake itself. The Eyilikler gudgeon (G. battalgilae) is known only from Eyilik stream in the northern Lake Beysehir basin. The Hittitic spring minnow (Pseudophoxinus hittitorum) is known only from a spring at Eflatunpınar, east of Lake Beysehir, and Bakaran stream, which drains from the south of the lake. Two spined loaches (Cobitis) are confined to the Lake Beysehir basin. The Beysehir spined loach (C. battalgili) is known only from three streams in the Lake Beysehir basin, and in the Manavgat stream that flows to the Mediterranean. Bilsel’s spined loach (C. bilseli) is confined to a few streams in the Lake Beysehir basin.

Lake Egirdir Lake Egirdir (Egirdir Golu in Turkish) is a large freshwater lake located in south-western Anatolia. The introduction of the predatory pike-perch (Sander lucioperca) has had a decimating impact upon the native fish species. Hanlirsch’s spring minnow (Pseudophoxinus handlirschi) has not been recorded since the 1980s despite intensive fieldwork, and is thought to be extinct. The Egirdir spring minnow (P. egridiri) is restricted to two densely vegetated tributaries and one large shoreline spring in the Lake Egirdir basin. At one time feared extinct, it was rediscovered in 1993. The Egirdir longsnout scraper (Capoeta pestai) was formerly found throughout Lake Egirdir, but is now confined to a single inflowing stream (Caykoy Creek) along with a small area of adjacent lake where it is considered highly threatened.

Lake Burdur Lake Burdur (Burdur Golu in Turkish) is a large, deep saline lake located in south-western Turkey. The Sureyan toothcarp (Aphanius sureyanus) and the Burdur toothcarp (A. burduricus) are both confined to Lake Burdur, where they are threatened by water extraction, damming of the lake’s sources, and drought. The Burdur sportive loach (Oxynoemacheilus anatolicus) is confined to three separate, short, spring-fed streams that used to drain into Lake Burdur.

Lake Tuz Lake Tuz (Tuz Golu in Turkish) is located in central Turkey. It is the second largest lake in the country and one of the largest hypersaline lakes in the world.

The Eurasian Region

The Cappadocian chub (Squalius cappadocicus) is known only from the Melendiz River, a small inflow of Lake Tuz. The Cappadocian gudgeon (Gobio gymnostethus), Taurus gudgeon (G. hettitorum), and Cihanbeyli gudgeon (G. insuyanus) are all confined to streams and marshes within the Lake Tuz basin. The Tuz sportive loach (Oxynoemacheilus eregliensis) is confined to streams and springs within the Lake Tuz basin. The Tuz spined loach (Cobitis turcica) is confined to a few springs and streams in the Lake Tuz basin.

Lake Vistonis Lake Vistonis is a shallow coastal lagoon located in northeastern Greece. General threats include damming, pollution, and water extraction for irrigation. The Vistonis bleak (Alburnus vistonicus) is confined to the Lake Vistonis drainage. The Thracian shad (Alosa vistonica) is confined to the Lake Vistonis drainage.

Lake Trichonida Lake Trichonida is a large lake located within the Acheloos River drainage of western Greece. It is connected to the smaller Lake Lysimachia. The Trichonida dwarf goby (Economidichthys trichonis) is confined to Lake Trichonida and Lake Lyssimachia. The Trichonida combtooth blenny (Salaria economidisi) is confined to Lake Trichonida. It is threatened by loss of habitat.

Lake Skadar Lake Skadar is located on the border between Albania and Montenegro. It is the largest lake in southern Europe. The Skadar nase (Chondrostoma scodrense) is known only from nine specimens collected in the late nineteenth century from the Lake Skadar basin. Surveys in recent decades have failed to find the species, and it is now considered to be extinct. The Skadar gudgeon (Gobio skadarensis) is known only from Zeta stream and the lower Moraca River, within the Lake Skadar basin.

Lake Ammer Lake Ammer (Ammersee in German) is a large lake located in the upper Danube basin of southern Germany (Bavaria). The Ammersee char (Salvelinus evasus) is confined to deep waters in Lake Ammer. The Ammersee ruffe (Gymnocephalus ambriaelacus) is a type of perch confined to Lake Ammer, where it is threatened mainly by introduced species. The Ammersee whitefish (Coregonus bavaricus) is confined to deep waters in Lake Ammer.

Lake Ladoga Lake Ladoga is a freshwater lake located in north-western Russia near Saint Petersburg. It is the largest lake in Europe.

The Ladoga ringed seal (Pusa hispida ladogensis) is confined to Lake Ladoga, where it is threatened by human disturbance. The Ladoga whitefish (Coregonus baerii) is threatened by overfishing and damning of its spawning rivers.

The Tagus River The Tagus River (Rio Tajo in Spanish/Río Tejo in Portuguese) is the longest river on the Iberian Peninsula. It has its origins in east-central Spain and flows directly west through Portugal to the Atlantic Ocean. The Lisbon arch-mouthed nase (Iberochondrostoma olisiponensis) is confined to three small tributaries within the lower Tagus River drainage of Portugal. The Arrago River The Arrago River is located in western Spain. The Alagón spined loach (Cobitis vettonica) is confined to the Arrago River. The Gallo River The Gallo River is located in central Spain. The Gallo chub (Squalius castellanus) is known from specimens collected from the Gallo River and its main tributaries, the Bullones and Arandilla rivers. It is threatened mainly by pollution from the city of Molina de Aragon.

The Guadiana River The Guadiana River flows through central Spain before turning south along a southern stretch of the Portugal–Spain border and emptying into the Atlantic. The small-headed barbel (Luciobarbus microcephalus) is largely confined to the Guadiana River drainage, where it is threatened by dam construction, pollution, water extraction, and introduced species. It occurs as well in a small stretch of the Tajo River, where it was most likely introduced. The Iberian minnowcarp (Anaecypris hispanica) is confined to the Guadiana River drainage.

The Guadalquivir River The Guadalquivir River is located in south-central Spain. The Guadalquivir chub (Iberocypris palaciosi) is confined to the Guadalquivir River drainage. The Jándula River The Jándula River is a tributary of the Guadalquivir River. The Jándula arch-mouthed nase (Iberochondrostoma oretanum) is confined to the Robledillo and Fresnada rivers, small tributaries of the Jándula River.

The Danube River The Danube is Europe’s second largest river drainage, being roughly twice the size of California. The river flows over 2857 km from Germany’s Black Forest to the shores of the Black Sea.

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The Danube salmon (Hucho hucho) is found patchily throughout the Danube River drainage, with very few selfsustaining populations. The species has been locally introduced to Lake Constance in Germany, the Vistula River in Poland, the Tagus River in Spain, the Rhône River in France, and most likely other drainages where it apparently maintains only through stocking. The Arges River The Arges River is located in southern Romania. The sculpin-perch (Romanichthys valsanicola) was formerly known from the upper Arges River and its tributaries, the Vâlsan and Râul Doamnei. It is now known only from a single 1-km stretch of the upper Vâlsan, where the population is entirely reliant upon the amount of water released by a reservoir immediately upstream. The Arges sculpin (Cottus transsilvaniae) is known only from its original collection from the upper Arges River in 1998. The Beli Vit River The Beli Vit River is located in north-western Bulgaria. The Beli Vit sculpin (Cottus haemusi) is known only from its original collection from the Beli Vit River in the 1980s. The Danube Delta The Danube delta is located in south-western Romania and south-eastern Ukraine. Antipa’s gudgeon (Romanogobio antipai) was historically found in the Danube delta. Last recorded in the 1960s, it is now considered to be extinct although the reason for this remains unknown. The Danube delta dwarf goby (Knipowitschia cameliae) is known only from a single small, brackish lagoon south of the Danube delta in Romania. Last recorded in 1994, it is possibly extinct.

The Lake Constance deepwater char (Salvelinus profundus) is confined to the deeper waters of Lake Constance. It was thought to have gone extinct during the 1970s due to eutrophication, but was unexpectedly rediscovered in 2016.

The Rhône River One of the major rivers of Europe, the Rhône rises in the Swiss Alps. After passing through Lake Geneva, it runs down through south-eastern France before dividing near its mouth with the Mediterranean at Arles. The resulting delta constitutes the Camargue wetland region. The Rhône streber (Zingel asper) is confined to four fragmented subpopulations in the rivers Durance and Beaume, Rhône River drainage, with no hope of connectivity between them owing to dam construction. It is threatened by pollution.

The Neretva River The Neretva River is located in Bosnia-Herzegovina and Croatia. The Neretva nase (Chondrostoma knerii) is confined to the main channel of the Neretva River. The Imotski chub (Squalius microlepis) is confined to two lakes and a few small karstic streams within the Neretva River drainage. The Mostar minnow (Phoxinellus pseudalepidotus) is known only from a single stream within the Neretva River drainage of Bosnia-Herzegovina. The Neretva spined loach (Cobitis narentana) is confined to the Neretva River drainage. The Norin River The Norin River is a small karstic stream located in southern Croatia. The Norin dwarf goby (Knipowitschia radovici) is confined to the Norin River.

The Rhine River

The Pinios River

The Rhine is the second longest river in central and western Europe (after the Danube). It runs from Switzerland to the North Sea.

The Pinios River is located in east-central Greece. The Thessaly gudgeon (Gobio feraeensis) was historically found in the Pinios and Karla lakes within the Pinios River drainage, but has been extirpated from the latter when it was largely drained for agriculture. The Thessaly dwarf goby (Knipowitschia thessala) is confined to the Pinios River catchment.

Lake Constance Lake Constance (Bodensee in German) is a lake on the Rhine at the northern foot of the Alps in Germany, Switzerland, and Austria. It consists of three bodies of water: Upper Lake Constance, Lower Lake Constance, and a connecting stretch of river. Gmelin’s whitefish (Coregonus gutturosus) was historically endemic to the deeper waters of Lake Constance. An important commercial species in the 1960s, it is thought to have become extinct in the 1970s when eutrophication of the lake reached its peak. Kottelat’s whitefish (C. arenicolus) still survives in the lake, where it is threatened by hybridization with a more common species.

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The Krka River The Krka River is located in coastal Croatia. Tursky’s dace (Telestes turskyi) is confined to Lake Busko Blato in Bosnia-Herzegovina and to the Cikola River, a tributary of the Krka River in southern Croatia. It was thought to be extinct until rediscovered in 2002. The Vrba minnow (Phoxinellus dalmaticus) is confined to Vrba Creek, on the Cikola tributary of the upper Krka River drainage.

The Eurasian Region

Lake Visovac Lake Visovac is located in the lower Krka River drainage. Mrakovcic’s dwarf goby (Knipowitschia mrakovcici) is confined to Lake Visovac. The Cikola River The Cikola River is located in north-western Croatia. The Cikola dace (Telestes polylepis) is today confined to a single karstic stream within the Cikola River drainage, but was formerly more widespread.

The Cetina River The Cetina River is located in southern Croatia and western Bosnia-Herzegovina. The Cetina dace (Telestes ukliva) is confined to the Cetina River drainage in Croatia, where it was thought to have gone extinct in the 1980s. It was rediscovered in 1997. The Cetina spined loach (Cobitis dalmatina) is confined to the Cetina River drainage.

The Menderes barbel (Luciobarbus kottelati) is confined to the Buyuk Menderes drainage. The Isikli nase (Chondrostoma meandrense) is confined to the Buyuk Menderes drainage, where it is most abundant in Lake Isikli. The Apamean spring minnow (Pseudophoxinus maeandri) is confined to two springs within the upper Buyuk Menderes drainage. The Isikli gudgeon (Gobio maeandricus) appears to be absent from Lake Isikli itself, but is known from two separate inflowing spring systems and some additional spring-fed streams elsewhere. The Isparta algae-eater (Crossocheilus klatti) is known from the Kopu River drainage and Lake Isikli basin in the upper Buyuk Menderes drainage. It formerly inhabited lakes Egirdir and Golcuk, but is now believed to have been extirpated there. The Menderes sportive loach (Oxynoemacheilus cinicus) is known only from a single imprecise type locality, most likely within the Buyuk Menderes drainage.

The Ceyhan River The Ceyhan River (historically known as the Pyramus) is located in south-central Turkey. The Ceyhan spring minnow (Pseudophoxinus zekayi) is confined to streams, springs, and lakes within the middle Ceyhan River drainage. The Ceyhan chub (Squalius seyhanensis) is known only from two localities within the upper Ceyhan River drainage. The Ceyhan spined loach (Cobitis evreni) is known only from three stream catchments of the middle Ceyhan River drainage. The Cilician sportive loach (Oxynoemacheilus seyhanicola) and the Elbistan stone loach (O. ceyhanensis) are both endemic to the Ceyhan River drainage.

Lake Isikli Lake Isikli (Isikli Golu in Turkish) is located within the upper Buyuk Menderes drainage. The chocolate chub (Squalius carinus) is confined to the Lake Isikli basin, where it appears to be absent from the lake itself but is known from two separate inflowing spring systems and some spring-fed streams near Dinar. The Isikli sportive loach (Oxynoemacheilus mesudae) is known from four springs and spring-fed streams within the Lake Isikli basin. While still abundant, the water level is declining steadily due mainly to extraction, and several small springs where it presumably once occurred have already dried out.

The Zamanti River The Zamanti River is a headwater stream in the Ceyhan River watershed. The Zamanti sportive loach (Oxynoemacheilus seyhanensis) and the Zamanti stone loach (Barbatula samantica) are confined to the Zamanti River, where they are threatened by water extraction, pollution, and drought.

The Euphrates is the longest river in western Asia. It originates on the Armenian Plateau of Armenia, Azerbaijan, northwestern Iran, and eastern Turkey, from where it flows south through Syria and Iraq to the Persian Gulf. Firat’s spring minnow (Pseudophoxinus firati) is known only from two widely separated localities in central and eastern Turkey. The Erzurum sportive loach (Oxynoemacheilus araxensis) and the Mancilik stone loach (O. paucilepis) are each confined to a few streams within the upper Euphrates drainage of Turkey.

The Caysuyu River The Caysuyu River is a tributary stream of the Ceyhan River located in Kayseri province. The Caysuyu stone loach (Barbatula tschaiyssuensis) is confined to the Caysuyu River, where it is threatened by water extraction, pollution, and drought.

The Buyuk Menderes River The Buyuk Menderes River (Büyük Menderes Irmagi in Turkish, and historically known as the Maeander) is located in south-western Turkey. General threats include pollution, water extraction, and dam construction.

The Upper Euphrates River

The Upper Tigris River The upper Tigris is located primarily within the Taurus Mountains of south-eastern Turkey. The Yuksekova River is an endorheic basin nested within the upper Tigris River catchment of south-eastern Turkey (Hakkari province). The Yuksekova chub (Petroleuciscus kurui) is confined to the Yüksekova River drainage.

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The Batman River is a major tributary of the upper Tigris River located in south-eastern Turkey. The Diyarbakir loach (Paraschistura chrysicristinae) is known from two localities within the Batman River, but has not been recorded since the 1970s. It is possibly extinct, though the reason for its disappearance is unknown. The Göksu spined loach (Cobitis kellei) is known only from its original collection from the Göksu stream in 1974. Not recorded during more recent surveys, it is thought that the construction of a dam may have driven it extinct. The Queiq River (also known as the Aleppo River) is located in south-eastern Turkey and northern Syria. It has been largely desiccated by drought and water extraction. The Halap sportive loach (Oxynoemacheilus tigris) was historically found throughout the Quieq River drainage but is now known only from a single stream, only a few hundred metres long between two reservoirs, in south-eastern Turkey.

The Chornaya River The Chornaya River is a small river on the Crimean Peninsula. The Chornaya tubenose goby (Proterorhinus tataricus), Chornaya gudgeon (Gobio delyamurei) and Chornaya spined loach (Cobitis taurica) are all confined to a very limited stretch (about 1 km in length) of this drainage, below the Chornaya Gorge. Water extraction and drought threaten to dry up their habitat entirely.

The Amur River The Amur River flows for nearly 3000 km from the mountains of northern China and south-eastern Russia before finally emptying into the North Pacific. During the summer monsoon rains flood parts of the river, although for six months in the winter and spring large portions are frozen. The kaluga (Huso dauricus), a type of sturgeon, is one of the largest freshwater fishes in the world and capable of attaining a length of 5.6 m and at least 1000 kg in weight. Still found in small numbers throughout the Amur River drainage and perhaps in coastal waters, it has been fished to near extinction for its valuable roe. The Amur sturgeon (Acipenser schrenckii) is another enormous freshwater fish species that often reaches a length of up to 3 m and a weight of over 190 kg. Still found throughout the entire Amur River drainage from its estuary to the upper tributaries, it has nevertheless declined steadily since the end of the nineteenth century due to overfishing and is now considered to be seriously threatened.

Miscellaneous Lakes, Rivers, and Marshes Lake Bourget (Lac du Bourget in French) is a large, deep lake located in the southernmost end of the Jura Mountains in France. The Bourget whitefish (Coregonus bezola) was a deep-water species that went extinct in the 1960s according to local fishermen. Lake Morat (Lac de Morat in French/Murtensee in German) is located in western Switzerland. The Morat

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whitefish (Coregonus restrictus) was last recorded in 1890. Surveys in the 1950s found no whitefish at all, and in recent years a related species has been introduced to the lake. Lake Breiter Luzin (Breiter Luzin in German) is located in northern Germany. The Breiter Luzin whitefish (Coregonus lucinensis) is confined to the lake. During the 1970s the population declined heavily due to eutrophication, but has since improved along with the water quality. Lake Cheim (Cheimsee in German) is a glacial, pre-alpine lake located in southern Germany. The Cheimsee whitefish (Coregonus hoferi) was last recorded in the 1940s, although fishermen indicated its presence in the lake up until the late 1980s. It is likely extinct. Lake Atter (Attersee in German) is a large lake located in northern Austria. The Attersee whitefish (Coregonus atterensis) is historically confined to the lake, with an additional introduced population in nearby Lake Mond. It is threatened by competition from introduced fish species. Lake Traun (Traunsee in German) is located in northern Austria. The Traunsee whitefish (Coregonus danneri) is confined to the lake, where it is threatened by competition from introduced fish species. Lake Garda (Lago di Garda in Italian) is a large glacial lake in northern Italy. The Garda trout (Salmo carpio) is historically endemic to the lake, where it is seriously threatened by introduced species. It has, however, been introduced into a number of other lakes both in Italy and elsewhere. Lake Posta Fibreno (Lago di Posta Fibreno in Italian) is located in central Italy. It is an elongated karstic lake rich in underground springs and caves. The Fibreno trout (Salmo fibreni) is confined to the lake and its tributaries. Lake Doiran (Límni Dhoiráni in Greek) is located on the Greece/North Macedonia border. The Doiran bleak (Alburnus macedonicus) is confined to Lake Doiran. Lake Pamvotida is a large lake located in north-western Greece. The Pamvotida minnow (Pelasgus epiroticus) is confined to Lake Pamvotida, where it has declined considerably in recent decades due to overfishing, pollution, and introduced species. Lake Yliki (Limni Yliki in Greek) is located in central Greece. The Greek rudd (Scardinius graecus) is confined to the lake, where it is threatened by large fluctuations in water level. Lake Volvi (Límni Vólvi in Greek) is located in northeastern Greece. Ongoing threats include introduced species, water extraction for irrigation, and eutrophication. Two species, the Macedonian shad (Alosa macedonica) and the Volvi bleak (Alburnus volvitticus), are nowadays endemic to the lake. Formerly both were also found in Lake Koronia, but in 1995 the latter dried up, killing all the fish. Lake Techirghiol is a hypersaline lake located in coastal south-eastern Romania. The Techirghiol stickleback (Gasterosteus crenobiontus) was historically confined to freshwater springs of Lake Techirghiol. Hybridization with the three-spined stickleback (G. aculeatus) led to its extinction in the 1960s.

The Eurasian Region

Lake Mandras (Mandrensko ezero in Bulgarian) is located in eastern Bulgaria. The Mandras bleak (Alburnus mandrensis) is confined to the Lake Mandras basin, where it is threatened by pollution and the impoundment of its spawning streams. Lake Iznik (Iznik Golu in Turkish) is located in northwestern Turkey (Bursa province). The Iznik bleak (Alburnus nicaeensis) was known only from the Lake Iznik basin. Alien species stocked into the lake to improve fisheries yields likely led to its extinction. It has not been reported since the late twentieth century despite several intensive searches. The Iznik sportive loach (Oxynoemacheilus phoxinoides) is confined to just one small stream in the Lake Iznik basin. Lake Acigol (Acigol Golu in Turkish) is a located in westcentral Anatolia, Turkey. The Acigol toothcarp (Aphanius transgrediens) was, prior to the 1970s, found throughout Lake Acigol and its catchment basin. The lake has since dried up, and today the species is confined to a single small spring field and a few short feeder streams. Lake Apolyond (Apolyond Golu in Turkish) is located in western Anatolia, Turkey, within the Sea of Marmara basin. The Apolyond sprat (Clupeonella muhlisi) is confined to Lake Apolyond. Lake Golcuk (Golcuk Golu in Turkish) is a small mountain lake in east-central Turkey (Izmir province). The Golcuk toothcarp (Aphanius splendens) was known only from Lake Golcuk. The year of its extinction is unknown, but it had already disappeared by the 1980s. Lake Kezenoi-am is located in the Caucasus Mountains of south-western Russia (Chechnya). The Kezenoi-am trout (Salmo ezenami) was historically endemic to Lake Kezenoiam, although it appears to have been introduced into Lake Mochokh (Daghestan) in the early 1960s. Lake Abrau is a small karst lake located near the Black Sea coast of south-western Russia. The Abrau sprat (Clupeonella abrau) is confined to Lake Abrau, where it is threatened by water extraction and introduced species. Lake Balkhash is located mainly in south-eastern Kazakhstan. One of the largest lakes in Asia, like the Aral Sea it is shrinking owing to the diversion of water from the rivers that feed it. It used to have a rich fish fauna with a number of endemic species, but since the 1970s biodiversity has been declining. The Balkhash perch (Perca schrenkii) occurs in Lake Balkhash and the Alakol lake group, where it is threatened by the introduction of a predatory fish. Lake Elgygytgyn is an impact crater lake located in the Chukchi region of north-eastern Siberia. The long-finned char (Salvethymus svetovidovi) and the Elgygtgyn char (S. elgyticus) are confined to the lake. The Mira River (Rio Mira in Portuguese) is located is southern Portugal. The Torgal chub (Squalius torgalensis) is confined to the Torgal tributary of the Mira River. The Douro River (Rio Doero in Spanish/Rio Douro in Portuguese) is located in north-western Spain and northern Portugal. The Sarda minnow (Achondrostoma salmantinum)

is confined to the Águeda, Yeltes, Turones, and Uces tributaries of the Douro River drainage of western Spain (Salamanca province). The Júcar River (Rio Jucar in Spanish) is located in eastcentral Spain. The Júcar nase (Parachondrostoma arrigonis) is confined to the Júcar River drainage. The Lez River (Le Lez in French) is located in southern France near Montpellier. The Lez sculpin (Cottus petiti) is confined to a short (3 km) stretch of this coastal river, from its karstic springs source to the Lirou tributary. The Hérault River (L’Hérault in French) is a small coastal drainage in southern France. Rondelet’s sculpin (Cottus rondeleti) is confined to three stretches of stream (each only a few hundred metres long) within the Hérault River. The Jadova River is located in western Croatia. The Jadova minnow (Delminichthys jadovensis) and the Jadova spined loach (Cobitis jadovaensis) are both confined to a single ephemeral stream within the Jadova River drainage. The Dragonja River is located in Slovenia and Croatia. The Istrian chub (Squalius janae) is confined to the upper Dragonja River. The Vardar River (historically known as the Axios River) is located in Greece and North Macedonia. The Balkan streber (Zingel balcanicus) is known only from a few specimens collected in the Vardar River drainage. The Macedonian trout (Salmo macedonicus) is confined to the upper Vardar River drainage. The Moraca River is located in Montenegro, where it arises in the mountains of the north-east and empties into Lake Skadar. The Zeta trout (Salmo taleri) is confined to the upper Moraca drainage and its tributary, the Zeta River. It is threatened by hybridization with introduced brown trout (S. trutta). The Provadiskaya River is located in Bulgaria. The Varna gudgeon (Gobio kovatschevi) is confined to the Provadiskaya River. The Struma River is located in Bulgaria and Greece. The Struma spined loach (Cobitis punctilineata) is confined to the Aggitis stream, a tributary of the Struma River in northern Greece. The Aoös River is located in north-western Greece and south-western Albania. The Pindus sportive loach (Oxynoemacheilus pindus) is confined to the Aoös River drainage. The Arachthos River is located in north-western Greece. The Arachthos spined loach (Cobitis arachthosensis) is confined to the Arachthos River drainage. The Louros River is located in north-western Greece. The Louros spined loach (Cobitis hellenica) is confined to the Louros River drainage. The Acheron River is located in western Greece. The Acheron dwarf goby (Knipowitschia milleri) is confined to the Acheron River delta. The Eurotas River (also known as the Evrotas) is located on the Peloponnese Peninsula of southern Greece. The

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The Palearctic Realm

Eurotas chub (Squalius keadicus) is confined to the Eurotas River. It is threatened by pollution and the seasonal drying of the river. The Kucuk Menderes River is located in western Turkey. It is heavily desiccated in its upper reaches and heavily polluted in the lower ones. The Ephesus dwarf goby (Knipowitschia ephesi) is confined to the delta and marshes of the lowermost Kucuk Menderes. The Bakircay River is located in western Turkey. The Bakircay bleak (Alburnus attalus) is confined to a few tributaries of the Bakircay River, where it is threatened by water extraction and pollution. The Tahtali River is located in western coastal Turkey. Kosswig’s chub (Squalius kosswigi) is nowadays confined to the Tahtali River, although it may have formerly occurred (or still occur) in the Kucuk Menderes River drainage as well. The Onaç River is located in south-western Turkey. The Onaç spring minnow (Pseudophoxinus ninae) is confined to the Onaç stream drainage. The Gediz River (Gediz Nehri in Turkish) is located in south-western Turkey. The Simav sportive loach (Oxynoemacheilus simavica) is confined to Simav stream, a tributary of the Gediz River. The Koprucay River is located in south-central Turkey, where it drains into the Bay of Antalya. The Pisidian spring minnow (Pseudophoxinus fahrettini) is confined to some headwater tributaries and a spring within the Koprucay River drainage. The Akgol-Eregli Marshes are located in south-central Turkey. The Akgol-Eregli stone loach (Barbatula eregliensis) is confined to the Akgol-Eregli marshes. The Aksu River is located in north-eastern Turkey. The Aksu goby (Ponticola turani) is confined to a few streams within the Aksu drainage. The Lenkoran River is located within the Caspian Sea basin of Azerbaijan. The Lenkoran sportive loach (Oxynoemacheilus lenkoranensis) is confined to the Lenkoran River. The Lena River is located in eastern Siberia. One of the longest rivers in the world, it rises west of Lake Baikal in southern Russia and flows north for some 4400 km before finally emptying into the Arctic Ocean. At its mouth into the Laptev Sea in northern Siberia, the river forms a huge delta of 32,000 km2, which is the largest Arctic delta and the most extensive protected wilderness area in Russia. Baunt’s whitefish (Coregonus sardinella baunti) is confined to the Vitim River, a tributary of the Lena River. The Penzhina River is located in Kamchatka, Russia. The Penzhina whitefish (Coregonus subautumnalis) lives in the Sea of Okhotsk, from where it migrates up the Penzhina River to spawn. It is threatened by overfishing.

Coasts and Satellite Islands This section includes the coastal areas and islands of western and north-western Europe, the Russian Far East, the

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Mediterranean shore of southern Europe, and the majority of the islands within the Mediterranean Sea. The sandy mole-rat (Spalax arenarius) is found only along the lower Dnepr River sands in southern Ukraine, with the main part of the population lying within the Black Sea Biosphere Reserve. The Atlantic puffin (Fratercula arctica) is a type of seabird that can still be found throughout the northern Atlantic on rocky coasts and offshore islands, but has experienced rapid declines across its range. The spotted greenshank (Tringa guttifer) is a type of wading bird that breeds along the south-western and northern coasts of the Sea of Okhotsk, and possibly along western Kamchatka and Sakhalin. At other times the species migrates south across south and South East Asia as far as Sri Lanka and Australia. The total population is very small, however, and threatened by development of coastal wetlands throughout its range. The Yelkouan shearwater (Puffinus yelkouan) is a type of seabird that breeds on islands and coastal cliffs along the central and eastern Mediterranean coast. It is threatened by habitat destruction, fisheries by-catch, and introduced species. Audouin’s gull (Larus audouinii) is found patchily in breeding colonies along the coasts and islands of the Mediterranean, where it was at one time threatened by eggrobbing fishermen. Numbers have since recovered to safe levels.

The Azores The Azores are a remote archipelago located in the North Atlantic some 1360 km west of Portugal. Like so many other volcanic islands, they appear to rise directly from the ocean floor. First discovered by the Portuguese in 1431 and long colonized, the unique laurel forests that once covered them have been almost completely destroyed. The Azores noctule (Nyctalus azoreum) is a type of bat confined to the Azores, where it is still relatively abundant although vulnerable owing to its small range. Monteiro’s storm petrel (Hydrobates monteiroi) is only known to breed on a few small islets in the Azores. It is thought to remain in the vicinity of these islands during the non-breeding season. The Azores wood pigeon (Columba palumbus azorica) is rare but still occurs on a number of islands. São Miguel São Miguel is the largest and most populous island in the Azores. The São Miguel scops owl (Otus frutuosoi) probably became extinct after European settlement due to habitat destruction and the introduction of alien species. The Azores bullfinch (Pyrrhula murina) is confined to one small area of remnant cloud forest in the mountains of São Miguel, where it is considered stable. The total population is around 250.

The Eurasian Region

The British Isles The British Isles are an archipelago in the northern Atlantic consisting of the islands of Great Britain, Ireland, the Isle of Man, and thousands of smaller ones. Animal and plant life are similar to that of the north-western Europe. Great Britain Great Britain is the largest of the British Isles and indeed the largest island in Europe. Threatened vertebrate species consist almost entirely of freshwater fishes. The chars (Salvelinus) are a group of salmonid fish with a circumpolar distribution. Most are typically cold-water fish that primarily inhabit freshwater, though some also migrate to the sea. The following species have very localized distributions and are considered threatened. The golden char (S. youngeri) is known from Loch Eck and possibly a few other lakes in Scotland. The haddy char (S. killinensis) is confined to Loch Killin, Loch Doine, possibly Loch Builg, and a few other Scottish lakes. Malloch’s char (S. mallochi) is confined to Loch Scourie and Loch Shin, Scotland. The Ben Hope char (S. maxillaris) was long known only from its original description, but has since been found in several lakes in northern Scotland. Peris’ char (S. perisii) is known from six locations in northern Wales. The Struan char (S. struanensis) is confined to Loch Rannoch and Loch Ericht in Scotland. Lonsdal’s char (S. lonsdalii) is confined to Haweswater in north-western England. Willoughby’s char (S. willoughbii) is known from Lake Windermere and possibly Ennerdale Water in northwestern England. A number of whitefish (Coregonus) are similarly restricted in distribution. The powan (C. clupeoides) was historically confined to Loch Lomond and Lock Eck in west-central Scotland. It has since been introduced into two reservoirs in the Loch Lomond basin. The gwyniad (C. pennantii) was historically confined to Llyn Tegid (Lake Bala) in northern Wales, where it is threatened by declining water quality and introduced species. It has been introduced into nearby Llyn Arenig. The vendace (C. vandesius) was historically found in lakes Derwentwater and Bassenthwaite in north-western England, and Castle Loch and Mill Loch in south-western Scotland. The two Scottish populations have since been extirpated, although another has been introduced into Loch Skene. The schelly (C. stigmaticus) is confined to four lakes in north-western England (Haweswater, Ullswater, Brotherswater, and Red Tarn). The Shetland Islands are a subarctic archipelago located north-east of Scotland. The Shetland char (Salvelinus gracillimus) was long known only from Loch of Girlsta in Shetland, but may occur in Loch More and possibly other lakes in northern Scotland. The Inner Hebrides are a small archipelago located off the western coast of Scotland. The Orkney char (Salvelinus inframundus) is nowadays confined to Loch Mealt on the Isle of Skye. It formerly occurred in Heldale Water on Hoy Island, in the Orkney Islands, but has not been recorded there since 1908.

The St. Kilda Islands are an isolated archipelago located 64 km west of the Outer Hebrides. The St. Kilda house mouse (Mus musculus muralis) evolved on the tiny island of Hirta, where it was entirely dependent upon the presence of humans. It died out soon after all the people were evacuated from Hirta in 1930. The St Kilda wren (Troglodytes troglodytes hirtensis) is confined to the St. Kilda Islands, where the total population is around 500. Fair Isle is a tiny island located between the main Shetland Islands and Orkney. It is the most remote inhabited island in the United Kingdom. The Fair Isle wren (Troglodytes troglodytes fridariensis) has a breeding population of from 10 to 50 pairs adapted to life on boulder beaches. The slightest environmental interference would likely prove to be devastating. Ireland Ireland is located west of Great Britain and is the second largest of the British Isles. As with Great Britain all of the threatened vertebrate species are freshwater fishes. The pollan (Coregonus pollan) is a type of whitefish confined to just five lakes (Lough Neagh, lower Lough Erne, Lough Ree, Lough Derg, and Lough Allen) in central Eire and Northern Ireland. However, the only remaining sustainable populations are those of Lough Neagh and Lough Allen, the rest relying upon continued restocking. The blunt-snouted char (Salvelinus obtusus) was historically found in a number of lakes in eastern and south-western Eire, but has disappeared from most if not all of them due to eutrophication and pollution. It was last recorded from Lough Muckross in 1904, Lough Tay in 1908, Lough Dan in 1988, and Lough Leane in 1999. Surveys in Lough Accose in 1983, meanwhile, failed to find any char. Lough Melvin is located in north-western Ireland, on the border between Eire and Northern Ireland. Gray’s char (Salvelinus grayi) is confined to Lough Melvin, where it has been declining for decades due to eutrophication and introduced species. Two species of trout, the gillaroo (Salmo stomachicus), and the sonaghen (S. nigripinnis) are also endemic to Lough Melvin. Lough Coomasaharn is a small oligotrophic lake located in south-western Ireland (Eire). The Coomsahar char (Salvelinus fimbriatus) is confined to Lough Coomasaharn. Lough Leane is located in south-western Ireland (Eire). The Killarney shad (Alosa killarnensis) is confined to Lough Leane, where it is seriously threatened by eutrophication and introduced species.

Sakhalin Sakhalin is a large island located in the northern Pacific off the eastern coast of Russia and north of Japan. The Sakhalin musk deer (Moschus moschiferus sachalinensis) is confined to Sakhalin.

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The Palearctic Realm

The Sakhalin vole (Microtus sachalinensis) is confined to northern and central parts of the island, where it may be potentially affected by oil infrastructure development projects.

Bering Island Bering Island is located in the Bering Sea off the Kamchatka Peninsula. The spectacled cormorant (Phalacrocorax perspicillatus), the largest known species of cormorant, was almost flightless and therefore easy prey for hunters. It was exterminated by about 1852.

Sicily Located south of the Italian Peninsula, Sicily (Sicilia in Italian) is the largest island in the Mediterranean Sea. It is an often-quoted example of man-made deforestation, which has occurred since Roman times when the island was turned into an agricultural region. This gradually dried the climate, leading in turn to a decline in rainfall and the drying up of rivers. The central and western parts are practically devoid of forest. Not surprisingly, there is little in the way of endemic fauna remaining.

The Sicilian grey wolf (Canis lupus cristaldii) was a slender, short-legged subspecies that was likely driven extinct due to human persecution in the 1920s, although there were several possible sightings up until the 1970s. The Sicilian pond turtle (Emys trinacris) is confined to the remaining wetlands of Sicily, where it appears to be relatively abundant.

Corsica and Sardinia The islands of Corsica and Sardinia and here considered together, as they are geographically close and share much of the same fauna. The Corsican red deer (Cervus elaphus corsicanus) is a small subspecies that is thought to have been introduced to Corsica and Sardinia by humans around 8000 years ago. It underwent a dramatic decline on Corsica and was extirpated there in 1969, after which the only remaining population (around 100) lived on Sardinia. Captive breeding and better protection enabled the animals to slowly recover on the latter island and, eventually, to be successfully reintroduced to Corsica. The Sardinian pika (Prolagus sardus), a type of small, tailless mammal, historically occurred on Corsica, Sardinia, and a few satellite islands. Last seen in 1774, it is considered to be extinct. Bedriaga’s rock lizard (Archaeolacerta bedriagae) has a highly fragmented distribution on Corsica, Sardinia, and a few satellite islands. Corsica Corsica (Corse in French) is located south-east of France and west of Italy. Roughly two-thirds of this large island is comprised of a single mountain chain. The Corsican nuthatch (Sitta whiteheadi) is confined to areas of Corsican pine, which occurs in fragments on the island’s mountain ridges. The Corsican painted frog (Discoglossus montalentii) is endemic to the central mountains.

Figure 2.7 Spectacled cormorant. (Credit: Joseph Wolf.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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Sardinia Sardinia (Sardegna in Italian) is located south of Corsica. It is the second largest island in the Mediterranean Sea. The Sardinian long-eared bat (Plecotus sardus) is confined to forest fragments on Sardinia. The Sardinian barred grass snake (Natrix helvetica cetti) is confined to Sardinia. The Sardinian brook salamander (Euproctus platycephalus) is a rare species found only in eastern Sardinia between the Limbara Mountains in the north and the Sette Fratelli Mountains in the south. Several species of European cave salamander (Speleomantes) are endemic to various areas of Sardinia, where they are threatened by loss of habitat and illegal collection for the international pet trade. The Supramonte cave salamander (S. supramontis) is confined to central-eastern Sardinia. Gene’s cave salamander (S. genei) occurs in south-western Sardinia.

The Eurasian Region

The imperial cave salamander (S. imperialis) is confined to central and eastern Sardinia. The Monte Albo cave salamander (S. flavus) is found within the Monte Albo Mountains of north-eastern Sardinia. The Sarrabus cave salamander (S. sarrabusensis) is found around Monte dei Sette Fratelli in extreme south-eastern Sardinia.

The Aegean Islands The Aegean Islands are located in the Aegean Sea between Greece and Turkey. They are traditionally comprised of seven different island groups. Euboea Euboea (Evvoia in Greek) is the second largest island in Greece, after Crete. The Manikiotikos barbel (Barbus euboicus) is a type of freshwater fish confined to a single stream on the southern part of Euboea, where during the dry season it is often reduced to only a few intermittent pools. The Sporades Islands The Sporades (Vóries Sporádhes in Greek) are an archipelago along the eastern coast of Greece, north-east of the island of Euboea in the Aegean Sea. They consist of 24 islands, only 4 of which are permanently inhabited. Two subspecies of Sporades wall lizard (Podarcis gaigeae) are endemic to a few of the Sporades Islands. The Skyros wall lizard (P. g. gaigeae) is confined to Skyros and associated islets, while Weigand’s wall lizard (P. g. weigandi) is confined to the island of Piperi. Crete Crete (Kríti in Greek) is located in the southern part of the Aegean Sea and is the largest of the Greek islands. It is elongated in shape and mountainous, with a large number of islets surrounding the coast. The island has long been isolated from mainland Europe, Asia, and Africa, and this is reflected in the diversity of its fauna. Dwarf forms of elephant, mammoth, hippopotamus, and deer, along with giant flightless owls, were all native here during the Pleistocene Period. The Cretan shrew (Crocidura zimmermanni) is known from a few localities in the mountains. Two species of wall lizard (Podarcis) are endemic to Crete and its satellite islands. The Cretan wall lizard (P. cretensis) is found on western Crete and a few small islets. The Leventis wall lizard (P. levendis) is confined to the uninhabited islets of Pori and Lagouvardos, north of the island of Antikythira, where its total population is presumably very small. The Cretan water frog (Pelophylax cretensis) is patchily distributed in the lowlands and not particularly abundant. The Cyclades The Cyclades (Kiklaoes in Greek) are a group of some 220 small islands in the Aegean Sea, south-east of mainland Greece.

Three subspecies of Cyclades wall lizard (Podarcis milensis) are endemic to various islands. The Milos wall lizard (P. m. milensis) is confined to Milos. Adolf Jordans’ wall lizard (P. m. adolfjordansi) is confined to Ananes Island west of Milos. The Gerakunia wall lizard (P. m. gerakuniae) is confined to Gerakunia (Falconera) and Velopoula. Schweizer’s blunt-nosed viper (Macrovipera schweizeri) is confined to the western Cycladic islands of Milos, Kimolos, Polyaigos, and Syphnos. The Dodecanese Islands The Dodecanese (Dodekánisa in Greek) are a group of 15 larger plus 150 smaller islands located in the south-eastern Aegean Sea, off the coast of Turkey. Helversen’s Lycian salamander (Lyciasalamandra helverseni) is confined to the islands of Karpathos, Kasos, and Saria, where it is still fairly common within its restricted range. Rhodes (Ródos in Greek) is the largest of the Dodecanese Islands. The Rhodes minnow (Ladigesocypris ghigii) is endemic to freshwater streams, springs, marshes, reservoirs, and pools on Rhodes, where it is threatened by habitat destruction. Karpathos is located about 47 km south-west of Rhodes. The Karpathos water frog (Pelophylax cerigensis) is only known with certainty from a single river in the mountains of Karpathos. Kastellorizo lies roughly 2 km off the south coast of Turkey and some 125 km east of Rhodes. The Kastellorizo Lycian salamander (Lyciasalamandra luschani basoglui) is confined to Kastellorizo.

The Balearic Islands The Balearic Islands (Islas Baleares in Spanish) are an archipelago of around 50 islands in the western Mediterranean near the eastern coast of the Iberian Peninsula. The larger islands (Mallorca, Menorca, Ibiza, and Formentera) are popular tourist destinations, although many of the smaller islands and islets are uninhabited. The Balearic shearwater (Puffinus mauretanicus) is a type of seabird that breeds exclusively in the Balearic Islands. Lilford’s wall lizard (Podarcis lilfordi) was historically found on the larger islands of Mallorca, Menorca, and Ibiza, as well as in the Cabrera Archipelago. It is believed that the introduction of cats and other predators was responsible for the extirpation of the species from the main islands, but a number of subspecies still survive on various rocky islets. These will be discussed below. Mallorca Mallorca (also known as Majorca) is the largest of the Balearic Islands. The Sargantana wall lizard (Podarcis lilfordi sargantanae) is confined to four islets off the northern coast of Mallorca (Sargantana, Ravells, Bledes, and Tusqueta).

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The Palearctic Realm

The Mallorcan midwife toad (Alytes muletensis) is confined to the Sierra Tramuntana in northern Mallorca. Dragonera is an islet off the northern coast of Mallorca. The Dragonera wall lizard (Podarcis lilfordi gigliolii) is confined to Dragonera. Toro is an islet off the coast of Mallorca. The Toro wall lizard (Podarcis lilfordi toronis) is confined to Toro. La Guardia is an islet off the coast of Majorca. The La Guardia wall lizard (Podarcis lilfordi jordansi) is confined to La Guardia. Malgrats is an islet off the south-western coast of Mallorca. The Malgrats wall lizard (Podarcis lilfordi hartmanni) is confined to Malgrats. Menorca Menorca (also known as Minorca) is located north-east of Mallorca. Ratas was a tiny, rocky islet within the bay of Mahón, Menorca. The Ratas wall lizard (Podarcis lilfordi rodriquezi) was confined to Ratas Island. It went extinct in 1950 after the island was destroyed during harbour reconstruction. Rey is another tiny islet located within the bay of Mahón, Menorca. The Rey wall lizard (Podarcis lilfordi hospitalis) is confined to Rey Island. Another subspecies, Bedriaga’s wall lizard (P. l. balearica), was historically endemic to Rey Island but has been introduced to Minorca. Addaya is an islet off the eastern coast of Menorca. The Addaya wall lizard (P. l. addayae) is confined to Addaya. Aire is an islet off the south-eastern coast of Menorca. The Aire wall lizard (P. l. lilfordi) is confined to Aire. Colom is an islet off Menorca. The Colom wall lizard (P. l. brauni) is confined to Colom. Carbonera is an islet off Menorca. The Carbonera wall lizard (Podarcis l. carbonerae) is confined to Carbonera. Colomer is an islet off the north-eastern coast of Menorca. The Colomer wall lizard (P. l. colomi) is confined to Colomer. Binicondrell is an islet off the southern coast of Menorca. The Binicondrell wall lizard (P. l. codrellensis) is confined to Binicondrell. Sanitja is an islet off northern Menorca. The Sanitja wall lizard (P. l. fenni) is confined to Sanitja. Porros is an islet off the northern coast Menorca. The Porros wall lizard (P. l. porrosicola) is confined to Porros. Ibiza Ibiza is the third largest of the Balearic Islands. The Ibiza wall lizard (P. l. zenonis) was historically confined to Ibiza, where it went extinct at some unknown date after the introduction of invasive predators. The Cabrera Archipelago The Cabrera Archipelago is located south of Majorca. The Cabrera wall lizard (P. l. kuligae) is confined to Cabrera, Fonoi Gros, Fonoi Petit, and Ses Rates. The Na Redonda wall lizard (P. l. conejerae) is confined to Na

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Rodonda. The Xapat Gros wall lizard (P. l. xapaticola) is confined to Xapat Gros, Xapat Petit, and La Teula islands. The L’Esponge wall lizard (P. l. espongicola) is confined to L’Esponge. The Na Pobra wall lizard (P. l. pobrae) is confined to Na Pobra. The L’Imperial wall lizard (P. l. imperialensis) is confined to L’Imperial. The Na Plana wall lizard (P. l. planae) is confined to Na Plana. The Sas Bledas wall lizard (P. l. nigerrima) is confined to Sas Bledas. The Horadada wall lizard (P. l. fahrae) is confined to Horadada. The Estel de Fora wall lizard (P. l. estelicola) is confined to Estel de Fora.

The Aeolian Islands The Aeolian Islands (Isole Eolie in Italian) are a volcanic archipelago in the Tyrrhenian Sea, north of Sicily. The Aeolian wall lizard (Podarcis raffonei) is confined to a few isolated areas on Volcano Island, along with a few small rocky islets (Strombolicchio, La Canna, and Scoglio Faraglione).

Miscellaneous Islands Santo Stefano Island (Isola di Santo Stefano in Italian) is located off the western coast of Italy. The Santo Stefano wall lizard (Podarcis siculus sanctistephani) was confined to Santo Stefano, where it became extinct in 1965 most likely due to predation by feral cats and other factors. Corfu (Kerkyra in Greek) is located off the north-western coast of Greece. The Corfu dwarf goby (Knipowitschia goerneri) was long known only from a single freshwater spring, where it was last recorded in 1983. Surveys in the 1990s failed to find any there. In 2014 nine specimens were collected from Korission Lagoon in southern Corfu. The Berlengas Archipelago is a group of small islands off the coast of western Portugal. The Berlengas wall lizard (Podarcis carbonelli berlengensis) is confined to the Berlenga Islands. Shedao Island is located off the coast of Liaotung, northeastern China. The Shedao pit viper (Gloydius shedaoensis) is confined to Shedao Island. Paramushir Island is located in the northern Kuril Islands off the Kamchatka Peninsula. The Paramushir shrew (Sorex leucogaster) is confined to Paramushir Island.

Balance for the Eurasian Region Widely dispersed, isolated finds of fossils and stone artefacts suggest that Homo erectus had migrated across Eurasia from Africa by around three million years ago, but he and his eventual successor Heidelberg Man (H. heidelbergensis) apparently remained quite rare. Between 600,000 and 350,000 years ago Neanderthals (H. neanderthalensis) first emerged, and are considered to be the first modern Europeans and the first to leave behind a substantial tradition, as evidenced by their cave paintings and burial practices. Modern humans (H. sapiens) arrived in Mediterranean Europe and southern Siberia from

The Eurasian Region

the Levant between 45,000 and 43,000 years ago, whereupon both species coexisted for several thousand years until the assimilation or extinction of the Neanderthals between 40,000 and 28,000 years ago. Humans subsequently proceeded to populate the entire continent and advanced north, following the retreating ice sheets of an ice age which spanned from 26,500 to 19,000 years ago and surviving as hunter-gatherers. After the last ice age ended around 12,500 B . C . temperatures and sea levels began to rise, changing the environment and creating the British Isles. Finally, about 8000 years ago a wave of ‘farmers’ arrived from the Near East and permanent settlement began. Despite this long human settlement, particularly in western Eurasia, there had been relatively little environmental destruction within the Eurasian Region up until the modern era. While it is true that the Mediterranean countries had suffered almost complete forest loss and later soil erosion due to goats, an ecological balance was more or less maintained. Beginning in the late sixteenth century, Russian explorers began to probe ever further into the vast Siberian wilderness, reaching the Lena River by the 1620s, Lake Baikal and Sakhalin by 1643, and the Kuril Islands by 1706. However, it was not until the nineteenth century that we begin to see a major human impact. Indeed, it was primarily in Europe that

the concept of conservation first originated. In 1810, the English poet William Wordsworth described the Lake District as ‘a sort of national property, in which every man has a right and interest who has an eye to perceive and a heart to enjoy’. Even earlier, the Naples government had undertaken laws to protect natural areas, which could be used as a game reserve by the royal family. Over the past few decades there has been an ever-growing awareness of environmental issues throughout Europe and ambitious attempts at rewilding more remote areas. In recent historical time (i.e. since A . D . 1500), the Eurasian Region has lost at least 18 species/12 subspecies of vertebrates. Among the extinct forms 1 species/9 subspecies are mammals, 2 species are birds, 3 subspecies are reptiles, and 15 species are freshwater fishes. Another 6 species are possibly extinct. In addition, there are 459 species/94 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 55 species/47 subspecies are mammals, 19 species/5 subspecies are birds, 42 species/33 subspecies are reptiles, 38 species/7 subspecies are amphibians and 305 species/2 subspecies are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

1 species

~ species

~ species

55 species

9 subspecies

~ subspecies

~ subspecies

47 subspecies

10 taxa

~ taxa

~ taxa

104 taxa

2 species

~ species

~ species

19 species

~ subspecies

~ subspecies

~ subspecies

5 subspecies

2 taxa

~ taxa

~ taxa

24 taxa

~ species

~ species

~ species

42 species

3 subspecies

~ subspecies

~ subspecies

33 subspecies

3 taxa

~ taxa

~ taxa

75 taxa

~ species

~ species

~ species

38 species

~ subspecies

~ subspecies

~ subspecies

7 subspecies

~ taxa

~ taxa

~ taxa

45 taxa

15 species

6 species

~ species

305 species

~ subspecies

~ subspecies

~ subspecies

2 subspecies

15 taxa

6 taxa

~ taxa

307 taxa

18 species

6 species

~ species

459 species

12 subspecies

~ subspecies

~ subspecies

94 subspecies

30 taxa

6 taxa

~ taxa

553 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

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The Palearctic Realm

The Sino-Himalayan Region The Sino-Himalayan Region extends from the Tibetan Plateau and the Himalayas through the temperate and subtropical, mostly mountainous areas of northern India, China, and northern Indochina to the Japanese Archipelago and Taiwan. It is bordered on the north by the drier parts of the Eurasian Region and on the south by the lowland rainforests and grasslands of the Indo-Malaysian Realm. The South China tiger (Panthera tigris amoyensis) was formerly distributed in a large part of eastern and central China northward to 38–40° N. latitude. Heavily hunted for decades, it is now the most critically endangered subspecies of tiger and, indeed, one of the most threatened animals in the world. Despite unconfirmed reports and some evidence of footprints, there have been no confirmed sightings since the 1970s, leading experts to consider it already functionally extinct in the wild. The entire known population of roughly 65 individuals is currently held in captivity. The Himalayan wild dog (Cuon alpinus laniger) occurs in southern Tibet, Nepal, Sikkim, Bhutan, and Kashmir. The thin-tailed wild dog (C. a. lepturus) is found in China south of the Yangtze River. Both are threatened by loss of habitat, depletion of their prey base and human persecution. The Indochinese clouded leopard (Neofelis nebulosa) is a medium-sized wild cat found patchily over much of southern and eastern Asia, where it has long been hunted for its beautiful fur. The western black crested gibbon (Nomascus concolor) lives primarily in the mountainous areas of southern China, northern Vietnam, and northern Laos, and is seriously threatened by habitat destruction and hunting. The Tonkin black crested gibbon (N. c. concolor) is found in southern China (south-western Yunnan) and northern Vietnam (Lao Cai, Yen Bai, Son La, and Lai Chau provinces), roughly between the Black and Red rivers. The Tonkin snub-nosed monkey (Rhinopithecus avunculus) has long been confined to areas east of the Red River in far north-eastern Vietnam. Since the mid-twentieth century it has suffered a serious decline due to loss of habitat and hunting, and is now confined to a few areas of steep, karstic forest in Tuyen Quang, Bac Kan, Ha Giang, and Thai Nguyen provinces. In 2008 fewer than 250 were thought to exist. The white-headed langur (Trachypithecus leucocephalus) is a type of leaf-eating monkey found patchily within a small area of south-western China (Guangxi), where the total population is thought to be less than 250. Shortridge’s langur (T. shortridgei) occurs in south-western China (Yunnan) and north-eastern Myanmar. Both are threatened by loss of habitat and hunting for food as well as use in ‘traditional medicine’. Perhaps the most dramatic history of any animal faced with extinction is that of Père David’s deer (Elaphurus davidianus). In ancient times this deer had a wide range in China and occurred in Japan as well, but as early as the Shang

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dynasty (1766–1122 B . C .) it seems to have been exterminated in the wild. Like so many other deer in China, however, it survived in parks. It was in one of these reserves, the imperial hunting park near Peking (present-day Beijing), that a French missionary, Father David, came upon the last remaining herd. Father David managed to send a number of the animals to European zoos, so that by about 1870 there were small herds at several zoological gardens. This came none too soon, because when floods broke through the imperial walls of the hunting park in 1894, many deer escaped and were killed by peasants. And in 1900 foreign troops sent to Peking during the Boxer Rebellion killed almost all the deer. The few that remained were sent to the Peking Zoo, but by 1921 these, too, were dead. After the events of 1900, the eleventh Duke of Bedford at Woburn Abbey in Bedfordshire, England undertook to help the deer by collecting a herd of 16 animals from European zoos. Today, all survivors derive from this herd. By the early 1960s there were some 400 deer in over 40 zoos all over the world, including Beijing. In the mid-1980s the species was reintroduced into semi-captive facilities in China, prior to being established in protected areas of intact coastal grassland. The sambar deer (Rusa unicolor) is a large species found over a wide area of southern and south-eastern Asia, where it is divided into a number of subspecies. Dejean’s sambar deer (R. u. dejeani) is confined to southern and south-western China. The Chinese goral (Naemorhedus griseus) is a small, goatlike ungulate found in the mountainous regions of China,

Figure 2.8 Père David’s deer. (Credit: Josèphe Huët.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The Sino-Himalayan Region

Myanmar, India, Thailand, Vietnam, and possibly Laos. It is threatened by overhunting. Milne-Edwards’ serow (Capricornis milneedwardsii) is a goat-like species found widely but patchily in forests and rocky areas throughout Central and South East Asia, where it is threatened by habitat destruction and hunting. The pangolins or scaly anteaters are a group found throughout the tropical and subtropical regions of Africa and southern Asia. They are among the most illegally traded animals in the world today, as their protective scales and other body parts are highly valued for use in ‘traditional medicine’. The Chinese pangolin (Manis pentadactyla) is still found over much of the northern Indian subcontinent, northern South East Asia and southern China (including Taiwan and Hainan), but has been extirpated from many areas. Reeves’ pheasant (Syrmaticus reevesii) is endemic to temperate forest fragments in central and north-eastern China, where it is threatened by habitat destruction, hunting, and overcollection of its eggs. The species has been successfully introduced to various parts of Europe and to Hawaii. Elliot’s pheasant (S. ellioti) is endemic to south-eastern China (Guizhou, Hubei, Anhui, Zhejiang, Fujian, Jiangxi, Hunan, Guangxi, and Guangdong), where it lives in both lowland and montane forests. While not immediately threatened, habitat destruction is a concern, as is hunting for food. The brown-chested jungle flycatcher (Cyornis brunneatus) is a rare species that breeds in the lowland forests of southeastern China and winters in the southern Malay Peninsula. The fairy pitta (Pitta nympha) is a brightly coloured passerine bird that breeds in Japan, South Korea, south-eastern China and Taiwan, from where it migrates in winter mainly to the island of Borneo. The total population is very small and seriously threatened by loss of habitat and trapping for the cage-bird trade. The Naung Mung wren-babbler (Rimator naungmungensis) is known only from a small area of far northern Myanmar (Kachin state). The Chinese grass-babbler (Graminicola striatus) is found disjunctly in wetland areas across much of South and South East Asia, with small resident populations in central and south-eastern China (Guangxi, Guangdong, Hainan, and Hong Kong) and central Myanmar. Vagrants have been found in Thailand, Vietnam, and Cambodia. It is everywhere under threat by loss of habitat. The beautiful nuthatch (Sitta formosa) is a rare species found patchily over a wide area of north-eastern India (Arunachal Pradesh, Assam, Nagaland, and West Bengal), Bhutan, northern and eastern Myanmar, south-western China (Yunnan), and extreme north-western Thailand. It is threatened by the loss of large trees upon which it depends. Several species and subspecies of Asian box turtle (Cuora) number among the most heavily trafficked and seriously threatened in the world, being much sought after both by the international pet trade and for use in ‘traditional medicine’. Zhou’s box turtle (C. zhoui) is known only from Chinese food

market specimens and from a relatively small number of captive individuals in Germany and the United States. The species appears most likely to originate from a very small area of extreme northern Vietnam and possibly adjacent southern China (south-eastern Yunnan or western Guangxi), although it has never been scientifically recorded in the wild. The Chinese yellow-headed box turtle (C. aurocapitata) is known only from a small area of central-eastern China (Anhui). McCord’s box turtle (C. mccordi) was originally described in 1988 from a single specimen obtained from the Chinese pet trade. It was unknown from the wild until 2007, when a small population was discovered in southern China (central Guangxi). Pan’s box turtle (C. pani) is confined to central China (Shaanxi, Sichuan, and Hubei). The Chinese threestriped box turtle (C. trifasciata) was historically widespread and common in southern China (Guangxi, Guangdong, and Fujian) as well as on the islands of Hainan, Macau, and Hong Kong (the latter two populations most likely having been introduced). The species has been extirpated from mainland China, and now survives only on the aforementioned islands. The Vietnamese three-striped box turtle (C. cyclornata) was historically found in southern China (Guangxi), northern Vietnam, and northern Laos. Nearly extinct in the wild, it is well-established on Chinese turtle farms. The yellow-margined box turtle (C. flavomarginata) as a species is widespread in central and eastern China (Hunan, Henan, Anhui, Hubei, Chongging, Sichuan, Zhejiang, and Jiangsu) as well as Taiwan and southern Japan. The Chinese yellow-margined box turtle (C. f. flavomarginata) is the subspecies found over the majority of this range, but is everywhere seriously threatened. Bourret’s skink (Sphenomorphus tritaeniatus) is known only from two localities within the Red River delta region of northern Vietnam. The Chinese cobra (Naja atra) is still found over a relatively wide area of south-eastern China (including the islands of Hainan, Hong Kong, and Macao), northern Vietnam, and northern Laos. It is threatened by human persecution and overcollection for use in ‘traditional medicine’. The Assamese bug-eyed tree frog (Theloderma moloch) is known only from a few localities in north-eastern India (Arunachal Pradesh and Assam). Khare’s gliding frog (Pterorana khare) in known only from a few disjunct localities in north-eastern India (Assam and Nagaland).

Mountains and Highlands Mountains and highlands within the Sino-Himalayan Region include the Himalayas along with certain of the southern foothills, as well as the Tibetan and Yunnan-Guizhou plateaus and various smaller mountain ranges and plateaus in China, northern Myanmar, and northern Vietnam. The golden snub-nosed monkey (Rhinopithecus roxellana) is found in the temperate mountainous forests of central China, where it is divided into three subspecies. The Moupin golden

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The Palearctic Realm

snub-nosed monkey (R. r. roxellana) is confined to western Sichuan, southern Gansu, and southern Shaanxi. The Myanmar snub-nosed monkey (R. strykeri) was unexpectedly discovered in 2010 in high-altitude forest in northern Myanmar (Kachin State), and has since been found in the Gaoligong Mountains of neighbouring south-western China (Yunnan). The species is seriously threatened by loss of habitat and hunting. The iconic giant panda (Ailuropoda melanoleuca) was historically widespread throughout central and south-eastern China, being found as far north as Beijing and south into parts of South East Asia. The better-known black and white form (A. m. melanoleuca) now has a very restricted and fragmented range in the high mountains of Sichuan, Shaanxi, and Gansu provinces, where it exists exclusively in middle-elevation forests with a dense understory of bamboo. The largest populations are currently found in the Min and Qionglai ranges, with smaller, more isolated ones remaining in the Liangshan, Daxiangling, and Xiaoxiangling mountains. The takin (Budorcas taxicolor) is a muskox-like goat-antelope found in heavily forested montane areas of north-eastern India (Arunachal Pradesh and Sikkim), Bhutan, western China (Gansu, Sichuan, Shannxi, Tibet, and Yunnan), and northern Myanmar. Several subspecies are threatened by hunting and habitat destruction. The Mishmi takin (B. t. taxicolor) is found in western China (Tibet and Yunnan) and northern Myanmar. The Sichuan takin (B. t. tibetana) is found in Tibet and western China (Yunnan). The Tibetan argali (Ovis ammon hodgsoni) is a type of wild sheep found in western and central China. It is threatened by poaching and competition with introduced livestock. The red goral (Naemorhedus baileyi) is a type of goatantelope confined to the temperate mountains of southwestern China (Tibet and Yunnan), north-eastern India (Arunachal Pradesh), and northern Myanmar. It is threatened by habitat destruction and illegal hunting. Two subspecies of elk or wapiti (Cervus canadensis) occurring on the Tibetan Plateau and nearby regions are also threatened. MacNeil’s deer (C. c. macneilli) has long been intensively hunted for its antler velvet, which is believed to have aphrodisiac properties. The Tibetan deer (C. c. wallichi) of south-eastern Tibet and Bhutan was thought to be extinct until its rediscovery in 1995. The South China sika deer (Cervus nippon kopschi) was historically found throughout the Yangtze River drainage east to the coast and as far south as the border with Vietnam. Today, the surviving population of around 300 is widely scattered across its former range in remote mountains isolated by heavily populated lowlands. About 30 exist in the Tianmu Mountains of northern Zhejiang, 70–100 in southern Anhui, and 150 in northern Jiangxi. The population size in southern Guangxi is unknown, and a tiny number may still exist in northern Guangdong as well. Ongoing poaching as well as inbreeding are the main threats. The tufted deer (Elaphodus cephalophus) is a small, muntjaclike species historically found widely within central and southern

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China and northern Myanmar, where it has suffered significant declines due to hunting. The south-western tufted deer (E. c. cephalophus) is found in south-western China and (at least historically) northern Myanmar, although it has perhaps been extirpated from the latter country. The south-eastern tufted deer (E. c. michianus) is confined to south-eastern China. The central tufted deer (E. c. ichangensis) is found in central China. The black muntjac (Muntiacus crinifrons) is a type of deer confined to the montane forests of Zhejiang, Anhui, Jiangxi, and Fujian in south-eastern China. Up until 1975 it was known only from a few specimens and was considered the rarest deer in the world. It has since been shown to be more populous than previously thought but nevertheless threatened. Heavily harvested throughout the twentieth century (at least 2000 were killed in 1978 alone), the current population is estimated at between 5000 and 10,000. Several species and subspecies of musk deer (Moschus) are seriously threatened by hunting for use in ‘traditional medicine’, as well as loss of habitat. The black musk deer (M. fuscus) is found in south-western China (Yunnan and Tibet), northern Myanmar, Bhutan, north-eastern India (Arunachal Pradesh), and eastern Nepal. The dwarf musk deer (M. berezovskii) is found widely in the mountains of central and southern China, northern Vietnam, and possibly northern Laos. A number of ill-defined subspecies have been described, including Berezovski’s dwarf musk deer (M. b. berezovskii), the Bijang dwarf musk deer (M. b. bijanensis), the Cao Bang dwarf musk deer (M. b. caobangis), and the Yunnan-Guizhou dwarf musk deer (M. b. yanguiensis). The alpine musk deer (M. chrysogaster) is found in the mountains of northern and central China. The red panda (Ailurus fulgens) is a racoon-like species that, despite its name, is not closely related to the giant panda. It is native to the eastern Himalayas and the mountains of south-western China, where it is usually divided into two subspecies. The Himalayan red panda (A. f. fulgens) is found in north-eastern India (Sikkim, Assam, West Bengal, and Arunachal Pradesh), Nepal, and Bhutan, while Styan’s red panda (A. f. styani) is found in northern Myanmar (Kachin State) and south-western China (Sichuan, Yunnan, and Tibet). Both are threatened by habitat destruction. The Sichuan weasel (Mustela russelliana) is known only from three collection records within a small area of northcentral China (Sichuan). The Muong Bang stripe-bellied rat (Pseudoberylmys muongbangensis) is known only from a single locality in northern Vietnam (Son La province). The Sichuan dormouse (Chaetocauda sichuanensis) is known only from five specimens collected from the mountains of central China (northern Sichuan). Crump’s mouse (Diomys crumpi) is known only from a few widely separated localities in northern India (Bihar and Manipur), southern Nepal, and northern Myanmar. The Duke of Bedford’s vole (Proedromys bedfordi) is known only from a small area in the mountains of central China (southern Gansu and northern Sichuan).

The Sino-Himalayan Region

The dusky shrew (Sorex sinalis) is known only from a few mountaintop localities spread over a relatively wide area of central China (Gansu, Sichuan, and Shaanxi). The Gansu shrew (S. cansulus) is known only from a few areas of Gansu, Qinghai, and Tibet. The dusky tube-nosed bat (Murina fusca) is known only from a single locality in north-eastern China (Heilongjiang). Anthony’s pipistrelle bat (Hypsugo anthonyi) is known only from a single specimen collected in the 1940s in northeastern Myanmar. Several species and subspecies of eared pheasant (Crossoptilon) are endemic to small areas of China. The brown-eared pheasant (C. mantchuricum) is confined to a few scattered areas of montane forest in northern central China (Shanxi, Hebei, and Shaanxi). Those populations outside of protected areas are threatened by habitat destruction. The white-eared pheasant (C. crossoptilon) is found in westcentral and south-western China, where it is divided into three subspecies. The Sichuan white-eared pheasant (C. c. crossoptilon) is confined to Sichuan. The Yunnan whiteeared pheasant (C. c. lichiangnse) is confined to Yunnan. Drouyn’s white-eared pheasant (C. c. drouyni) is found in Tibet and Qinghai. All are threatened by deforestation and hunting for food. Cabot’s tragopan (Tragopan caboti) is a type of pheasant found patchily in the montane forests of south-eastern China, where it is divided into two subspecies. The eastern Cabot’s tragopan (T. c. caboti) occurs in Fujian, Jiangxi, Zhejiang, and Guangdong, while the western Cabot’s tragopan (T. c. guangxiensis) is confined to north-eastern Guangxi and southern Hunan. The grey-bellied tragopan (T. blythii) is divided into two subspecies. Blyth’s grey-bellied tragopan (T. b. blythii) occurs in the Himalayas of north-eastern India through south-western China and north-western Myanmar. All are threatened by loss of habitat and high hunting and snaring pressure. Sclater’s monal (Lophophorus sclateri) is a type of pheasant divided into two subspecies. The eastern Sclater’s monal (L. s. sclateri) is found in the Himalayas of north-eastern India, northern Myanmar, and south-western China (Tibet and Yunnan). The western Sclater’s monal (L. s. arunachalensis) is confined to north-eastern India (Arunachal Pradesh). The Sichuan partridge (Arborophila rufipectus) is confined to a few montane forest fragments in central China (southcentral Sichuan and possibly north-eastern Yunnan). It is threatened by hunting and habitat destruction. The grey-sided thrush (Turdus feae) breeds in a small mountainous area of north-eastern China (Shanxi, Hebei, and Beijing), from where it migrates in winter to north-eastern India, Bangladesh, Myanmar, Thailand, and Laos. It is threatened by deforestation. The white-spotted laughingthrush (Garrulax bieti) is confined to the mountains of central China (north-western Yunnan and south-western Sichuan).

The rufous-headed robin (Larvivora ruficeps) breeds only in a few areas of northern China (north-central Sichuan and southern Shaanxi), from where it migrates to the Malay Peninsula and Cambodia. It is threatened by forest loss and overcollection for use as cage birds. The black-throated blue robin (Calliope obscura) was first discovered in 1891 in the mountains of north-central China (Sichuan, Gansu, and Shaanxi). Only a handful of individuals were seen again until 2011, when its breeding grounds were discovered in the Qinling Mountains. The species is thought to migrate to southern China and northern Thailand. The grey-hooded parrotbill (Sinosuthora zappeyi) is confined to the mountains of central China (south-central Sichuan and western Guizhou). The Emei liocichla (Liocichla omeiensis) is a type of passerine bird known from the mountains of central China (south-central Sichuan and extreme north-eastern Yunnan). The golden-fronted fulvetta (Schoeniparus variegaticeps) is a type of passerine bird known only from a few localities in the mountains of south-central China (Sichuan and Guangxi). The giant nuthatch (Sitta magna) was historically found throughout the mountains of south-western China (Yunnan), north-western Thailand, and central and eastern Myanmar, but may now be extirpated from the latter country. It is threatened by the loss of large trees upon which it depends. The Yunnan gecko (Gekko scabridus) is a poorly known species from the mountains of south-western China (Sichuan and Yunnan). The Sichuan pit viper (Sinovipera sichuanensis) is known only from two specimens collected in western China (southern Sichuan). The Sichuan rat snake (Euprepiophis perlacea) is known only from a few mountainous areas of western China (western Sichuan). Angel’s keelback (Rhabdophis angeli) is a type of snake known only from a few specimens collected in the 1930s from a small area of northern Vietnam (Thai Nguyen and Vinh Phuc provinces). The Sichuan hot-spring keelback (Thermophis zhaoermii) is confined to a small area of central China (Sichuan). The Chapa flat-nosed keelback (Hebius chapaensis) is known only from a small area of montane forest in northern Vietnam, where it was last reported in the early twentieth century. The Wa keelback (H. metusium) is known only from two localities in central China (Sichuan). The Yunnan mountain snake (Plagiopholis unipostocularis) is known only from a single specimen collected from an imprecise locality in southern China (Yunnan). The Ningshan kukri snake (Stichophanes ningshaanensis) is known only from a single locality in central China (Shaanxi). Lacroix’s kukri snake (Oligodon lacroixi) is known only from three localities across a relatively wide area of southwestern China (Yunnan and Sichuan) and northern Vietnam. Blyth’s reticulated snake (Blythia reticulata) appears to have been found historically across a wide area of north-

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eastern India (Arunachal Pradesh, Assam, Manipur, and Meghalaya), northern Myanmar, and possibly southern China (Tibet), although in recent decades it has only been recorded from Myanmar (Chin State). The Tam Dao stream snake (Opisthotropis tamdaoensis) is known only from Tam Dao National Park in northern Vietnam (Vinh Phuc province). The Lichuan bell toad (Bombina lichuanensis) is known only from two localities in northern China (Hubei and Sichuan). It probably occurs more widely, especially in area between the two known sites, but is nevertheless thought to be threatened by loss of habitat. Several lazy toads of the genus Oreolalax are endemic to small areas of China and Indochina. The Liangbei lazy toad (O. liangbeiensis) is known only from a single small stream in central China (southern Sichuan). Weigold’s lazy toad (O. weigoldi) is known only from a single specimen collected in the 1920s from an imprecise locality in central China (Sichuan). The Chuanbei lazy toad (O. chuanbeiensis) is known only from a small area of central China (northern Sichuan). The Puxiong lazy toad (O. puxiongensis) is known only from a single locality in west-central China (Sichuan). The dotted lazy toad (O. multipunctatus) and the Emei lazy toad (O. omeimontis) are both confined to a small area of central China (south-western Sichuan). Several lazy toads of the genus Scutiger are threatened by habitat destruction and degradation. The Chinting lazy toad (S. chintingensis) is known only from a small area of central China (Sichuan). The Ningshan lazy toad (S. ningshanensis) is known only from two specimens collected from a single locality in central China (southern Shaanxi). The Muli lazy toad (S. muliensis) is known only from a single locality in central China (south-western Sichuan). The Jiulong lazy toad (S. jiulongensis) is known only from a single locality in central China (southern Sichuan). The spotted lazy toad (S. maculatus) is known only from two localities in western China (northwestern Sichuan and eastern Tibet). The Pingwu lazy toad (S. pingwuensis) is known only from two localities in central China (north-eastern Sichuan and southern Gansu). The Liupan lazy toad (S. liupanensis) is confined to a few localities in central China (Gansu province and Ningxia Autonomous Region). The bumpy lazy toad (S. tuberculatus) is confined to a small area of central China (southern Sichuan). The Adung lazy toad (S. adungensis) is known for certain only from two specimens collected from the Adung Valley in northern Myanmar in 1931. The Chapa bug-eyed tree frog (Theloderma bicolor) occurs in south-western China (Yunnan) and north-western Vietnam (Quang Tri province), where it is threatened by loss of habitat. Hu’s gliding tree frog (Rhacophorus hui) is known only from two, relatively distant montane areas of central China (Sichuan and Hubei), but presumably occurs more widely. The Hoanglien gliding tree frog (R. hoanglienensis) is known from two localities in northern Vietnam (Lao Cai and Ha Giang provinces).

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Dubois’ gliding tree frog (Zhangixalus duboisi) is known only from two localities in northern Vietnam and southwestern China (Yunnan). The Jinxiu foam-nest tree frog (Gracixalus jinxiuensis) is known only from a few widely scattered montane localities in southern China (south-eastern Yunnan, north-eastern Guangxi, and southern Hunan) and northern Vietnam (Mount Fan Si Pan). It likely represents a cluster of species, some as yet undescribed. The Htingnan shrub frog (Philautus tytthus) is known only from its original collection in northern Myanmar in 1940. The Darjeeling shrub frog (P. dubius) is known only from its original collection during the late nineteenth century from an undefined locality in north-eastern India (West Bengal or Meghalaya). Several odorous frogs of the genus Odorrana are threatened by habitat destruction and overcollection for food. The Jingdong odorous frog (O. jingdongensis) occurs over a relatively wide area of south-western China (Yunnan and Guangxi) and northern Vietnam. The geminated odorous frog (O. geminata) is known from a few localities in southwestern China (south-eastern Yunnan) and north-eastern Vietnam (Ha Giang and Cao Bang provinces). The Trankien odorous frog (O. trankieni) is known only from a single locality in north-eastern Vietnam (Son La province). Ahl’s odorous frog (O. sinica) is known only from a single specimen collected in the early twentieth century from an undefined locality in China. The Tay Yen Tu odorous frog (O. yentuensis) is known only from the Tay Yen Tu Nature Reserve in north-eastern Vietnam (Bac Giang province). The Guangwu odorous frog (O. kuangwuensis) is known from a few localities in central China (Sichuan and Hubei). The Nanjiang odorous frog (Oreolalax nanjiangensis) is confined to a few isolated localities in central China (Sichuan, Gansu, and Shaanxi), where it is threatened by loss of habitat. Three species of horned frog (Megophrys) are endemic to small areas of central China, where they are threatened by loss of habitat. The Nankiang horned frog (M. nankiangensis) is known only from a few localities in Sichuan and Gansu. The convex-tailed horned frog (M. caudoprocta) is confined to parts of Hunan and Hubei. The Shuicheng horned frog (M. shuichengensis) is known only from a single locality in western Guizhou. The Kambaiti cascade frog (Amolops longimanus) is known only from its original collection in 1939 from northern Myanmar. The vitreous cascade frog (A. vitreus) is known only from northern Laos (Phongsaly province) and northern Vietnam (Dien Bien and Son La provinces). The Assamese cascade frog (A. assamensis) is known only from the Mayeng Hill Reserve Forest in north-eastern India (Assam). The Lolokou cascade frog (A. loloensis) is confined to a small area of central China (Sichuan and Yunnan). The minute cascade frog (A. minutus) is known only from Mu Ham Mountain in north-western Vietnam (Lai Chau province), but likely ranges as well into south-western China (Yunnan). The splendid

The Sino-Himalayan Region

cascade frog (A. splendissimus) is known only from a small area of north-western Vietnam (Lai Chau province). Boring’s large-eyed litter frog (Leptobrachium boringii) is known from a few localities in central China (Sichuan, Guizhou, and Hunan), where it is threatened by overcollection for human consumption and the international pet trade, as well as by habitat destruction. The Dawei large-eyed litter frog (L. promustache) is confined to a small area of southern China (Yunnan) and adjacent north-western Vietnam (Lào Cai province). Two species of spiny frog (Quasipaa) are threatened by habitat destruction and overcollection for human consumption. The Jiulong spiny frog (Q. jiulongensis) is known from a few localities in east-central China (Zhejiang and Fujian). Shin’s spiny frog (Q. shini) occurs in central China (Sichuan, Guizhou, Hunan and Guangxi). Two species of tiny frog (Nanorana) are threatened by overcollection for food and subsistence wood extraction outside of protected areas. Anderson’s tiny frog (N. yunnanensis) occurs over a wide area of central and south-western China (Sichuan, Yunnan, Guizhou, Hunan, and possibly Hubei), northern Myanmar, and northern Vietnam. The Tonkin tiny frog (N. unculuanus) occurs in south-western China (Yunnan), northern Vietnam, and possibly northern Laos. The Chuanan short-legged litter frog (Brachytarsophrys chuannanensis) is confined to a small area of central China (southern Sichuan). Liu’s wart frog (Limnonectes liui) is known only from a small area of south-western China (southern Yunnan), but may extend into adjacent areas of Myanmar and Laos. The Tsinpa salamander (Liua tsinpaensis) is known only from three localities in the mountains of central China (southern Shaanxi and north-eastern Sichuan). The Yichang salamander (Hynobius chinensis) is known only from a single locality in central China (Hubei province). Three salamanders of the genus Pseudohynobius are threatened by loss of habitat and overcollection for food and the international pet trade. The Schuicheng salamander (P. shuichengensis) is known only from a single locality in southwestern China (Guizhou). The Kuankuoshui salamander (P. kuankuoshuiensis) is known only from the PuchangKuankuoshui Nature Reserve in Guizhou. The yellow-spotted salamander (P. flavomaculatus) is confined to two localities in central China (Hubei and Hunan). Three species of mountain stream salamander (Batrachuperus) are threatened by overcollection for food and ‘traditional medicine’. Schmidt’s mountain stream salamander (B. tibetanus) occurs in central China (Sichuan, Shaanxi, Qinghai, Gansu, and Tibet). Pinchon’s mountain stream salamander (B. pinchonii) is currently found in western Sichuan and north-western Yunnan, having been extirpated from south-eastern Guizhou many decades ago. The Yenyuan mountain stream salamander (B. yenyuanensis) is known from the Daliang, Daxue, and Xiaoxianglin mountain ranges of south-western Sichuan.

The red-tailed knobby newt (Tylototriton kweichowensis) is confined to the mountains of central China (western Guizhou and north-eastern Yunnan).

The Tibetan Plateau The Tibetan Plateau (also known as the Qinghai-Tibet Plateau or Himalayan Plateau) is located in Central and East Asia. It covers much of western China (Tibet Autonomous Region, southern Xinjiang, northern Yunnan, western Sichuan, western Gansu, and Qinghai), as well as parts of northern India (Ladakh and Himachal Pradesh), Pakistan (Gilgit-Baltistan), northern Nepal, Bhutan, eastern Afghanistan, eastern Tajikistan, and southern Kyrgyzstan. The world’s highest and largest plateau, it stretches approximately 1000 km from north to south, some 2500 km from east to west and averages over 4500 m in elevation. Often referred to as the ‘Roof of the World’ or the ‘Third Pole’, it is essentially a vast, high-altitude arid steppe interspersed with mountain ranges and large, brackish lakes. It is from here that several major rivers (including the Yangtze, Mekong, and Indus) begin their long journeys to the sea. Owing to its remoteness much of the environment remains intact, with species and subspecies like the Tibetan wild ass (Equus kiang), Tibetan gazelle (Procapra picticaudata), and Tibetan grey wolf (Canis lupus filchneri) all still relatively common. The wild yak (Bos mutus) is a large, shaggy bovid and the ancestor of the domestic yak (B. grunniens). Today it is primarily found in the cold treeless uplands of northern Tibet and western Qinghai, with a few populations extending into the southernmost parts of Xinjiang, China, and Ladakh, India. Further small, isolated populations are also found in western Tibet and eastern Qinghai. Historically the species was also found in central China, Nepal, Sikkim and Bhutan, but has been extirpated there due to hunting. The white-lipped or Thorold’s deer (Cervus albirostris), one of the largest of all deer, formerly ranged across much of the eastern Tibetan Plateau but has been much reduced by intensive hunting, both for meat as well as ‘traditional medicine’. It now occurs only in fragmented populations in remote mountainous areas of eastern Tibet, Gansu, Qinghai, Sichuan, and Yunnan. The total population is estimated at around 7000. The Tibetan brown bear (Ursus arctos pruinosus) is a littleknown subspecies with long, bluish fur found on the eastern Tibetan Plateau. Rarely sighted in the wild, it is threatened mainly by hunting for use in Chinese ‘traditional medicine’ and loss of habitat. The Chinese mountain cat (Felis bieti) is a small, rare species found only on the north-eastern edge of the Tibetan Plateau. It is threatened by the organized poisoning of pikas (Ochotona sp.), which both diminishes prey and kills the cats unintentionally. Przewalski’s gazelle (Procapra przewalskii) was formerly widespread across the high plateaus of north-western China, from the area around Lake Qinhai through Gansu to Ningxia, possibly Shanxi and Inner Mongolia. It underwent a significant

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decline in the 1950s due to hunting, and today is confined to around a dozen sites within five isolated areas around Lake Qinhai. Kozlov’s pika (Ochotona koslowi) is known only from three localities in northern Tibet. Kozlov’s shrew (Sorex kozlovi) is known only from a single locality in eastern Tibet. The Linzhi mountain vole (Neodon linzhiensis) is known only from the Gongbu Nature Reserve in south-eastern Tibet. The Chinese monal (Lophophorus lhuysii) is a type of pheasant confined to the mountains of central China (western Sichuan, eastern Tibet, south-eastern Qinghai, southern Gansu, and possibly north-western Yunnan). In 2000 the total population was estimated to be around 12,000 and declining. The giant babax (Pterohinus waddelli) is a rare, thrush-like bird confined to southern Tibet and extreme north-eastern India (Sikkim). The Sichuan jay (Perisoreus internigrans) is a rare species confined to the mountains of central China (western Sichuan, eastern Tibet, south-eastern Qinghai, and southern Gansu). Sillem’s rosefinch (Carpodacus sillemi) was long known only from two specimens collected in 1929 from southern Xinjiang, China. In 2012 it was photographed at another locality some 1500 km away in western Qinghai province. The Medog forest lizard (Calotes medogensis) is only known from a single locality in south-eastern Tibet. Kaulback’s lance-headed pit viper (Protobothrops kaulbacki) is known only from Medog County (south-eastern Tibet) and from northern Myanmar. The Zamda toad (Bufotes zamdaensis) is known only from a single high-elevation wetland locality on the south-western Tibetan Plateau. The Medog gliding tree frog (Rhacophorus verrucopus) is known only from a single locality on the south-eastern Tibetan Plateau, but may occur more widely. The Medog foam-nest tree frog (Gracixalus medogensis) is only known from a single specimen collected in southeastern Tibet. The Medog frog (Liurana medogensis), Xizang frog (L. xizangensis), and the alpine frog (L. alpina) are all known only from small areas of south-eastern Tibet. The reticulated frog (Ingerana reticulata) is known only from a small area of south-eastern Tibet. Two species of cascade frog (Amolops) endemic to the Tibetan Plateau are threatened by loss of habitat. The Medog cascade frog (A. medogensis) and the Aniqiao cascade frog (A. aniqiaoensis) are each known only from a single locality in Medog County, on the south-eastern edge of the Tibetan Plateau. The Medog tiny frog (Nanorana medogensis) is known only from a small area of south-eastern Tibet. The Medog horned frog (Megophrys medogensis) is known only from a small area of south-eastern Tibet, where it is threatened by loss of habitat due to road construction and urbanization.

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The Himalayan Mountains Located in Central Asia, the Himalayas include many of the Earth’s highest peaks and separate the plains of Indian subcontinent from the Tibetan Plateau. As here defined, they are spread across Bhutan, south-western China, northern India, Nepal, and northern Pakistan and are divided into two main subranges. The Himalayan brown bear (Ursus arctos isabellinus) is a relatively small subspecies that lives in high-altitude forests and alpine meadows in northern Nepal, northern and northwestern India, and northern Pakistan. It is threatened by loss of habitat. The Himalayan black bear (U. thibetanus laniger) is a long-furred subspecies that, during the summer, can be found at high altitudes of the Himalayas and surrounding areas. In winter it tends to descend to warmer lowerelevation forests. The Himalayan Pallas’ cat (Otocolobus manul nigripectus) is a rare subspecies that occurs in the Himalayas from Kashmir to Bhutan. The Himalayan tahr (Hemitragus jemlahicus) is a large wild goat native to the Himalayas of southern Tibet, northern India, and Nepal, where it is threatened by hunting and habitat destruction. The species has been widely introduced to Argentina, New Zealand, South Africa, and the United States. The Himalayan musk deer (Moschus leucogaster) occurs in parts of Afghanistan, Pakistan, Tibet, Nepal, Bhutan, and northern India. It is ruthlessly hunted for use in ‘traditional medicine’ and has suffered a considerable decline. The satyr tragopan (Tragopan satyra) is a type of pheasant found patchily in the Himalayas of northern India, southern Tibet, Nepal, and Bhutan, where it is threatened by hunting and habitat destruction. The wood snipe (Gallinago nemoricola) occurs as a vagrant throughout a wide area of Indochina and South Asia, but breeds only in the high alpine meadows of the Himalayas of northern India, Nepal, Bhutan, and southern China. It declined drastically during the early twentieth century due to hunting, which continues to be a threat in some areas. The Western Himalayas The Western Himalayas stretch from Badakhshan in northeastern Afghanistan/southern Tajikistan, through India (Jammu and Kashmir, Himachal Pradesh and Uttarakhand) to central Nepal. The Himalayas capture moisture from the monsoons that sweep in from the Bay of Bengal, and most of this rainfall is expended in the eastern Himalayas. Therefore, the western Himalayas are drier, a trend reflected in the timberline that declines from 4000 m in the east to about 3500 m in the west. The Chamba sacred langur (Semnopithecus ajax) is confined to a small area of north-western India, primarily in the Chamba Valley of Himachal Pradesh but extending somewhat into Jammu and Kashmir.

The Sino-Himalayan Region

The Kashmir markhor (Capra falconeri cashmiriensis) is a type of wild goat confined to a small area of north-western India (Jammu and Kashmir) and north-eastern Pakistan, where it is threatened by military activity and competition from introduced livestock. Two subspecies of mouflon (Ovis gmelini), a type of wild sheep previously discussed in this volume, are endemic to the Western Himalayas. The Punjab mouflon (O. g. punjabensis) is confined to the foothill forests of eastern Pakistan (Punjab). The Ladakh mouflon (O. g. vignei) is found in Ladakh and northern Pakistan, including Kashmir. The Kashmir stag (Cervus hanglu hanglu) is confined to dense riverine forests in the Kashmir Valley and northern Chambra district of Himachal Pradesh. Up until 1947 the total population of between 3000 and 5000 was regarded as the personal property of the Maharaja of Kashmir, which gave it adequate protection. In the years after Indian independence, however, it was heavily poached and much of its habitat destroyed by domestic livestock. By 1970 the population had been reduced to just 140–170 living in the Dachigam Sanctuary (now a national park) in Kashmir, with a few small groups in Himachal Pradesh. An international conservation project helped to increase this number to over 340 by 1980, but since then the animals have been under renewed threat from local shepherds and their dogs. As of 2018 only 186 were known to survive. The Kashmir musk deer (Moschus cupreus) is found sporadically in extreme north-western India (Jammu and Kashmir), northern Pakistan, and perhaps north-eastern Afghanistan (Nuristan province). It is seriously threatened by poaching for its prized scent glands. The Himalayan goral (Naemorhedus goral) is a type of goat-antelope divided into two subspecies. Hodgson’s Himalayan goral (N. g. goral) is found in southern Tibet, Bhutan, eastern Nepal, and north-eastern India (Sikkim and Arunachal Pradesh). Bedford’s Himalayan goral (N.

g. bedfordi) is confined to northern India (Jammu and Kashmir, Himachal Pradesh and Uttaranchal) and western Nepal. Both are threatened by loss of habitat and hunting for their meat. The central Kashmir vole (Alticola montosa) is known only from two localities in north-western India and northern Pakistan. The pale grey shrew (Crocidura pergrisea) is known only from a single locality in Pakistan-administered Kashmir. Peters’ tube-nosed bat (Harpiola grisea) is known only from a small area of northern India (Uttarakhand). The cheer pheasant (Catreus wallichi) occurs patchily in the western Himalayas from northern Pakistan through Kashmir and northern India to central Nepal. Widely shot for sport during the early twentieth century, it continues to be hunted and trapped for food and its eggs collected for local consumption. The western tragopan (Tragopan melanocephalus) has a disjunct distribution in the western Himalayas from the IndusKohistan district of Pakistan, east through Kashmir and Himachal Pradesh to Uttarakhand in north-western India. Although historically considered to be rare, a mid-1980s population estimate of 1600–4800 was revised in the mid-1990s to around 5000 birds following the discovery of several significant populations in northern Pakistan. It is threatened by habitat destruction and hunting. The Himalayan quail (Ophrysia superciliosa) is known only from the western Himalayas in north-western India (Uttaranchal), where about a dozen were collected near Mussooree and Naini Tal prior to 1877. It has not been recorded with certainty since then, despite numerous searches. However, quail are naturally difficult to observe and the species probably remains extant, although in small numbers. The Kashmir flycatcher (Ficedula subrubra) breeds in the Western Himalayas of north-western India and north-eastern Pakistan, from where it migrates south to winter primarily in

Figure 2.9 Himalayan quail painted by John Gould in 1883. (Credit: John Gould.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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the mountains of Sri Lanka as well as in the Western Ghats. At one time common within its restricted range, it has declined in many areas due to habitat destruction. The Agaupani mountain lizard (Japalura dasi) is known only from a single locality (Agaupani village in western Nepal). The Nepal lazy toad (Scutiger nepalensis) is confined to the alpine grasslands of north-western Nepal, where it is threatened mainly by diversion of water from breeding streams. The Chakrata cascade frog (Amolops chakrataensis) and Jaunsar’s cascade frog (A. jaunsari) are each known only from a single specimen collected in 1985 from a single locality near Chakrata in northern India (Uttarakhand). It seems likely that the two species in fact represent the same taxon, which was rediscovered in the same area in 2011. The Rara Lake tiny frog (Nanorana rarica) is confined to a small area of north-western Nepal. Rostand’s tiny frog (N. rostandi) occurs in western and central Nepal. Dubois’ tiny frog (N. minica) is found in northern India (Uttar Pradesh and Himacahal Pradesh) and in western Nepal. All are threatened by habitat destruction. The Eastern Himalayas The Eastern Himalayas extend from central Nepal and across north-eastern India (Sikkim, West Bengal, and Arunachal Pradesh), Bhutan, and south-eastern Tibet to parts of northern Myanmar and south-western China (Yunnan). The Arunachal macaque (Macaca munzala) is known only from a small area of north-eastern India (Arunachal Pradesh) and possibly adjacent areas of Bhutan. The white-cheeked macaque (M. leucogenys) was only recently described in 2015 from south-western China (south-eastern Tibet) and north-eastern India (Arunachal Pradesh). Both are threatened by hunting and loss of habitat. The golden langur (Trachypithecus geei) is confined to forest remnants in north-eastern India (western Assam) and southern Bhutan. The capped langur (T. pileatus) remains fairly common as a species over a large area of South Asia, in both lowland as well as highland areas. Nevertheless, it has suffered considerable population declines and is considered vulnerable. The Bhutan capped langur (T. p. tenebricus) is from north-eastern India and Bhutan. The Bhutan takin (Budorcas taxicolor whitei) is a type of goat-antelope found in high-altitude bamboo forests in northeastern India (Arunachal Pradesh and Sikkim), Bhutan, and south-western China (southern Tibet). It is threatened by hunting and habitat destruction. The Mechuka giant flying squirrel (Petaurista mechukaensis) is known only from an area of forest in north-eastern India (Arunachal Pradesh). Its range may extend into southwestern China. The giant mole shrew (Anourosorex schmidi) is known only from a few specimens collected on the south-eastern slopes of the Himalayas in Bhutan and north-eastern India (Arunachal Pradesh and Sikkim).

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Mandelli’s mouse-eared bat (Myotis sicarius) is known only from a few localities in north-eastern India (Sikkim and West Bengal) and Nepal, where it is threatened by deforestation. Csorba’s mouse-eared bat (M. csorbai) is known only from a few localities in Nepal. Both are threatened by loss of habitat. The sombre serotine bat (Eptesicus tatei) is known only from its original collection in 1951 from north-eastern India (West Bengal). Molesworth’s grey-bellied tragopan (Tragopan blythii molesworthi) is known only from eastern Bhutan, where it has not been recorded since the early 1970s. The chestnut-breasted partridge (Arborophila mandellii) occurs in the eastern Himalayas north of the Brahmaputra River in north-eastern India (West Bengal, Sikkim, and Arunachal Pradesh), Bhutan, and southern China (southeastern Tibet). It is threatened by hunting and loss of habitat. The rufous-throated wren-babbler (Spelaeornis caudatus) is a rarely observed species from Nepal, Bhutan, and northeastern India (Sikkim, West Bengal, and Arunachal Pradesh), where it is threatened by loss of habitat. The walnut kukri snake (Oligodon juglandifer) is known only from a few localities in north-western India (West Bengal), perhaps extending into western Bhutan and Sikkim. The few-scaled worm snake (Gerrhopilus oligolepis) is known only from a small area of north-eastern India (Sikkim and West Bengal). The Darjeeling blind snake (Indotyphlops meszoelyi) is known only from single specimen collected in north-eastern India (West Bengal). The Darjeeling caecilian (Ichthyophis sikkimensis) is known only from a small area of north-eastern India (Sikkim and West Bengal) and extreme eastern Nepal. The Daphla (or Dafla) Hills are a tract of hilly country in north-eastern India (Assam and south-western Arunachal Pradesh). The Daphla Hills capped langur (Trachypithecus pileatus brahma) is confined to the Daphla Hills. The Bugun liocichla (Liocichla bugunorum) is a rare type of passerine bird known only from a few localities within the Daphla Hills.

The Purvanchal Range The Purvanchal Range is located in north-eastern India south of the Brahmaputra Valley (Assam, Arunachal Pradesh, Manipur, Tripura, Nagaland, Meghalaya, and Mizoram) and north-western Myanmar (Chin State). The Namdapha gliding squirrel (Biswamoyopterus biswasi) is known only from a single specimen collected in 1981 from Namdapha National Park in north-eastern India (Arunachal Pradesh). The snowy-throated babbler (Stachyris oglei) is a rare species known from the Patkai and Mishmi Hills of northeastern India (Arunachal Pradesh) and adjacent northern Myanmar.

The Sino-Himalayan Region

The hornbill foam-nest tree frog (Chirixalus shyamrupus) is known only from Namdapha National Park in north-eastern India (Arunachal Pradesh). The Arunachal Pradesh shrub frog (Philautus namdaphaensis) is known only from two localities in north-eastern India (Arunachal Pradesh).

The Taihang Mountains The Taihang Mountains (Taihang Shanmai in Chinese) are a mountain range running down the eastern edge of the Loess Plateau in Shanxi, Henan, and Hebei provinces. The Taihang tiny frog (Nanorana taihangnica) is known only from a single locality in the Taihang Mountains (Henan).

The Naga Hills The Naga Hills are located in north-eastern India (Nagaland and Manipur) and north-western Myanmar (Chin State). The Samagutin worm snake (Indotyphlops tenuicollis) is known only from three specimens collected in the Naga Hills. The Nagaland bug-eyed tree frog (Theloderma nagalandense) is known only from a single locality within the Naga Hills of north-eastern India (Nagaland). The Zunheboto horned frog (Megophrys zunhebotoensis) is known only from a single locality within the Naga Hills of north-eastern India (Nagaland). The Mokochung tiny frog (Nanorana mokokchungensis) is known only from a single locality in north-eastern India (Nagaland).

The Qinling Mountains

The Mishmi Hills The Mishmi Hills are located in north-eastern India (Arunachal Pradesh). The rusty-throated wren-babbler (Spelaeornis badeigularis) was long known only from a single specimen collected in 1947 from the Mishmi Hills. It was rediscovered in the same small area in 2004, where it appears to be fairly common. The Patkai Hills The Patkai Hills are located in north-western Myanmar (Chin State) and north-eastern India (Mizoram and Tripura). The Lushai Hills (also known as the Mizo Hills) are located in north-eastern India (Mizoram and Tripura). The Serchhip horned frog (Megophrys serchhipii) is known only from a single specimen collected in the Lushai Hills.

The Loess Plateau The Loess Plateau, also known as the Huangtu Plateau (Huangtu Gaoyuan in Chinese), is located in central China. It covers almost all of the provinces of Shaanxi and Shanxi, and extends into parts of Gansu, Ningxia, and Inner Mongolia. The Luliang Mountains The Luliang Mountains (Luliang Shanmai in Chinese) are located in western Shanxi. The Shanxi sika deer (Cervus nippon grassianus) likely occurred at one time throughout the entire Loess Plateau, although in more recent years it has been confined to two populations in the upland forests of the Luliang Mountains. One of the most mercilessly hunted deer in China, there have been no reports of it for decades and it is now believed to be extinct in the wild. Pure-bred individuals remain in farms, but there is not enough suitable habitat left for reintroduction efforts.

The Qinling or Qin Mountains (Qinling Shanmai in Chinese) are a major east–west range in central China (southern Shaanxi). The mountains provide a natural boundary between northern and southern China, and support a large number of endemic species. The Qinling giant panda (Ailuropoda melanoleuca qinlingensis), notable for its light brown and white fur pattern, is confined to the Qinling Mountains. The Qinling golden snub-nosed monkey (Rhinopithecus roxellana qinlingensis) is confined to the Qinling Mountains. The golden takin (Budorcas taxicolor bedfordi) is a type of goat-antelope confined to high-elevation areas of the Qinling Mountains.

The Wuling Mountains The Wuling Mountains (Wuling Shanmai in Chinese) are located in central China (Guizhou and Hunan). Mount Fanjing At an altitude of 2570 m, Mount Fanjing (Fanjing Shan in Chinese) is the highest peak in the Wuling Mountains. The Fanjingshan National Nature Reserve was established in 1978. It was designated a UNESCO Biosphere Reserve in 1986, and a World Heritage Site in 2018. The grey snub-nosed monkey (Rhinopithecus brelichi) is confined to a block of forest centering on Fanjing Mountain, south of the Yangtze in the Wuling Mountains. There have been unconfirmed anecdotal reports of a population in Jinfoshan Nature Reserve in the Dalou Mountains, but this has yet to be resolved. The total population in 2005 was estimated at around 750 individuals.

The Qionglai Mountains The Qionglai Mountains (Qionglai Shanmai in Chinese) are located in central China (Sichuan). Marie’s vole (Volemys musseri) is known from a few specimens collected in the alpine meadows of the Qionglai Mountains.

The Min Mountains The Min Mountains (Min Shanmai in Chinese) are located in central China (southern Gansu and north-central Sichuan). The snowy-cheeked laughingthrush (Garrulax sukatschewi) is confined to a few forest fragments within the Min Mountains. The rusty-throated parrotbill (Sinosuthora przewalskii) is confined to a few forest fragments within the Min Mountains.

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The Daba Mountains The Daba Mountains (Daba Shanmai in Chinese) are located in east-central China (Sichuan, Chongqing, Shaanxi, and Hubei). The Hubei golden-snub-nosed monkey (Rhinopithecus roxellana hubeiensis) is confined to the Shennongjia forest region of the Daba Mountains.

The Tianmu Mountains The Tianmu Mountains (Tianmu Shanmai in Chinese) are located in east-central China (Zhejiang). Mount Tianmu Mount Tianmu (Tianmu Shan in Chinese) is made up of two peaks (West Tianmu and East Tianmu). It is nominally protected as a UNESCO Biosphere Reserve. The Tianmu loach (Leptobotia tchangi) is a type of freshwater fish known only from Tianmu Mountain.

The Dabie Mountains The Dabie Mountains (Dabie Shanmai in Chinese) are located in east-central China (Hubei, Henan, and Anhui). The western part has a relatively low elevation, though there are a few peaks rising to 900 m. The eastern part is higher, averaging 1000 m. The range is still heavily forested. The Anhui musk deer (Moschus anhuiensis) is confined to the Dabie Mountains of western Anhui province. The Shangcheng stout salamander (Pachyhynobius shangchengensis) is confined to the Dabie Mountains.

The Luoxiao Mountains The Luoxiao Mountains (Luoxiao Shanmai in Chinese) are located in east-central China (Jiangxi, Hubei, and Hunan). The Jinggang Range The Jinggang Range (Jingang Shanmai in Chinese) is located in the remote border region of Jiangxi and Hunan provinces. Zong’s odd-scaled snake (Achalinus jinggangensis) is known only from a few specimens collected in the Jinggang Range.

The Huangshan Mountains The Huangshan Mountains are located in eastern China (southern Anhui). The Huangshan horned frog (Megophrys huangshanensis) is known only from Mount Huangshan.

The Gaoligong Mountains The Gaoligong Mountains (Gaoligong Shanmai in Chinese) are located in south-central China (western Yunnan). They straddle the border between south-western China and northern Myanmar. The Xiaoheishan slug-eating snake (Pareas nigriceps) is known only from a type series collected on Xiaohei Hill, in the Gaoligong Mountains National Nature Reserve.

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The Pianma cascade frog (Amolops bellulus) is known only from a single specimen collected in 1940 from the western slopes of Gaoligongshan.

Mount Emei Mount Emei (Emei Shan in Chinese) is located at the western rim of the Sichuan basin in south-central China (Sichuan). It was declared a World Heritage Site in 1996. Anderson’s shrew mole (Uropsilus andersoni) appears to be largely confined to Mount Emei. The chevron-spotted brown frog (Rana chevronta) is known only from montane forest on Mount Emei, where it was last reported in 1983. The Longdong stream salamander (Batrachuperus longdongensis) is known only from the Longdong River on Mount Emei.

The Hengduan Mountains The Hengduan Mountains (Hengduan Shanmai in Chinese) are a group of mountain ranges in south-western China (Sichuan, north-western Yunnan, southern Qinghai, and eastern Tibet) and north-eastern Myanmar (Kachin State). Running north to south, they connect the south-eastern portions of the Tibetan Plateau with the Yunnan-Guizhou Plateau. The Liangshan vole (Proedromys liangshanensis) is known only from two nature reserves in south-western Sichuan. The Gongshan wolf snake (Lycodon gongshan) is known only from two localities in Yunnan. Ping’s lazy toad (Oreolalax pingii) is known only from two localities in the Daliang Mountains of west-central China (Sichuan). The Puxiong salamander (Pseudohynobius puxiongensis) is known only from a few specimens collected from a single locality in Sichuan province. The Yun Mountains (Yun Ling in Chinese) are located in south-western China (north-western Yunnan and southeastern Tibet). The black snub-nosed monkey (Rhinopithecus bieti) occurs only in the Yun Mountains. In 2006 the total known population was estimated at less than 2000, divided among 15 known subpopulations. The Yangbi Yi toad (Bufo aspinius) is known only from a small area of the Yun Mountains, where it is threatened by habitat destruction.

The Yunnan-Guizhou Plateau The Yunnan-Guizhou or Yungui Plateau (Yungui Gaoyuan in Chinese) is a highland area located in south-western China, primarily in the provinces of Yunnan and Guizhou. In the south-west it is a true plateau, while in the north-east it is a mountainous region of rolling hills, deep gorges, and karst topography. The binturong or bearcat (Arctictis binturong) is a large viverrid still found across much of South and South East Asia, but is everywhere uncommon and threatened by hunting and

The Sino-Himalayan Region

habitat destruction. A number of subspecies are recognized. The Yunnan binturong (A. b. menglaensis) is confined to Yunnan. The inquisitive shrew mole (Uropsilus investigator) is confined to high-altitude areas of south-western China (Yunnan). The Yunnan box turtle (Cuora yunnanensis) was long known only from a few specimens collected in south-western China (Yunnan), and thought possibly to be extinct. Rediscovered in recent years, it remains rare (or perhaps extinct) in the wild but has been established in captivity. The Puer gliding tree frog (Rhacophorus puerensis) is known only from a small area of montane forest in southwestern China (Yunnan). The great piebald horned frog (Megophrys giganticus) is confined to montane forest in south-western China (southwestern Yunnan). The Jingdong cascade frog (Amolops tuberodepressus) is known only from two mountains (Wuliang and Ailao) in south-western China (Yunnan). The piebald tiny frog (Nanorana maculosa) is known only from a few localities in south-western China (central Yunnan). The Chenggong firebelly newt (Cynops chenggongensis) is known only from a single locality in south-western China (Yunnan).

The Yunnan Plateau The Yunnan Plateau is located in south-western Yunnan, and represents the only actual flattened portion of the YunnanGuizhou Plateau. The white-handed gibbon (Hylobates lar) is divided into a small number of subspecies that were historically found throughout much of South East Asia. The Yunnan white-handed gibbon (H. l. yunnanensis) is known only from the Nangunhe Nature Reserve in south-western Yunnan, where a survey in 2007 failed to find any sign of the subspecies. It is most likely extinct. The Yunnan bush rat (Hadromys yunnanensis) is known only from a small area of the Yunnan Plateau. Boehme’s wolf snake (Lycodon synaptor) is known only from a small area of the Yunnan Plateau. The Dalou Mountains The Dalou Mountains (Dalou Shanmai in Chinese) are a limestone mountain range running some 300 km across the northern edge of the Yunnan-Guizhou Plateau in Guizhou and Sichuan provinces. The Jinfo salamander (Pseudohynobius jinfo) is confined to Mount Jinfo and Mount Baima.

The Dayao Mountains The Lijiang Mountains The Lijiang Mountains (Lijiang Shanmai in Chinese) are located in southern China (north-western Yunnan). The pygmy brown-toothed shrew (Chodsigoa parva) is known only from a few specimens collected in the Lijiang Mountains in the 1920s. The Lijiang pit viper (Gloydius monticola) is confined to the mountains of southern China (north-western Yunnan).

The Dayao Mountains (Dayao Shanmai in Chinese) are located in south-eastern China (eastern Guangxi). The Dayao gliding tree frog (Zhangixalus yaoshanensis) and the minimal gliding tree frog (Z. minimus) are both confined to the Dayao Mountains. The Dayao bug-eyed tree frog (Theloderma kwangsiense) is known only from the Dayao Mountains.

The Miaoling Mountains The Ailao Mountains The Ailao Mountains (Ailao Shanmai in Chinese) are located in south-central China (Yunnan). The Ailao toad (Bufo ailaoanus) is known only from a single specimen collected in the Ailaoshan National Nature Reserve in 1984. The Jingdong lazy toad (Oreolalax jingdongensis) is confined to the Ailao Mountains. Wulian Feng The Wulian Feng (literally ‘Five lotus peaks’) is located in south-western China (central Yunnan). More of an escarpment towering above the Jinsha (upper Yangtze) River than an actual mountain range, it forms the north-western edge of the Yunnan-Guizhou Plateau. The Wuliangshan slender litter frog (Leptolalax alpinus) is known only from the Wuliangshan National Nature Reserve, where it is threatened by subsistence wood extraction. Wuliang Mountain (Wuliang Shan in Chinese) is located in south-western Yunnan. The Wuliang reed snake (Calamaria yunnanensis) is known only from a few specimens collected from Wuliang Mountain.

The Miaoling Mountains (Miaoling Shanmai in Chinese) are located in south-central China (south-eastern Guizhou). The Leishan large-eyed litter frog (Leptobrachium leishanense) is known only from two localities in the Miaoling Mountains.

The Hoang Lien Son Range The Hoang Lien Son Range is located in north-western Vietnam (Lao Cai province), near the border with China. Part of it lies within Hoang Lien National Park. The Tonkin weasel (Mustela tonkinensis) is known only from a single specimen collected in the Hoang Lien Son Mountains. The white-throated wren babbler (Rimator pasquieri) is confined to the montane forests of the Hoang Lien Mountains. Mount Fansipan Mount Fansipan (Phan Xi Pang in Vietnamese) is the highest mountain on the Indochinese Peninsula, rising as it does some 3143 m. Sterling’s lazy toad (Oreolalax sterlingae) is known only from a single stream on Mount Fansipan.

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The green-backed gliding tree frog (Zhangixalus dorsoviridis) is known only from Mount Fansipan, where it has been recorded only once since its discovery in the 1930s. The Fansipan large-eyed litter frog (Leptobrachium echinatum) is known only from Mount Fansipan. Botsford’s slender litter frog (Leptobrachella botsfordi) is known only from a single high-elevation stream on Mount Fansipan. Bourret’s slender litter frog (L. bourreti) and the rainy slender litter frog (L. pluvialis) are similarly known only from Mount Fansipan, but may range more widely.

Miscellaneous Mountains and Highlands Mount Mang (Mang Shan in Chinese) is located in central China (southern Hunan and northern Guangdong). The Mangshan pit viper (Protobothrops mangshanensis) is known only from Mount Mang. Mount Taibei (Taibei Shan in Chinese) is located in central China (Shanxi). The Taibai mountain stream salamander (Batrachuperus taibaiensis) is known only from Taibei Shan. Mount Wawu (Wawu Shan in Chinese) is located in southcentral China (Sichuan). The Wawu horned frog (Megophrys wawuensis) is confined to Mount Wawu. Mount Qanning (Qanning Shan in Chinese) is located in south-central China (Sichuan). The Qanning lazy toad (Scutiger brevipes) is known only from Mount Qanning. Mount Longtou is located in south-central China (Guizhou). The Longtou odorous frog (Odorrana anlungensis) is known only from Mount Longtou. Mount Longdao (Longdao Shan in Chinese) is located in southern China (northern Guangdong). The Longdao odorous frog (Odorrana leporipes) is known only from its original collection in the early twentieth century from Mount Longdao. Mount Dawei (Dawei Shan in Chinese) is located in southcentral China (south-eastern Yunnan), near the border with Vietnam. The Dawei horned toad (Megophrys daweimontis) is known only from Mount Dawei. Mount Huanglian (Huanglian Shan in Chinese) is located in south-central China (Yunnan). The Huanglian cascade frog (Amolops caelumnoctis) is known only from Mount Huanglian, but may range more widely. Mount Mao’er (Mao’er Shan in Chinese) is located in south-eastern China (Guangxi). The Xingan salamander (Hynobius maoershanensis) is known only from Mount Mao’er. Mount Longwang (Longwang Shan in Chinese) is located in coastal eastern China (Zhejiang). The Longwang salamander (Hynobius amjiensis) is confined to five small pools atop Mount Longwang. Mount Yangming (Yangming Shan in Chinese) is located in south-eastern China (Hunan). The Yangming frog (Rana hanluica) is known only from Mount Yangming. Mount Tay Con Linh II is located in northern Vietnam (Ha Giang province). The Tay Con Linh cascade frog (Amolops iriodes) is known only from Mount Tay Con Linh II.

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Lowland Subtropical Moist Forests Much of southern China and northern Vietnam were once covered by subtropical moist forests, but most disappeared long ago and today only remnants remain. Those in karstic areas in particular provide an important last refuge for many species, particularly primates. The Cao-Vit crested gibbon (Nomascus nasutus) was historically found in north-eastern Vietnam and southern China (Guangxi and Guangdong), roughly from Ha Long Bay to the Red River delta. The species was most likely extirpated in China during the 1950s, and was feared to be possibly extinct by the early twenty-first century. However, since then a few populations have been discovered in northern Vietnam (Cao Bang province), and it is thought that more may occur in Hoa Binh and Bac Kan provinces as well. It remains one of the world’s most endangered primates. Two leaf-eating monkeys of the genus Trachypithecus are largely or entirely confined to karst forests, where they are highly threatened by loss of habitat and hunting for use in ‘traditional medicine’. Delacour’s langur (T. delacouri) is confined to a small area of north-central Vietnam (Ninh Bình, Ha Nam, Hòa Binh, Thanh Hoa, and Ha Tay provinces), where less than 250 still survived as of 2010. There are very few in captivity, and it is feared that the only remaining viable wild population may now be in the Van Long Nature Reserve in Ninh Bình province. François’ langur (T. francoisi) occurs widely but patchily in south-central China (Chongqing, Guangxi, Guizhou, and Sichuan) and northern Vietnam. The Cuc Phuong ferret-badger (Melogale cucphuongensis) is known only from two specimens collected from a heavily degraded forest area in northern Vietnam. It is most likely threatened by hunting. Daovantien’s limestone rat (Tonkinomys daovantieni) is known only from the Huu Lien Nature Reserve in northeastern Vietnam (Lang Son province). The blue-crowned laughingthrush (Pterohinus courtoisi) is a seriously threatened species with an extremely small and fragmented range in south-eastern China (Jiangxi). The bird is known in the international pet trade, and a small captivebreeding programme has been established. The Nonggang babbler (Stachyris nonggangensis) is confined to a small area of south-western China (south-western Guangxi) and adjacent northern Vietnam. The crocodile lizard (Shinisaurus crocodilurus) is a semiaquatic species restricted to karst mountain drainages. Two subspecies are generally recognized. The Chinese crocodile lizard (S. c. crocodilurus) is found in south-eastern China (Hunan, Guangxi, and Guizhou). The Vietnamese crocodile lizard (S. c. vietnamensis) is confined to a small area of northeastern Vietnam (Quang Ninh province). Both are threatened by habitat destruction and harvesting for the international pet trade. The Maolan pit viper (Protobothrops maolanensis) is known only from the Maolan National Nature Reserve in

The Sino-Himalayan Region

south-eastern China (Guizhou). The Trung Khanh pit viper (P. trungkhanhensis) is known only from two specimens collected from the Trung Khanh Nature Reserve in northern Vietnam (Cao Bang province). Moellendorf’s trinket snake (Elaphe moellendorffi) is known from a few lowland karst areas in southern China (Guangdong and Guangxi) and north-western Vietnam (Hoa Binh province). It is heavily traded in animal markets. The Zhaoping tree frog (Hyla zhaopingensis) is known only from a small area in south-central China (eastern Guangxi). The Hong Kong cascade frog (Amolops hongkongensis) is confined to a few localities in southern China (Guangdong) and Hong Kong. The Vietnamese knobby newt (Tylototriton vietnamensis) is known only from two areas of forest in north-eastern Vietnam (Bac Giang and Quang Ninh provinces). The species is present in captivity and in the international pet trade.

Lowland Temperate Deciduous Forests As with lowland subtropical forests there are very few areas of lowland temperate forest remaining within the SinoHimalayan Region, the vast majority having long ago been converted for agriculture. Patches remain in central and north-eastern China. The North China sika deer (Cervus nippon mandarinus) historically inhabited lowland forests of northern and northeastern China. Loss of habitat and intense hunting pressure had reduced it to remote areas of north-eastern China and the Qing Imperial Hunting Grounds for centuries. There have been no sightings for many decades, however, and it is now thought to be extinct in the wild. It remains fairly common in zoos and on Asian antler farms, although lack of suitable habitat makes reintroduction efforts impossible. The Chinese water deer (Hydropotes inermis inermis) was historically found across a relatively wide area of coastal eastern China, but has since been extirpated from much of this range and is now confined to the eastern Yangtze River drainage, where it continues to decline due to hunting and habitat destruction. It was introduced to England in the 1870s, where it has thrived and indeed become something of a pest. A second introduced population in western France appears to have become extirpated. Graham’s lizard (Diploderma grahami) is known only from a single specimen collected in the 1920s from an unspecified locality in central China (Sichuan). The Shanxi gecko (Gekko auriverrucosus) is known only from a single specimen collected from north-central China (Shanxi). The species is possibly extinct due to loss of habitat.

Isolated Caves, Springs, and Pools Southern China and northern Vietnam have extensive areas of karst and cave systems, providing habitat to a number of amphibians and fish.

The red-spotted lazy toad (Oreolalax rhodostigmatus) is known from a few limestone caves in central China (Hubei, Sichuan, Guizhou, and Hunan). The Wuchuan odorous frog (Odorrana wuchuanensis) is known only from a single, isolated limestone cave near Baicun in south-western China (Guizhou). The total population is very small. The Na Hang dwarf litter frog (Leptobrachella nahangensis) is known only from a cave entrance in the Na Hang Nature Reserve of northern Vietnam (Tuyen Quang province). The Maolan salamander (Paramesotriton maolanensis) is known only from a single deep pool in the Maolan National Nature Reserve in south-eastern China (Guizhou), where it probably also lives in underground rivers. The Jianshui cavefish (Typhlobarbus nudiventris) is confined to a single cave in southern China (Yunnan). Several golden-line barbels (Sinocyclocheilus) are endemic to isolated caves in southern and south-central China. The duckbilled golden-line barbel (S. anatirostris) is known only from two caves within the Bailang subterranean river in southcentral China (Guangxi). The Daxiao Dong golden-line barbel (S. cyphotergous) and the angular golden-line barbel (S. angularis) are confined to south-eastern China (Guizhou). The hyaline golden-line barbel (S. hyalinus) and the eyeless golden-line barbel (S. anophthalmus) are confined to southern China (Yunnan). The small-eyed golden-line barbel (S. microphthalmus) occurs in an undefined locality in southern China. Zheng’s stone loach (Oreonectes anophthalmus) is confined to a few caves in south-central China (Guangxi and Guizhou). The Gejiu stone loach (Triplophysa gejiuensis) is a blind, subterranean species confined to a single underground river in south-western China (Yunnan). The Changping stone loach (Paranemachilus genilepis) is known only from a single subterranean river in south-central China (southern Guangxi). The Nan Tong stone loach (Yunnanilus parvus) is known only from an imprecise type locality (a cave outlet) in southern China (Yunnan). The Guangxi cave loach (Protocobitis typhlops) is confined to a cave in south-central China (Guangxi).

Lakes, Rivers, and Marshes The Sino-Himalayan Region is not as rich as the Eurasian Region in terms of its lakes, rivers, and marshes, although those that do exist tend to be much more important in terms of biodiversity. They include the upper reaches of many important southern and South East Asian rivers, along with the Yangtze, Pearl, Yellow, and Red river systems. There are also a great many small, montane lakes in southern China along with miscellaneous other wetlands. The white-eared night heron (Oroanassa magnificus) is found widely but patchily in southern China, Hainan, and

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northern Vietnam, where it is threatened by habitat destruction, hunting, and trapping. The Japanese night-heron (Gorsachius goisagi) breeds in Japan and on Jeju Island off the Korean Peninsula, occurring at other times in the Philippines, Indonesia, parts of China, and the Russian Far East. Swinhoe’s giant softshell turtle (Rafetus swinhoei) historically occurred in the lakes and rivers of eastern and southern China. Today, only a single living captive individual is known, an old male in the Suzhou Zoo in China. Another at the Beijing Zoo died in 2005, and a third at the Shanghai Zoo the following year; both had been caught at Gejiu in the 1970s. A few wild individuals continue to be reported. One was caught in 1998 in the Red River and subsequently released. Another semi-wild specimen of questionable taxonomy lived in a lake in Hanoi, Vietnam for many years until its death in 2016. More recently one was captured and observed several times in Dong Mo Lake west of Hanoi, and another from a nearby lake. The species is clearly very close to extinction. The Chinese soft-shelled turtle (Pelodiscus sinensis) is commercially farmed in vast numbers and has been introduced to areas outside its native habitat. However, wild populations continue to be exploited for food, resulting in serious declines. The wattle-necked softshell turtle (Palea steindachneri) is still fairly widespread in south-eastern China (Guangdong, Guangxi, Guizhou, Yunnan, and the island of Hainan), northern Laos, and northern Vietnam, but is everywhere threatened by overcollection for food. It has been introduced to Hawaii and Mauritius. The big-headed turtle (Platysternon megacephalum) is divided into three subspecies, all of which are threatened by overcollection for use as food. The nominate form (P. m. megacephalum) is found in southern China. Several species and subspecies of pond turtle (Mauremys) are threatened by loss of habitat, hybridization with introduced turtle species, and overcollection for use as food or pets. The Chinese stripe-necked pond turtle (M. sinensis) is found in south-eastern China (Guangdong, Fujian, and the island of Hainan), Taiwan, Laos, and northern and central Vietnam. The red-necked pond turtle (M. nigricans) is confined to southern China (Guangxi and Guangdong). Reeves’ pond turtle (M. reevesii) is native to most of temperate and subtropical China and the Korean Peninsula as well as Taiwan, Hong Kong, and parts of Japan, although the island populations may have been introduced. Populations on Timor and Palau certainly originated from human introductions. At least two subspecies of yellow pond turtle (M. mutica) occur in South East Asia. The nominate form (M. m. mutica) ranges from central Vietnam north through central and southern China, with additional insular populations on Taiwan and Hainan. Beal’s eyed turtle (Sacalia bealei) has a relatively wide distribution in mainland China (Anhui, Fujian, Guangdong, Guangxi, Guizhou, and Jiangxi), along with the island of Hong

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Kong, but is everywhere heavily exploited for use in ‘traditional medicine’. The short-legged horned frog (Megophrys brachykolos) is known for certain only from Hong Kong, but may possibly also occur in other parts of south-eastern China (Fujian) and in northern Vietnam (Lang Son and Ha Bac provinces). The Hejiang odorous frog (Odorrana hejiangensis) is known only from a small area of north-eastern Vietnam (Bac Giang province). The Junlian odorous frog (O. junlianensis) is confined to a small area of west-central China (Sichuan and Guizhou). The concave-eared odorous frog (O. tormota) is confined to a small area of eastern China (Anhui, Zhejiang, Jiangsu, and Jiangxi). All are threatened by loss of habitat and overcollection for use as food. The Irrawaddy golden-backed frog (Hylarana margariana) is known only from two specimens collected in 1875 on the Myanmar/southern China border. Zheng’s frog (Rana zhengi) is known only from wetlands near the village of Zhangcun in north-western China (Sichuan). The Fujian frog (Glandirana minima) is confined to a few areas of remaining habitat in south-eastern China (eastern Fujian). Several species of spiny frog (Quasipaa) are threatened by loss of habitat and overcollection for use as food. The little spiny frog (Q. exilispinosa) is found patchily in south-eastern China (Fujian, Hunan, Guangdong, Guangxi, and the island of Hong Kong). Boulenger’s spiny frog (Q. boulengeri) is found in central and southern China and northern Vietnam. Ye’s spiny frog (Q. yei) is confined to the Dabie Mountains of eastern China (Hubei, Henan, and Anhui). David’s spiny frog (Q. spinosa) is found in central, southern, and south-western China (including the island of Hong Kong) and in northern Vietnam. Kaulback’s cascade frog (Amolops kaulbacki) is known only from its original collection in 1940 from northern Myanmar. The Van Ban cascade frog (A. cucae) is known only from a small area of north-western Vietnam (Lao Cai province). Two species of fully aquatic giant salamander (Andrias) are seriously threatened by overcollection for food and ‘traditional medicine’. Père David’s giant salamander (A. davidianus) is still widespread in the hillstreams of central, south-western, and southern China, primarily within the Yangtze River drainage, its range is now very fragmented. The South China giant salamander (A. sligoi) is the largest amphibian in the world, reaching as it does over a metre in length. Historically found in the Pearl River drainage south of the Nanling Mountains, it is extremely threatened and may no longer exist in the wild state. The Chiala mountain stream salamander (Batrachuperus karlschmidti) is a largely aquatic species confined to a small area of central China (Sichuan). The Guabangshan salamander (Hynobius guabangshanensis) is known only from a single locality in central China (Hunan province).

The Sino-Himalayan Region

The Shancheng stout salamander (Pachyhynobius yunanicus) is known only from a small area of central-eastern China (Henan). The Chinhai spiny newt (Echinotriton chinhaiensis) is confined to a small area of central coastal China (Zhejiang). Of the three known subpopulations, one has already been extirpated. The Wanggao warty newt (Paramesotriton fuzhongensis) and the Paiyangshan warty newt (P. guangxiensis) are both confined to small areas of south-eastern China (Guangxi), where they are threatened by loss of habitat and overcollection for the international pet trade. Three species of knobby newt (Tylototriton) are threatened by loss of habitat, pollution, and overcollection for use as food and in ‘traditional medicine’. The Anhui knobby newt (T. anhuiensis) is confined to a small area of central-eastern China (Anhui). The Dabien knobby newt (T. dabienicus) is confined to the Mount Dabien area of central-eastern China (Henen). The Wenxian knobby newt (T. wenxianensis) is found patchily in central China (southern Gansu, northern Sichuan, eastern Guizhou, and Chongqing). The Dayang newt (Cynops orphicus) is a rare species confined to a few small areas of south-eastern China (eastern Guangdong and central Fujian). The Yunnan carp (Cyprinus chilia) was historically known from a number of lakes on the Yunnan Plateau of southern China, but has been extirpated from most of them due to overfishing, pollution, and invasive species. The Assamese kingfish (Cyprinion semiplotum) is confined to hillstreams in north-eastern India, Nepal, and Bhutan, where it is threatened by overfishing, habitat destruction, and pollution. Langson’s shuttle-like carp (Luciocyprinus langsoni) occurs in both lakes and rivers in northern Vietnam (Lang Son province) and southern China (Yunnan, Gunagxi, and Guizhou). It has declined dramatically due to loss of habitat and by-catch fishery pressure. Chu’s naked carp (Gymnocypris chui) is known only from four endorheic mountain lakes in southern and western Tibet. Günther’s naked carp (G. dobula) is known only from Lake Peigu and Lake Chuocuolong in southern Tibet. Both are threatened by overfishing. Three species of mandarin fish (Siniperca) are threatened by overfishing and pollution. The golden mandarin fish (S. scherzeri) is found widely but patchily in China, the Korean Peninsula, and Vietnam. The big-eyed mandarin fish (S. kneri) is confined to the Yangtze and Pearl river drainages of China. The slender mandarin fish (S. roulei) is found patchily in eastern and south-eastern China. The Yunnan snowtrout (Schizothorax yunnanensis) is known from the upper Yangtze and Mekong river drainages and from Lake Caohai in south-western China (Yunnan and Guizhou). The species historically spawned in Lake Erhai as well, but has not done so since the 1970s. Richardson’s snowtrout (S. richardsonii) is widely distributed in the Himalayan foothills of northern India, Nepal, Bhutan,

Pakistan, and Afghanistan, but has declined everywhere due to overfishing, dams, and the introduction of exotic fish species. The Nam Mau barb (Spinibarbus nammauensis) is confined to Lake Ba Bê and the Nang River drainage in northern Vietnam. The Bac Can chub (Parazacco babeensis) and Vinh’s chub (P. vinhi) are both known only from a few creeks and rivers in northern Vietnam (Bac Can province). The Chinese large-lipped minnow (Parasinilabeo assimilis) is known from the Pearl and Yangtze river drainages, where it is threatened by loss of habitat, overfishing, and pollution. The White Cloud Mountain minnow (Tanichthys albonubes) is a popular aquarium fish with a relatively wide but highly disjunct distribution in southern China, including the island of Hainan. It is everywhere rare in the wild due to habitat degradation and pollution. Peters’ minnow (Pseudohemiculter dispar) is relatively widespread in south-eastern China, Vietnam, and Laos, but is threatened throughout much of its range by dam construction and pollution. The Tsinan gudgeon (Gnathopogon tsinanensis) is confined to a small area of north-eastern China (Shangdong). Nguyen’s goby (Rhinogobius imfasciocaudatus) is known only from a few creeks and rivers in the mountains of northern Vietnam (Ha Giang and Lai Chau provinces). The Ky-Phu loach (Liniparhomaloptera monoloba) is known only from Ky-Phu creek in northern Vietnam. The Nan Pang loach (Paralepidocephalus yui) is known from Yangzong and Yilong lakes and from the outflow of Jiuxiang cave in southern China (Yunnan). It is threatened by pollution. Inglis’ stone loach (Nemacheilus inglisi) is known only from a few rivers in north-eastern India (Sikkim). The Phong Tho stone loach (Schistura phongthoensis) is known only from a few creeks and rivers in north-western Vietnam (Lai Chau province). Hodgart’s swamp eel (Monopterus hodgarti) is known only from its original description during the early twentieth century from north-eastern India (Arunachal Pradesh).

Lake Dianchi Lake Dianchi (Dianchi Hu in Chinese; also known as Kunming Lake) is a large, high-elevation lake located in south-central China (Yunnan). Historically home to a number of endemic species, it has been devastated by overfishing, pollution, and introduced species. The Yunnan lake newt (Cynops wolterstorffi) was confined to the shores of Lake Dianchi. It has not been reported since 1979 despite extensive surveys and is now considered extinct, a victim of habitat destruction, pollution, and introduced species. The Dianchi carp (Cyprinus micristius) was historically found in Lake Dianchi and its tributaries, where it was a

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common commercial species. Not seen for many years, it is most likely extinct but may survive in the Songhuaba Reservoir. The Dianchi nase (Xenocypris yunnanensis) began to decline in the 1970s, and was last collected in 1985. It is now most likely extinct. The Dianchi snowtrout (Schizothorax grahami) was historically found throughout Lake Dianchi as well as its tributaries and connecting springs, but is now confined to the Muyang and Lengshui rivers and to the Black Dragon and Green Dragon springs within the Songhuaba Reservoir drainage. The Dianchi golden-line barbel (Sinocyclocheilus grahami) has been extirpated from the lake itself due to heavy pollution, but survives in a single lake tributary and in some 20 small temple ponds. The elongated bitterling (Acheilognathus elongatus) was historically confined to shallow inshore areas of Lake Dianchi, where it may have disappeared by the 1980s due to pollution and loss of habitat. Two species of silver minnow (Anabarilius) historically occurred in Lake Dianchi. The big silver minnow (A. polylepis) was at one time a major commercial species, but likely disappeared from the lake by the 1970s. It may survive in the Songhuaba Reservoir. The Dianchi silver minnow (A. alburnops) has had a similar history, and is today only caught occasionally. The Dianchi catfish (Silurus mento) was historically common, but began to be rare by the mid-twentieth century. It has not been collected since the 1970s and is likely extinct, although it may survive in some lake-connected reservoirs. The Dianchi bullhead (Pseudobagrus medianalis) is a type of catfish that was historically common within the Lake Dianchi basin, but is now known for certain only from White Dragon Spring and Longwantan Spring (which may be connected by small, underground karst rivers) and to the Zhangjiu River. The species may also occur in a few other tributary rivers and reservoirs around the lake. It is threatened by pollution as well as from conversion of springs into fish ponds. Two other species of bullhead (genus Liobagrus) were historically found in Lake Dianchi. King’s bullhead (L. kingi) appears to have disappeared from the lake basin in the 1960s, but may persist in two tributaries of the Jinsha River in the upper Yangtze River drainage. The black-bellied bullhead (L. nigricauda) still survives in the Lake Dianchi basin. The Dianchi stone loach (Sphaerophysa dianchiensis) is confined to Lake Dianchi. It has not been recorded in many decades and is possibly extinct, although it may persist in isolated springs and tributaries. The White Dragon stone loach (Yunnanilus discoloris) is entirely confined to White Dragon Spring, where the total population is thought to be less than 500. It has yet to be protected and has been heavily modified and degraded. Two other species, the black-spotted stone loach (Y. nigromaculatus), and the side-striped stone loach (Y. pleurotaenia), are

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also found in the Lake Dianchi and perhaps elsewhere, where they are thought to be threatened.

Lake Fuxian Lake Fuxian (Fuxian Hu in Chinese) is located in southcentral China (Yunnan). It is the third largest lake in China, as well as the deepest. Its numerous endemic fish species have been seriously impacted by overfishing and introduced species. The Fuxian carp (Cyprinus fuxianensis) was historically confined to the lake. Not seen after 1995, it is most likely extinct. The Fuxian mahseer (Folifer yunnanensis) was last seen in the 1990s. The Fuxian snowtrout (Schizothorax lepidothorax) was last recorded in the 1990s. The Fuxian barb (Poropuntius chonglingchungi) was last recorded in the 1980s, and is most likely extinct. The Fuxian gudgeon (Discogobio longibarbatus) was last recorded in the 1990s, and is most likely extinct. The Fuxian golden-line barbel (Sinocyclocheilus tingi) is confined to Lake Fuxian, where it has undergone a drastic decline. The species has been successfully bred in captivity, however, and reintroduced to the lake. Graham’s silver minnow (Anabarilius grahami) is confined to Lake Fuxian, where it is nearing extinction. The Fuxian stone loach (Triplophysa fuxianensis) is confined to Lake Fuxian. Two other species of stone loach (Yunnanilus), Chu’s stone loach (Y. chui) and the West Dragon Spring stone loach (Y. obtusirostrisa), were last seen in the 1990s and are possibly extinct.

Lake Qilu Lake Qilu (Qilu Hu in Chinese) is located on the Yunnan Plateau in south-central China (Yunnan). Introduction of non-native species and overfishing have had a devastating effect on the endemic species. Tchang’s carp (Cyprinus yunnanensis) and the Qilu carp (C. ilishaestomus) have not been reported since the late 1970s, and are likely extinct. The Qilu silver minnow (Anabarilius qiluensis) has not been seen since the early 1980s, and is most likely extinct.

Lake Yangzong Lake Yangzong (Yangzong Hai in Chinese) is located on the Yunnan Plateau in south-central China (Yunnan). It has been heavily impacted by industrial pollution and overfishing. The Yangzong golden-line barbel (Sinocyclocheilus yangzongensis) is known only from Lake Yangzong. The Yangzong silver minnow (Anabarilius yangzonensis) is endemic to Lake Yangzong, where it is very rare or possibly already extinct. The Yangzong stone loach (Paracobitis oligolepis) is known only from Lake Yangzong.

The Sino-Himalayan Region

Lake Erhai Lake Erhai (Er Hai Hu in Chinese) is a large lake located in the mountains of south-central China (Yunnan). Its native fishes have been decimated by overfishing, pollution, and introduced species. Four species of carp (Cyprinus) were historically endemic to Lake Erhai. The Dali carp (C. daliensis) has not been recorded since the 1960s, and is most likely extinct. The large-eyed carp (C. megalophthalmus) has not been recorded since the early 1980s, and is possibly extinct. The bearded carp (C. barbatus) and the long-finned carp (C. longipectoralis) both went unrecorded from the early 1980s and were also thought to be extinct, although a few reports since the early 2000s indicate that they may still survive in very small numbers. The little barb (Poropuntius exiguus) and the Dali barb (P. daliensis) have not been recorded in decades, and are possibly extinct. The Erhai sprat (Zacco taliensis) was historically confined to Lake Erhai, where it is now possibly extinct. The Erhai stone loach (Paracobitis erhaiensis) was historically confined to Lake Erhai, where it is now possibly extinct.

The Upper Mekong River The upper Mekong (Lancang Jiang in Chinese) is located in southern China (Tibet and Yunnan) and northern Laos. General threats include overfishing, pollution and dam construction. The red mahseer (Tor sinensis) is known for certain only from southern China (Yunnan), but may extend into Laos, Cambodia, and Thailand. The gold-striped danio (Devario chrysotaeniatus) is confined to tributaries and smaller streams within the upper Mekong drainage. The upper Mekong bat-catfish (Oreoglanis setiger) is known only from a few records in the upper Mekong in south-western China (Yunnan) and the Ma Oun River in north-western Laos (Louang Namtha province). Kloetzli’s stone loach (Schistura kloetzliae) and the largeheaded stone loach (S. macrocephalus) are both known only from the Mengla River in southern China (Yunnan) and the Youan River in north-western Laos (Luang Namtha province). The Mengla River is located in southern China (Yunnan). The Mengla mahseer (Tor polylepis) is known only from the Mengla River. The Nanla River is located in southern China (Yunnan) and north-western Laos (Luang Namtha province). Its major tributary is the Youan River. The Nanla rasbora (Rasbora septentrionalis) is confined to the Nanla and Youan rivers. The Youan danio (Devario apopyris) is known only from a single stream in the Youan River watershed. The Nanla goby (Rhinogobius maculicervix) is confined to the Nanla and Youan rivers.

The Nanla stone loach (Sectoria heterognathos) is confined to the Nanla and Youan rivers.

The Upper Irrawaddy River The Irrawaddy or Ayeyarwady River flows from north to south from south-western China (Yunnan) through Myanmar to the Indian Ocean. The upper Irrawaddy, as here defined, includes the upper tributaries of south-western China and northern Myanmar (i.e. those before the confluence with the Chindwin River). Tsao’s snowtrout (Schizothorax meridionalis) is known only from two tributaries of the upper Irrawaddy in southwestern China (Yunnan). The Qiaojie garra (Garra qiaojiensis) is known from only two localities in south-western China (Yunnan). The Dracula fish (Danionella dracula) was only described in 2007 from a consignment of aquarium fishes shipped to the United Kingdom, said to have originated from a stream near Sha Du Zup in northern Myanmar. The marvellous fish (D. mirifica) is similarly known only from a single locality in northern Myanmar (Kachin State). The Putao torrent minnow (Psilorhynchus brachyrhynchus) is known only from a few localities within the upper Irrawaddy River drainage of northern Myanmar. Malaise’s catfish (Glyptosternon malaisei) is known only from its original collection from the upper Irrawaddy River in the 1940s. The Irrawaddy naked catfish (Batasio procerus) is known only from a few specimens collected from the upper Irrawaddy River drainage in northern Myanmar. The Tengchong stone loach (Schistura polytaenia) is known only from a single tributary of the upper Irrawaddy in south-western China (Yunnan). The Taping River (also known as the Daying River in China and the Ta Hkaw Kha in Myanmar) is located in south-western China (Yunnan) and northern Myanmar. The Taping snowtrout (Schizothorax malacanthus), Taping garra (Garra bispinosa), and Taping stone loach (Schistura yingjiangensis) are all known only from the Taping River drainage. The Nampoung River is located in south-western China (Yunnan) and northern Myanmar. The Nampoung barb (Poropuntius margarianus) is known only from the Nampoung River.

The Upper Salween River The Salween River is one of the largest river drainages in South East Asia, flowing some 2815 km through China, Myanmar, and Thailand to the Andaman Sea. The upper Salween River (known as the Nu Jiang in China) originates in the Tanggula Mountains on the Tibetan Plateau, and runs through southern China (Yunnan) before finally dropping into a deep gorge into north-eastern Myanmar. A series of massive dams have long been planned.

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The Nujiang barb (Poropuntius opisthoptera) is confined to the upper Salween River in southern China (Yunnan). The bihorned barbel (Tariqilabeo bicornis) is known only from a single specimen collected from the upper Salween River in the 1970s. The Nujiang snowtrout (Schizothorax nukiangensis) is confined to the upper Salween River in Yunnan. The Tibetan osman (Schizopygopsis thermalis) is confined to the upper Salween River drainage in Tibet. Fea’s catfish (Pareuchiloglanis feae) is known for certain only from the upper Salween River of southern China and north-eastern Myanmar. The Nujiang river loach (Hemimyzon nujiangensis) is confined to the upper Salween River in Yunnan. The Nanpenghe River The Nanpenghe River is located in southern China (Yunnan). The Nanpenghe garra (Garra nujiangensis) is known only from the Nanpenghe River.

The Upper Ganges River The upper Ganges (Ganga in Hindustani) rises in the western Himalayas of northern India and Nepal and flows south. The Bonar baril (Barilius bonarensis) is a type of cyprinid fish known only from its original description in 1912 from north-western India (Uttarakhand). The dark mahseer (Naziritor chelynoides) is confined to the headwaters of the Ganges River. Edds’ catfish (Pseudecheneis eddsi) is known only from the Ganges River drainage in central Nepal. The Nepalese naked catfish (Batasio macronotus) is known only from a few specimens collected from the Ganges River drainage in eastern Nepal. The Narayani River The Narayani River (also known as the Gandaki River) is located in Nepal. The Rapi catfish (Erethistoides cavatura) is confined to the Rapi River, a tributary of the Narayani River in southcentral Nepal. The Mewa Khola River The Mewa Khola River is located in eastern Nepal. The thicktail catfish (Pseudecheneis crassicauda) is known only from the Mewa Khola River. The Song River The Song River is located in north-western India (Uttarakhand). The dimorphic baril (Barilius dimorphicus) is known only from the Song River. The Giri River The Giri River is located in north-western India (Himachal Pradesh).

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The Giri catfish (Pseudecheneis suppaetula) is known only from the upper reaches of the Giri River.

The Upper Brahmaputra River The upper Brahmaputra River, as here defined, begins in the mountains of Tibet as the Tsangpo River and flows down through Bhutan and north-eastern India (Arunachal Pradesh, Sikkim, West Bengal, and Assam). The blunt-nosed snowtrout (Schizothorax molesworthi) is known only from the upper Brahmaputra River drainage, where it is heavily exploited for food. The Brahmaputra torrent catfish (Amblyceps arunchalensis) is known only from the Subansiri and Dikrong rivers in north-eastern India (Arunachal Pradesh). Tikader’s stone loach (Aborichthys tikaderi) is known only from a few streams within the Brahmaputra River drainage of north-eastern India (Arunachal Pradesh). The Tsangpo River The Tsangpo River (Yarlung Tsangpo in Tibetan) is the upper stream of the Brahmaputra River, and the longest river in Tibet. The Sattar snowtrout (Schizothorax curvilabiatus) is confined to the lower reaches of the Tsangpo River. The Teesta River The Teesta or Tista River is a 315-km river that rises in the eastern Himalayas. It flows through north-eastern India (Sikkim and West Bengal) before joining the Brahmaputra River in Bangladesh. The Teesta brook carp (Neolissochilus spinulosus) is known only from the Teesta River in Sikkim, where it is considered rare. The Teesta catfish (Pseudolaguvia foveolata) and the giant fennel catfish (P. ferula) are known only from the Teesta River drainage. The Teesta eel-loach (Pangio apoda) is known only from the Teesta River. The Raidak River The Raidak River is located in Bhutan, north-eastern India (West Bengal), and Bangladesh. The Raidak catfish (Pseudolaguvia ferruginea) is known only from a single locality within the Raidak River of northeastern India (West Bengal). The Siren River The Siren River is located within the upper Brahmaputra River drainage of north-eastern India (Arunachal Pradesh). The Siren River catfish (Pseudecheneis sirenica) is confined to the Siren River. The Kameng River The Kameng River is a tributary of the Brahmaputra River located in the eastern Himalayas of north-eastern India (Arunachal Pradesh and Assam).

The Sino-Himalayan Region

The Dirang torrent minnow (Psilorhynchus arunachalensis) is known only from the Dirang River, a tributary of the Kameng River. The Dikrong River The Dikrong River is located in north-eastern India (Assam and Arunachal Pradesh). The Senkhi catfish (Erethistoides senkhiensis) is known only from Senkhi stream, a tributary of the Dikrong River. The Dibru River The Dibru River is a small tributary of the Brahmaputra located in north-eastern India (northern Assam). The Dibru perch (Badis dibruensis) is known only from the Dibru River.

The Yangtze River The Yangtze River (Chang Jiang in Chinese) is the longest river in Asia and the third longest in the world. It rises in the mountains of the Tibetan Plateau and flows some 6300 km to its mouth at the East China Sea, near Shanghai. It has been severely affected by pollution and dam construction. The Three Gorges Dam, the world’s largest hydroelectric project, will greatly alter the flow of the river and seriously affect many species. The Yangtze River dolphin or baiji (Lipotes vexillifer) was once common along much of the central and lower Yangtze but is now considered to be functionally extinct, a victim of industrialization. The last known living individual died in 2002, although a videotape taken in 2007 appears to show one of the animals. Any remaining survivors, however, would be insufficient to save the species. The narrow-ridged finless porpoise (Neophocaena asiaorientalis) is divided into two subspecies. The Yangtze narrow-

ridged finless porpoise (N. a. asiaorientalis) is a freshwater species confined to the middle and lower reaches of the Yangtze River along with Poyang and Dongting lakes and their tributaries, although historically it was found as far as 1600 km upstream. It is threatened by habitat destruction and degradation, boat traffic, and pollution. The Chinese alligator (Alligator sinensis) was historically widespread in the lower reaches of the Yangtze River, but has declined dramatically in recent decades due to human persecution. Today it is known only from a small area of south-eastern Anhui province, mainly in small ponds. A relatively large captive population and reintroduction efforts may not be enough to ensure its survival. The Chinese sturgeon (Acipenser sinensis) is an anadromous fish that spends most of its life out at sea, but relies on freshwater rivers for spawning. Historically found throughout the waters surrounding China, Japan, and the Korean Peninsula, it has been extirpated almost everywhere due to habitat destruction and overfishing. It is also greatly prized for use in ‘traditional medicine’. At present it spawns only in the Yangtze River, where the construction of the Gezhouba Dam in 1981 blocked its migration route. The Chinese government has made some efforts to project it, including marine reserves specifically aimed at this species and restocking through the release of juveniles in the lower parts of the Yangtze. Dabry’s sturgeon (A. dabryanus) was, under the natural, unaltered conditions that existed until the middle of the twentieth century, an inhabitant of the middle and upper reaches of the Yangtze River and its major tributaries, along with a number of large lakes within the drainage. An important species in commercial fisheries, during the late twentieth century it declined drastically due to overfishing and habitat degradation. Since the mid-1990s it has been extirpated from the lower river, and is now restricted to the upper main stream

Figure 2.10 Qiqi, the last captive Yangtze River dolphin. (Credit: Institute of Hydrobiology, Chinese Academy of Science.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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of the Yangtze in Sichuan province as well as the Ming, Tuo, and Jialing rivers. The Chinese paddlefish (Psephurus gladius), one of the world’s largest freshwater fishes, is (or was) endemic to the Yangtze River, including the brackish water at its mouth. Its population had already greatly decreased due to overfishing and habitat degradation when, in 1981, the construction of the Gezhouba Dam in the middle reaches of the Yangtze blocked its spawning route to the upper reaches of the river system. Juveniles were last recorded in the mid-1990s and only two adult specimens have been recorded since 2002. The species is likely already extinct. The Hubei yellowfin (Xenocypris hupeinensis) is known only from a single locality within the Yangtze River drainage in east-central China (Hubei). Banarescu’s loach (Parabotia banarescui) is confined to the middle reaches of the Yangtze in east-central China (Hubei). Chen’s loach (P. lijiangensis) is known only from the Li and Xiang rivers of south-eastern China (Guangxi and Hunan). The imperial flower loach (Leptobotia elongata) is confined to the upper and middle Yangtze River drainage in Hubei and Hunan provinces, where it has been extirpated from many areas due to disruption of its migration paths, habitat degradation, and pollution. The Jinsha River The Jinsha River (Jinsha Jiang) is the Chinese name for the upper stretches of the Yangtze in central and south-western China (Qinghai, Sichuan, and Yunnan). The Sichuan taimen (Hucho bleekeri) is a type of salmon endemic to the headwaters of the Yangtze River drainage, where it is threatened by habitat destruction and overfishing. Tchang’s sharpbelly (Hemiculter tchangi) is known only from the upper reaches of the Yangtze River in central Chinga (Sichuan). The Jinsha stone loach (Triplophysa grahami) is confined to the Jinsha River and tributaries in Yunnan. Lake Quinshui is located within the Jinsha River drainage in south-western China (Sichuan). The Quinshui silver minnow (Anabarilius xundianensis) is confined to Lake Quinshui. The Yuan River The Yuan River is located in southern and south-eastern China (Guizhou and Hunan). The Yuan stone loach (Triplophysa xiangxensis) is a cavedwelling species confined to the Yuan River in Hunan. Lake Dongting is a large, shallow lake located in northeastern Hunan. The Dongting mountain catfish (Glyptothorax sinensis) is known only from Lake Dongting. The Li River The Li River (Li Jiang in Chinese) is located in south-eastern China (Guangxi province).

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The Li zebra loach (Sinibotia zebra) is known only from the Li River.

The Pearl River The Pearl River (Zhu Jiang in Chinese, and formerly known as the Canton River) is an extensive river system in south-eastern China and northern Vietnam. The name is essentially a catchall for the combined watersheds of the Xi, Bei, and Dong rivers, which share a common delta. Reeves’ noodlefish (Salanx reevesii) is known only from the Pearl River drainage in Guangdong province. The Pearl River bangana (Bangana decorus) is known from the Pearl River drainage in Yunnan, Guangdong, and Guangxi provinces, where it has been impacted by damming and pollution. Richardson’s armoured catfish (Cranoglanis bouderius) was historically widespread and common within the Pearl River drainage, but has suffered significant declines due to overfishing and other factors. The Pearl River loach (Sinibotia robusta) is confined to the Pearl River drainage. The Lia Jiang loach (Vanmanenia homalocephala) is known only from a single locality within the Pearl River. Lin’s spined loach (Cobitis arenae) is known only from the Xi and Dong rivers, tributaries of the Pearl River in southern China. The Xi River The Xi River (Xi Jiang in Chinese) is the largest and westernmost of the Pearl River tributaries. It is formed by the confluence of the Gui and Xun rivers in Guangxi, and flows east through Guangdong before entering the Pearl River delta just east of the Lingyang Gorge in Zhaoqing. The Xiangzhou icefish (Neosalanx argentea) is known only from its original description in the 1930s from the city of Xiangzhou (Guangdong). Hoffmann’s amblatikas (Toxabramis hoffmanni) is known only from a single locality in Guangxi province, China. The Xi gudgeon (Gobiobotia meridionalis) is confined to the Xi River drainage. Jordan’s ratmouth barbel (Ptychidio jordani) and the bigeye ratmouth barbel (P. macrops) are both found patchily within the Xi River drainage, where they are seriously threatened by overfishing. The Xi loach (Erromyzon sinensis) is confined to the Xi River drainage. The golden zebra loach (Sinibotia pulchra) is known from the Liu and Gui rivers, tributaries of the Xi River. The Xi stone loach (Oreonectes furcocaudalis) is a type of cavefish known from a single locality in the Xi River (an outlet of subterranean waters near Rongshui in Guangxi) The Nanpan River (Nanpan Jiang in Chinese) is located in southern China (Yunnan, Guizhou, and Guangxi). Part of its course is blocked by the Tianshengqiao Dam, from which Lake Wanfeng is formed.

The Sino-Himalayan Region

The Nanpan garra (Garra yiliangensis) is known only from a single specimen collected in the 1960s from a hillstream in southern China (Yunnan). The Nanpan silver minnow (Anabarilius maculatus) is confined to the Nanpan River and a few associated reservoirs. The Zuo River (Zuo Jiang in Chinese) is located in southern China (Guangxi). The Zuo carp (Cyprinus longzhouensis) is known for certain only from the upper Zuo River drainage. The Hongshuihe River is located is located in southern China (Guangxi). Wu’s fish (Discocheilus wui) is known only from the Hongshuihe River drainage. The Na-Ri River is located in northern Vietnam (Lang Son and Cao Bang provinces). The Na-Ri shovel-jaw carp (Onychostoma uniforme) is known only from the Na-Ri River. The Dong River The Dong River is located in south-eastern China (Guangdong). It is the most easterly tributary of the Pearl River. The Dong River mountain catfish (Glyptothorax pallozonus) is known only from the Dong River drainage. The Thuong River The Thuong River is located in north-eastern Vietnam (Lang Son province). The Thuong bitterling (Acheilognathus elongatoides) is known only from the Thuong River.

The Yellow River The Yellow River (Huang He in Chinese) is the second longest river in Asia. It originates in the Bayan Har Mountains of Qinqhai province, western China, and meanders east for about 1900 km before emptying into the Bohai Sea. The Tengger water frog (Pelophylax tenggerensis) is endemic to a small area along the banks of the Yellow River in north-central China (Ningxia Hui Autonomous Region), on the edge of the Tengerr Desert. The Yellow River gudgeon (Squalidus intermedius) is known only from its type locality near the town of Jinan, in the Yellow River drainage.

The Red River System The Red River (Hong He in Chinese/Song Hong in Vietnamese) is located in south-western China (Yunnan) and northern Vietnam. The confluence of its main tributaries, the Black and Lo rivers, forms a very broad main channel in north-eastern coastal Vietnam that flows past the city of Hanoi before spreading out again through the Red River delta and thence into the Gulf of Tonkin. The delta was once highly productive, but is now almost devoid of fish due to extensive flood control and the closure of floodplain fish breeding and nursery areas.

The Red River carp (Cyprinus quidatensis) is confined to the Red River drainage of northern Vietnam. The Red River carp-minnow (Pseudolaubuca hotaya) is confined to the Red River drainage of northern Vietnam. The Tonkin bangana (Vinalabeo tonkinensis) is confined to the Red River drainage. The Hong bitterling (Rhodeus vietnamensis) is known only from the Red River drainage on the Chinese/ Vietnamese border. The Red River sharpbelly (Hemiculter songhongensis) is confined to the Red River drainage of northern Vietnam. The Black River The Black River (Song Da in Vietnamese, and known as the Lixian River in China) is located in southern China (Yunnan province) and north-western Vietnam (Lai Chau, Son La, Lao Cai, Phu Tho, and Hoa Binh provinces). It is the principal tributary of the Red River. Nguyen’s carp (Cyprinus hyperdorsalis) and Da’s carp (C. dai) have not been recorded since the construction of the Hoa Binh dams in the 1990s, and are possibly extinct. The Suoi Rut barb (Poropuntius brevispinus) is known only from the Suoi Rut stream within the Black River drainage. The Hoa Binh barb (Spinibarbus vittatus) is known only from the Black River in north-western Vietnam (Hoa Binh province). The Lai Chau catfish (Hemibagrus songdaensis) is known only from the upper Black River in north-western Vietnam (Lai Chau province). The Black River stone loach (Schistura callichromus) is known only from a single tributary in Yunnan, China, but may occur in other parts of the Black River drainage. Mai’s hillstream loach (Beaufortia daon) is known only from a few creeks and small rivers within the Black River drainage of north-western Vietnam (Lai Chau province). The Bua River (Nam Bua in Vietnamese) is located in north-western Vietnam (Son La and Phu Tho provinces). The Bua suckerbelly loach (Pseudogastromyzon buas) is known only from the Bua River drainage. The Na River (Nam Na in Vietnamese) is located in northwestern Vietnam (Lai Chau province). The Na freshwater sleeper (Neodontobutis macropectoralis) is a type of goby known only from the Na River. The Phong Tho catfish (Pareuchiloglanis phongthoensis) is known only from the Na River drainage. The Na hillstream loach (Sinogastromyzon namnaensis) is known only from the Na River. The Mu River (Nam Mu in Vietnamese) is located in north-western Vietnam (Lai Chau province). The Nam Mu catfish (Pareuchiloglanis dorsoarcus) is known only from the Mu River drainage. The Dê River (Nam Dê in Vietnamese) is located in northwestern Vietnam (Lai Chau province). The Dê catfish (Pareuchiloglanis namdeensis) is known only from the Dê River drainage.

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The So River (Nam So in Vietnamese) is located in northwestern Vietnam (Lai Chau province). The So hillstream loach (Sinogastromyzon hypercorpus) is known only from the So River drainage. The Ngoi Bo River (Nam Ngoi Bo in Vietnamese) is located in north-western Vietnam (Lao Cai province). The Ngoi Bo catfish (Tachysurus longispinalis) is known only from the Ngoi Bo River drainage. The Tengtiao River (Tengtiao Jiang in Chinese) is located in southern China (Yunnan) and north-western Vietnam (Lai Chau province). The Tengtiao stone loach (Schistura macrotaenia) is known only from the Tengtiao River. The Lo River The Lo River (Song Lo in Vietnamese) is located in southern China (Yunnan) and northern Vietnam (Ha Giang, Lai Chai, Tuyen Quang, and Phu Tho provinces). The Lo steed (Hemibarbus songloensis) is a type of barb known only from the Lo River drainage. The Lo goby (Rhinogobius longipinnis) is known from the Lo River drainage and from the Nui Coc reservoir and its feeding streams. The Ha Giang catfish (Pareuchiloglanis rhabdurus) is known only from a small area of the Lo River. The Ha Giang hillstream loach (Sinogastromyzon hagiangensis) is confined to small rivers and creeks within the Lo River drainage. The Ha Giang stone loach (Schistura hagiangensis) is known only from the Lo River in northern Vietnam (Ha Giang province). Nguyen’s stone loach (Balitora nigrocorpa) is known only from the Lo River drainage. The Mu River is located in north-western Vietnam (Lai Chai and Ha Giang provinces). The Laichow garra (Vinagarra laichowensis) is known only from a few specimens collected from the Mu River. The Mu spined loach (Cobitis guttata) is known only from a few specimens collected from the Mu River. The Gam River (Song Gam in Vietnamese) originates in Guangxi, China and flows through northern Vietnam (Tuyen Quang province). The Gam barb (Spinibarbus ovalius), Gam steed (Hemibarbus thacmoensis), Gam catfish (Hemibagrus chiemhoaensis), Nahang loach (Vanmanenia nahangensis), Gam river loach (Hemimyzon songamensis), Vancuong’s loach (Parabotia vancuongi), Kimluan’s loach (P. kimluani), Vanlan’s stone loach (Balitora vanlani), Vanlong’s stone loach (B. vanlongi), and Haithanh’s stone loach (B. haithanhi) are all known only from the Gam River drainage. The Chay River is located in northern Vietnam (Tuyen Quang province). The Chay torrent catfish (Nahangbagrus songamensis) is known only from six specimens collected from the Chay River.

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The Cau River (Song Cau in Vietnamese) is located in northern Vietnam (Bac Thai, Bac Giang, and Bac Ninh provinces). The Cau loach (Vanmanenia ventrosquamata) is known only from the Cau River drainage.

Miscellaneous Lakes, Rivers, and Marshes Lake Xigong (Xigong Hu in Chinese) is an endorheic (closed) mountain lake located 1300 m above sea level in the lower Tsangbo (Brahmaputra) River drainage, Tibet. The Xigong snowtrout (Schizothorax integrilabiatus) is confined to Xigong Lake and nearby mountain streams. Lake Langtso (Langtso Hu in Chinese) is an endorheic (closed) mountain lake located 4300 m above sea level in southern Tibet. The Langtso naked carp (Gymnocypris scleracanthus) is confined to Lake Langtso. Lake Xingjun (Xingyun Hu in Chinese) is a plateau lake located in southern China (Yunnan). Anderson’s silver minnow (Anabarilius andersoni) is confined to Lake Xingjun, where it is threatened by habitat degradation and overfishing. Lake Hulun (Hulun Hu in Chinese, and also known as Lake Dalai) is a large but shallow lake located in Inner Mongolia, northern China. The Hulun stickleback (Pungitius stenurus) is confined to Lake Hulun, where it is potentially threatened by declining water levels. Lake Yilong (Yi Long Hu in Chinese) is a large lake in Yunnan, southern China. The Yilong carp (Cyprinus yilongensis) and the Yilong silver minnow (Anabarilius macrolepis) both became extinct in 1981 when water extraction for agriculture caused the lake to dry up for an extended period. Lake Qiong (Qiong Hai in Chinese) is located on the Yunnan Plateau in south-western China. The Qiong carp (Cyprinus qionghaiensis) is confined to Qiong Lake, where it is very rare or possibly already extinct owing to pollution, overfishing, and competition with introduced species. Lake Thac Ba (Ho Thac Ba in Vietnamese) is a man-made lake located in north-eastern Vietnam (Yen Bai province). It was created by the construction of the Thac Ba hydroelectric plant in the 1960s, and named for the Ba Falls which preexisted the dam. The Thac Ba minnow (Tanichthys thacbaensis) and Thac Ba spiny eel (Mastacembelus thacbaensis) are both confined to Lake Thac Ba. Lake Rara is a deep, high-elevation lake located in the Himalayas of north-western Nepal. While set within Rara National Park and a designated Ramsar site, pollution and overfishing remain threats. The Rara snowtrout (Schizothorax raraensis) and the Nepalese snowtrout (S. nepalensis) are both confined to Lake Rara and adjoining streams. Lake Nui Coc (Ho Nui Coc in Vietnamese) is a man-made lake located in northern Vietnam. The Nui Coc mangrove goby (Mugilogobius nuicocensis) is known only from Lake Nui Coc, although it would appear that the species originates from another freshwater lake or river. Lake Ho (Ho Tay in Vietnamese) is located near Hanoi in northern Vietnam. The Lake Ho gudgeon (Hypseleotris

The Sino-Himalayan Region

hotayensis) and the Hanoi amblatikas (Toxabramis hotayensis) are both confined to Lake Ho. The Datangze Marsh is a small endorheic basin located in southern China (Yunnan). The large-stomached stone loach (Yunnanilus macrogaster), black stone loach (Y. niger), and Datangze stone loach (Y. paludosus) are all confined to the Datangze Marsh. The Upper Indus River originates high on the western Tibetan Plateau and flows more than 3000 km through the Himalayas of north-western India and north-eastern Pakistan. The Kumaon snowtrout (Schizothorax kumaonensis) is a rare species from the upper reaches of the Indus River drainage. The Qiantang River (also known as the Fuchun River) is located in eastern China (Zheijang). Tung’s gudgeon (Gobiobotia tungi) is confined to the Qiantang River. The Nho Que River (Song Nho Que in Vietnamese) is located in northern Vietnam and southern China. The Nho Que barb (Acrossocheilus baolacensis) is confined to the Nho Que River. The Ky Cung River is located in northern Vietnam (Lang Son province). The Ky Cung sharpbelly (Hemiculter elongatus) is confined to the Ky Cung River. The Tam Duong River is located in north-western Vietnam (Lai Châu province). The Tam Duong garra (Garra bibarbatus) is confined to the Tam Duong River.

Coasts and Satellite Islands This section includes the coast of eastern and south-eastern China and its satellite islands (including Taiwan and Hong Kong), along with the Japanese Archipelago. The short-tailed or Steller’s albatross (Phoebastria albatrus) came perilously close to extinction. It was hunted on an almost industrial scale for its feathers during the latter half of the nineteenth century, when by some estimates upwards of ten million birds were killed, and was reduced to just a couple of thousand by the early twentieth century. By the 1930s the only survivors were on Torishima, south of Japan, where hunting continued until the Japanese finally declared a ban in 1933. This measure appeared to have come too late, however. In 1949 the species was declared extinct on the island, although a small number of birds, most likely juveniles, were still thought to survive at sea. When 23 of the latter eventually returned to the island they were carefully protected, and by 1954 they began to breed. The population recovered steadily thereafter and began to disperse into other parts of their historical range. As of 2014 the total was estimated at 4200, the majority of which still nest on Torishima, but there are now additional nesting colonies on at least two other Pacific islands (Minami-kojima and Kita-Kojima in the East China Sea). Some limited breeding activity has also recently been reported on Kure and Midway in the Hawaiian Islands, and during the non-breeding season the birds once again range across the northern Pacific. The Japanese murrelet (Synthliboramphus wumizusume) is a marine bird that breeds on uninhabited islands of central and southern Japan and other coastal areas of the Korean

Peninsula and perhaps the Russian Far East, where it has been much reduced in number. The Chinese crested tern (Thalasseus bernsteini) historically bred in island colonies along the East China coast and wintered in Thailand, Indonesia, and the Philippines. The species was decimated during the twentieth century by excessive hunting and egg collection and thought to be extinct. In 2000, however, four pairs along with four chicks were rediscovered nesting in a greater crested tern (T. bergii) colony on an islet in the Matsu Islands, off the coast of Fujian province. Two other small island populations have since been found in China and one in South Korea, though the total number is still thought to be less than 50. Bennett’s water snake (Enhydris bennettii) is known only from a small number of specimens collected from the coastal salt marshes and estuaries of southern China and the island of Hainan. The harpist frog (Babina okinavana) is confined to Taiwan and to Ishigaki and Iriomote in the Yaeyama Islands of Japan. Anderson’s crocodile newt (Echinotriton andersoni) is currently known from six islands in the Ryukyu Islands. There are old records from Mount Kuanyinshan on Taiwan as well, but the species is presumed to be extirpated there. It is threatened by loss of habitat and overcollection for the international pet trade.

The Japanese Archipelago The Japanese Archipelago is a group of 6852 islands extending over 3000 km along the north-eastern coast of the Eurasian continent, from the Sea of Okhotsk to the Philippine Sea. The northernmost islands are centered in the temperate zone, but the southern extremity reaches the subtropics. They are strongly affected by the monsoons and are characterized by humidity, high rainfall, and warm temperatures, all of which produce a rich vegetation. The great number of high volcanoes and mountains add a subarctic zone to this spectrum of climates and gives rise to a wide range of habitats. As much as 80 per cent of Japan is filled by mountains, of which the highest, Mount Fuji, is 3776 m. Long densely populated, the country’s remarkable industrial development has inevitably damaged its natural habitats. However, the Japanese were also pioneering in conservation in eastern Asia, their deplorable whaling record notwithstanding. Ijima’s leaf warbler (Phylloscopus ijimae) breeds mainly on the Izu Islands south of the Japanese main islands, as well as on Nakanoshima in the Tokara Islands, from where it winters in the Ryukyu Islands and perhaps Taiwan and the Philippines. The species has declined significantly since the 1970s due to loss of habitat and pesticide use. The well earthworm goby (Luciogobius pallidus) and the cave goby (L. albus) are both confined to coastal and island cave systems throughout Japan. The Japanese Main Islands The Japanese main islands include the northernmost and four largest – Kyushu, Shikoku, Honshu, and Hokkaido – within the Japanese Archipelago.

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Figure 2.11 Japanese grey wolf (Canis lupus hodophilax). (Credit: Unknown.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The Japanese grey wolf (Canis lupus hodophilax) lived on Honshu, Shikoku, and Kyushu, where it was much persecuted because it preyed on cattle. The last of these small wolves seems to have been killed in 1904 or 1905. The Japanese black bear (Ursus thibetanus japanicus) was historically found on Honshu, Shikoku, and Kyushu, but has been extirpated from the latter island. The Japanese river otter (Lutra lutra whiteleyi) was historically widespread and common in Japan, and indeed during the 1880s could even be seen in Tokyo. Heavily hunted for its pelt as well as for use in ‘traditional medicine’, the subspecies underwent a significant decline in the early twentieth century and after that was only rarely recorded. It was officially declared extinct in 2012, although in 2017 a wild otter was photographed on Tsushima Island. The frosted mouse-eared bat (Myotis pruinosus) is found patchily across Honshu, Shikoku, and Kyushu, where it is threatened by loss of habitat. Endo’s pipistrelle (Pipistrellus endoi) is a type of bat confined to a handful of scattered localities in Honshu and Shikoku, where it is declining.

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The copper pheasant (Syrmaticus soemmerringii) was historically common on Honshu, Shikoku, and Kyushu, but has declined considerably due to overhunting. During the 1970s the number of birds taken went from c.800,000 to c.300,000 per year, but appeared to stabilize at around 100,000 per year during the 1990s. It is now considered uncommon and difficult to find, if not actually threatened. There are five subspecies. Soemmering’s copper pheasant (S. s. soemmerringii) is found in northern and central Kyushu. The scintillating copper pheasant (S. s. scintillans) is found in northern and central Honshu. The Shikoku copper pheasant (S. s. intermedius) is found in south-western Honshu and Shikoku. The Pacific copper pheasant (S. s. subrufus) is found in south-eastern Honshu and south-western Shikoku. The Ijimia copper pheasant (S. s. ijimae) is confined to southeastern Kyushu. The yellow bunting (Emberiza sulphurata) breeds only on the Japanese main islands, from where it is thought to migrate mainly to the Philippines in winter. Generally uncommon, it declined significantly during the twentieth century due to habitat destruction, pesticides, and hunting. The Japanese giant salamander (Andrias japonicus) is confined to the rivers of western Honshu, Shikoku, and Kyushu, where it has greatly decreased in recent decades due to overharvesting (for food and ‘traditional medicine’), habitat destruction and degradation, and hybridization with introduced Père David’s giant salamanders (A. davidianus). Three species of salamander (Hynobius) are threatened by loss of habitat and overcollection for the pet trade. Kato’s salamander (H. katoi) is confined to the mountains of southern Honshu. The Oita salamander (H. dunni) is confined to small areas of Kyushu and Shikoku. The Odaigahara salamander (H. boulengeri) is confined to areas of southern Honshu, Shikoku, and Kyushu. The amago salmon (Oncorhynchus masou macrostomus) is endemic to freshwater streams in western Japan (Honshu, Shikoku, and Kyushu). The deepbody bitterling (Acheilognathus longipinnis) is a type of cyprinid fish confined to four rivers in central and southern Japan. It is threatened by channelization, dams, pollution, and competition with introduced species. Smith’s bitterling (Rhodeus smithii) was historically widespread in Kyushu and western Honshu prior to World War II, but has declined dramatically due to loss of habitat and introduced fish species. Honshu Honshu is the largest and most populous island of Japan. The Etigo mole (Mogera etigo) is confined to the Echigo Plain on Honshu, where it is threatened by agricultural expansion. The Honshu noctule (Nyctalus furvus) is a type of bat known only from a few records. The Honshu short-tailed bat (Eptesicus japonensis) is found patchily in the mountains of central Honshu. It is

The Sino-Himalayan Region

threatened by loss of habitat and human disturbance of its roosting sites. Four salamanders of the genus Hynobius are endemic to Honshu, where they are threatened by loss of habitat and overcollection for the pet trade. Abe’s salamander (H. abei), the Hokuriku salamander (H. takedai), and the Hakuba salamander (H. hidamontanus) are all known only from a few areas in south-central Honshu. The Tokyo salamander (H. tokyoensis) is confined to a small area of south-central Honshu. The Kirikuchi char (Salvelinus japonicus) is a type of freshwater fish confined to two headwater streams within the Totsu River drainage in central Honshu. The stumpy bullhead (Pseudobagrus ichikawai) is confined to a few rivers in Honshu. The Tokyo bitterling (Tanakia tanago) is confined to an area near Tokyo where it is threatened by urbanization. Lake Biwa (Biwa-ko in Japanese) is located in west-central Honshu. It is the largest freshwater lake in Japan. The Biwa salmon (Oncorhynchus masou masou) is confined to Lake Biwa. The kissing loach (Parabotia curtus) is confined to Lake Biwa and a few rivers.

16 named islands are permanently inhabited. Much of the archipelago lies within the borders of Daisen-Oki National Park. The Oki salamander (Hynobius okiensis) is confined to Dogo Island in the Oki Islands. The Izu Islands The Izu Islands (Izu-shoto in Japanese) are a volcanic archipelago stretching south and east from the Izu Peninsula of Honshu. The Izu robin (Larvivora tanensis) is confined to the Izu Islands. The Izu thrush (Turdus celaenops) is mostly confined to the Izu Islands, although a few birds move to adjacent parts of the Honshu and the Shikoku Islands during the winters. There are also small numbers on Yakushima and Tokara in the northern Ryukyu Islands.

The Ryukyu Islands

Tsushima Tsushima is a small island located in the Korea Strait, approximately halfway between the Japanese mainland and the Korean Peninsula. The Tsushima tube-nosed bat (Murina tenebrosa) is known for certain only from a single specimen collected in 1962 on Tsushima, although a bat collected on Yaku Island may represent the same species. There is only a small patch of forest remaining on Tsushima and very few caves available for roosting.

The Ryukyu Islands (Ryukyu-shoto in Japanese, and also known as the Nansei Islands) consist of numerous large and small subtropical islands lying in a chain between southern Japan and Taiwan. The larger of these islands are volcanic in origin, with mountainous terrain, while the smaller ones are mostly formations of coral. All contain forests of leafy trees that flourish in the wet, subtropical climate. The Ryukyu tree rat (Diplothrix legata) is found only in the mountains of the western Amami islands and northern Okinawa. The Ryukyu mouse-eared bat (Myotis yanbarensis) is confined to mature forests on Okinawa, Amami, and Tokunoshima. The Ryukyu tube-nosed bat (Murina ryukyuana) is confined to Okinawa, Tokuno, and Amami islands. The Japanese long-fingered bat (Miniopterus fuscus) is known from Honshu (where it was last collected in 1933) and sporadically throughout the Ryukyu Islands. It is threatened by disturbance of its cave roosting sites. The Ryukyu woodcock (Scolopax mira) is confined to the islands of Amami, Kakeromajima, Tokunoshima, Okinawa, and Tokashikijima. Kam’s yellow pond turtle (Mauremys mutica kami) is confined to the Ryukyu Islands. The Ryukyu odd-scaled snake (Achalinus werneri) is confined to Amami, Kakeromajimia, and Tokunoshima in the Amami Islands and to Okinawa and Tokashijima in the Okinawa Islands. The sword-tailed newt (Cynops ensicauda) is confined to 12 islands in the Amami and Okinawa island groups. The Ryukyu Ayu-fish (Plecoglossus altivelis ryukyuensis) is confined to the Ryukyu Islands.

The Oki Islands The Oki Islands (Oki-shoto in Japanese) are an archipelago located in the Sea of Japan. Volcanic in origin, only 4 of the

The Amami Islands The Amami Islands (Amami-gunto in Japanese) are a group of small, limestone coralline islands located between southernmost

Hokkaido Hokkaido is the northernmost and second largest of the main Japanese islands. The nominate form of Blakiston’s eagle-owl (Bubo blakistoni blakistoni), the largest living owl in the world, is restricted to a small area on Hokkaido and to Kunashir Island in the southern Kuril Islands. Kyushu Kyushu is the most south-westerly of the four main Japanese islands. The Kyushu bitterling (Rhodeus atremius) is confined to northern Kyushu, where it is threatened by loss of habitat, pollution, and collection for the international aquarium trade. The Ariake dwarf icefish (Neosalanx regani) is confined to estuarine areas in north-western Kyushu, where it is threatened by dam construction.

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Kyushu and Okinawa in the northern Ryukyu Islands. They include Amami, Kikaijima, Kakeromajima, Yoroshima, Ukejima, Tokunoshima, Okinoerabujima, Yoronjima, Edateku, Sukomobanare, Yubanare, Kiyama, and Eniya. The Amami rabbit (Pentalagus furnessi) is a small, darkfurred species confined to Amami and Tokuno islands. Often called a living fossil, it is a survivor of a group of ancient rabbits that once lived on the Asian mainland but have long since died out. Prior to 1921 it was heavily hunted and trapped, but since 1921 it has been completely protected, although loss of habitat and introduced predators remain a threat. The Amami spiny rat (Tokudaia osimensis) is confined to a few areas on Amami Island. The Tokunoshima spiny rat (T. tokunoshimensis) is confined to mountainous areas on Tokunoshima Island. The Amami shrew (Crocidura orii) is confined to the islands of Amami, Tokunoshima, and Kakeromajima. The Amami woodpecker (Dendrocopos owstoni) is confined to mature hill forests on Amami Island. The Amami jay (Garrulus lidthi) is confined to Amami, Kakeroma, Uke, and Edateku. In the 1970s the population was estimated at around 5800, but underwent a major decline in the 1990s from which it appears to be recovering once more. The Amami thrush (Zoothera major) is known only from the central and western parts of Amami and from Kakeromajima (where it may now be extirpated). The Otton frog (Babina subaspera) is confined to Amami and Kakeroma islands, where it is threatened by habitat destruction and introduced predators. Two species of odorous frog (Odorrana) endemic to the Amami Islands are threatened by loss of habitat. The splendid odorous frog (Odorrana splendida) is confined to Amami. The Amami odorous frog (O. amamiensis) is confined to Amami and Tokunoshima. The Okinawa Islands The Okinawa Islands (Okinawa-shoto in Japanese) are a group of large and small islands located roughly 640 km south of the Japanese main islands and 500 km north of Taiwan. The Ryukyu wood pigeon (Columba jouyi) was endemic to Okinawa and the Kerama and Daito islands. Known only from a few specimens collected during the nineteenth century, it apparently declined rapidly (and inexplicably). It was last recorded on Okinawa in 1904 and on Daito in 1936. The Ryukyu black-breasted leaf turtle (Geoemyda japonica) is confined to Okinawa, Kume, and perhaps Tokashiki. In 1975 it was designated a National Natural Monument of Japan. It is threatened by illegal collection for the international exotic pet trade. Two species of ground gecko (Gonturosaurus) are endemic to the Okinawa Islands. The spotted ground gecko (G. orientalis) is confined to four small islands (Tonakijima, Tokashikijima, Iejima, and Akajima). Kuroiwa’s ground gecko (G. kuroiwae) is confined to Okinawa and three adjacent islands (Kourijima, Sesokojima, and Yagajijima).

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Holst’s frog (Babina holsti) is confined to scattered localities on Okinawa and Tokashikijima. Okinawa (Okinawa-jima in Japanese) is the main island of the Okinawa Island group and the largest of all the Ryukyu Islands. The island’s subtropical climate supports a dense forest in the north. The Okinawa spiny rat (Tokudaia muenninki) is confined to forests of northern Okinawa, where it is very rare. The Okinawa flying fox (Pteropus loochoensis) is known only from three specimens of unknown provenance, but is thought to have been collected on Okinawa during the nineteenth century. The Okinawa rail (Hypotaenidia okinawae) is a nearly flightless bird only just discovered in 1981 in a small area of forest in northern Okinawa. It is threatened mainly by the introduced Javan mongoose (Herpestes javanicus) and habitat fragmentation. The Okinawa woodpecker (Dendrocopos noguchii) is confined to the remaining virgin montane and coastal forests of Okinawa, where it nearly went extinct during the 1930s. In the early 1990s the total population was estimated at between 150 and 585. The Okinawa odorous frog (Odorrana narina) and Ishikawa’s odorous frog (O. ishikawae) are both confined to northern Okinawa. Namiye’s wart frog (Limnonectes namiyei) is confined to northern Okinawa. Kume Island (Kumejima in Japanese) is located west of Okinawa. Yamashina’s ground gecko (Goniurosaurus yamashinae) is confined to Kume Island, where it is threatened by loss of habitat. Kikuzato’s stream snake (Opisthotropis kikuzatoi) is an aquatic species confined to two hilly streams located in the north and south of the island, respectively. Intervening habitats have been completely converted for cultivated lands and residential areas. Iheya Island (Iheya-jima in Japanese) is located in the East China Sea, north-west of Okinawa. Toyama’s ground gecko (Goniurosaurus toyami) is confined to this single small island, where it is threatened by deforestation due to development. The Senkaku Islands (Senkaku-shoto in Japanese) are a group of uninhabited islands located north-east of Taiwan in the East China Sea. The Senkaku mole (Mogera uchidai) is known only from a single specimen collected on Uotsuri Island in 1991. The Kerama Islands (Kerama-shoto in Japanese) are a group of 22 islands located 32 km south-west of Okinawa. The Kerama sika deer (Cervus nippon keramae) is confined to Yakabi Island, where the total population is only about 30. The Miyako Islands The Miyako Islands (Miyako-retto in Japanese) are located in the southern Ryukyus, near Taiwan.

The Sino-Himalayan Region

The Miyako kingfisher (Todiramphus cinnamomina miyakoensis) is known only from a single specimen collected on Miyako Island in 1887. It likely became extinct soon after. The Miyako grass lizard (Takydromus toyamai) is confined to the Miyako Island group. Pfeffer’s reed snake (Calamaria pfefferi) is confined to Miyakojima and Irabujima. The Yaeyama Islands Located south-west of the Miyako Islands, the Yaeyama Islands (Yaeyama-shoto in Japanese) are the most remote part of Japan. The Ryukyu yellow-margined box turtle (Cuora flavomarginata evelynae) is confined to Ishigaki and Iriomote. Iwasaki’s slug-eating snake (Pareas iwasakii) is confined to Ishigaki and Iriomote, where it is uncommon. Utsunomiyas’ odorous frog (Odorrana utsunomiyaorum) and the greater odorous frog (O. supranarina) are both confined to Ishigaki and Iriomote. Iriomote Island (Iriomote-jima in Japanese) is the largest of the Yaeyama Islands. The Iriomote leopard cat (Prionailurus bengalensis iriomotensis) is confined to Iriomote, where the total population is around 100. The imperial stiphodon (Stiphodon imperiorientis) is a type of freshwater goby confined to the Nakama River on Iriomote.

Taiwan Taiwan is a large subtropical island located about 180 km off the south-eastern coast of China. The eastern two-thirds of the island consist mostly of rugged mountains running in five ranges from north to south, with the remaining third being a flat coastal plain. It once had a rich diversity of habitats, but the human population explosion has destroyed most of the forests. The Taiwanese clouded leopard (Neofelis nebulosa brachyurus) was historically confined to Taiwan, where it gradually driven into remote mountain areas by extensive hunting pressure and habitat destruction. The subspecies was last definitely reported from Tawu Mountain Nature Reserve in 1983, although in 1989 the skin of a young individual was found within the area of Taroko National Park. There was further possible evidence of survival near Yushan National Park during the 1990s, but extensive camera-trap surveys since have failed to reveal any further trace of it. Alleged sightings in 2018 are doubtful. The Taiwanese black bear (Ursus thibetanus formosanus) is confined to areas of montane forest. The Taiwanese sambar deer (Rusa unicolor swinhoii) is confined to Taiwan. The Taiwanese sika deer (Cervus nippon taiouanus) was completely extirpated in terms of free-ranging populations due to hunting by the end of the 1960s. Fortunately, a large captive population existed and, beginning in the 1980s, groups have been established in protected areas on the island.

The Taiwanese yellow-throated marten (Martes flavigula chrysospila) is a rare and little-known subspecies confined to Taiwan. The lesser Taiwanese shrew (Chodsigoa sodalis) is known only from five specimens collected from montane forest. The Mikado pheasant (Syrmaticus mikado) is confined to the mountains of central and southern Taiwan, where it was at one time seriously threatened by hunting. The species has since recovered somewhat and is protected within Yushan National Park and other areas. Swinhoe’s pheasant (Lophura swinhoii) is also confined to the mountains of central and southern Taiwan. Much reduced in number by hunting in the mid-1970s, it has since recovered somewhat but remains threatened outside of protected areas by habitat destruction. Styan’s bulbul (Pycnonotus taivanus) is a type of passerine bird confined to eastern and southern Taiwan. The Taiwanese yellow-margined box turtle (Cuora flavomarginata flavomarginata) is confined to Taiwan, where it is seriously threatened by agricultural expansion. Maki’s keelback (Hebius miyajimae) is confined to central and northern Taiwan. The Koshun worm snake (Argyrophis koshunensis) is known only from a few specimens collected in the 1930s. Two tree frogs of the genus Zhangixalus are threatened by loss of habitat. The farmland tree frog (Z. arvalis) is confined to agricultural areas in south-western Taiwan. The orangebellied tree frog (Z. aurantiventris) is found patchily throughout the islands. The Taipa frog (Rana longicrus) is confined to northern and western Taiwan. The Taiwanese groove-toed frog (R. sauteri) is confined to lowland areas of western Taiwan. Both are threatened by habitat destruction and pollution. Stejneger’s paddy frog (Micryletta steinegeri) is known only from a few areas in central and southern Taiwan. Three salamanders of the genus Hynobius are endemic to the Central Mountains, where they are threatened by loss of habitat. The Formosa salamander (H. formosanus), Taichu salamander (H. sonani), and the Arisan salamander (H. arisanensis) are all very rare and confined to a few areas of undisturbed montane conifer forest. The Taiwanese salmon (Oncorhynchus masou formosanus) is a very rare, landlocked form known only from a few streams. The Taiwanese shovel-jaw carp (Onychostoma alticorpus) is confined to five river drainages on Taiwan, where it is threatened by siltation, dam construction, and pollution. The Taitung river loach (Hemimyzon taitungensis) is confined to streams of the Central Mountain Range. The Taiwanese hillstream loach (Sinogastromyzon puliensis) is confined to a few rivers in western Taiwan. Orchid Island Orchid Island (Lan Yu in Chinese) is a tiny island located off the south-eastern coast of Taiwan.

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The Botel gecko (Gekko kikuchii) is largely confined to coastal cliffs on the southern part of Orchid Island, where it remains common.

Hong Kong Hong Kong and its various satellite islands are located on the eastern side of the Pearl River estuary in southern China. Originally a lightly populated area of farming and fishing villages, today it is one of the most densely populated places on Earth. Remaining undeveloped terrain is largely hilly to mountainous with very little flat land. Hollinrake’s bronzeback tree snake (Dendrelaphis hollinrakei) is confined to Shek Kwu Chau, a small island located west of Hong Kong. Lazell’s blind snake (Indotyphlops lazelli) is known only from two specimens collected in a drainage ditch on Hong Kong Island. Romer’s shrub frog (Liuixalus romeri) was historically restricted to the islands of Lantau, Lamma, Chek Lap Kok, and Po Toi. With the construction of an airport on Chek Lap Kok, however, it was decided to translocate that population to suitable sites on Hong Kong Island and the Chinese mainland.

Cat Ba Island Cat Ba Island is the largest of the 367 islands of the Ca Ba Archipelago in Ha Long Bay, off the north-eastern coast of Vietnam. Approximately half its area is covered by a national park. The Cat Ba langur (Trachypithecus poliocephalus) is confined to a small area of Cat Ba Island mostly within the national park, where in 2006 the total population was just 64. The Cat Ba tiger gecko (Goniurosaurus catbaensis) is known only from Cat Ba Island, although it may possibly occur as well on other islands in Ha Long Bay.

Balance for the Sino-Himalayan Region Archaelogical evidence suggests that early hominids inhabited China some 2.25 million years ago. Remains of Peking Man, a subspecies of Homo erectus who used fire, were first discovered within the Zhoukoudian Cave, near Beijing in northern China, in 1926. He and his kind lived there in a cool, steppe, and forest region between 780,000 and 680,000 years ago alongside rhinos, elephants, bears, and big cats. The first modern human remains were discovered in Fuyan Cave, Hunan and date from 125,000 to 80,000 years ago. According to Chinese tradition the first dynasty emerged around 2100 B . C .. A Palaeolithic culture from around 30,000 B . C . constitutes the first known inhabitants of the islands of Japan. Owing to this long period of habitation, the Sino-Himalayan Region as a whole has been the scene of habitat destruction on a massive scale. Once largely covered by forests, today only a tiny fraction remains. One notable consequence of this is that they are no longer there to control the rivers, which frequently flood the lowlands, with catastrophic results.

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While there had been contact between China and the Western world for many centuries by way of the Silk Route, it was the Portuguese who were the first modern Europeans to truly explore the region. In 1513 Jorge Álvares became the first to reach China by sea, landing on Nei Lingding Island in the Pearl River delta. Thirty years later, in 1542 or 1543, an expedition reached the Osumi Islands of southern Japan, and in 1589 João da Gama visited ‘Yezo’ (Hokkaido). Meanwhile, further south, the missionary Bento de Góis travelled overland from India to China between 1602 and 1606 by way of Afghanistan and the Pamirs. The priest and explorer António de Andrade became the first known European to cross the Himalayas, reaching Tibet in 1624. Shortly thereafter, the Jesuit missionaries Estêvão Cacella and João Cabral would also cross the Himalayas and become the first to enter both Bhutan and Nepal. Dutch and Austrian expeditions as well would follow during the later seventeenth century. During the eighteenth century this part of Asia was opened up for trade, even as certain kingdoms remained stubbornly closed off to foreigners. In 1904 the British explorer Francis Younghusband led a notable expedition to Tibet, and in 1913 his countrymen Frederick Bailey and Henry Morshead explored the Tsangpo Gorge and the route of the Yarlung Tsangpo (upper Brahmaputra) River. More recently, in 1953 Edmund Hillary and Tenzing Norgay became the first people to ascend Mount Everest, and Lino Lacedelli and Achille Compagnoni the first to ascend K2 during the Italian Karakoram Expedition of 1954. While this area of Asia has long been heavily populated, it was until the mid-twentieth century that a truly massive population explosion began, one that shows no signs of levelling off. As a result, China’s lakes and rivers are heavily polluted. Moreover, a series of monumental hydroelectric projects have dammed the great rivers to the detriment of the fauna as well as the fertility of the surrounding plains. Most seriously of all, at least in terms of native species, is the overcollection of virtually anything that moves for both human consumption as well as ‘traditional medicine’, which is discussed in the introductory sections. In recent historical time (i.e. since A . D . 1500), the SinoHimalayan Region has lost at least 14 species/1 subspecies of vertebrates. Among the extinct forms, 1 subspecies is a mammal, 1 species is a bird, 1 species is an amphibian, and 12 species are freshwater fishes. Another 23 species/2 subspecies are possibly extinct. In addition, there are 606 species/58 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 92 species/42 subspecies are mammals, 61 species/9 subspecies are birds, 65 species/3 subspecies are reptiles, 181 species are amphibians, and 208 species/4 subspecies are freshwater fishes.

The Saharo-Arabian Region

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

~ species

2 species

~ species

92 species

1 subspecies

2 subspecies

~ subspecies

42 subspecies

1 taxon

4 taxa

~ taxa

134 taxa

1 species

~ species

~ species

60 species

~ subspecies

~ subspecies

~ subspecies

9 subspecies

1 taxon

~ taxa

~ taxa

69 taxa

~ species

1 species

~ species

65 species

~ subspecies

~ subspecies

~ subspecies

3 subspecies

~ taxa

1 taxon

~ taxa

68 taxa

1 species

1 species

~ species

181 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

1 taxon

1 taxon

~ taxa

181 taxa

12 species

19 species

~ species

208 species

~ subspecies

~ subspecies

~ subspecies

4 subspecies

12 taxa

19 taxa

~ taxa

212 taxa

14 species

23 species

~ species

606 species

1 subspecies

2 subspecies

~ subspecies

58 subspecies

15 taxa

25 taxa

~ taxa

664 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

The Saharo-Arabian Region The Saharo-Arabian Region is largely comprised of mountains and deserts and can be considered as something of a transition zone between the Palearctic, Afrotropical, and Indo-Malaysian realms. As here defined, it includes all of North Africa from the Sahara Desert to the Mediterranean and Atlantic coasts (including the Canary and Madeira islands as well as Cyprus), the Arabian Peninsula and the southern parts of the Levant, and south-western Asia east as far as the Indus River and north to the mountains of Central Asia. Once lush and green, most of this region is now semi-desert, although the topographic diversity gives the Mediterranean coastal strip at least a variety of habitats. There are stretches of maquis vegetation, small woods and, on the slopes of the Atlas Mountains, some temperate forests. The deteriorating valley of the northern Nile still has green areas. There are still many forms of life in the semidesert, but seen in the perspective of thousands of years, the impoverishment of the fauna, which has affinities to both Eurasia and the rest of Africa, is striking. The Atlas brown bear (Ursus arctos crowtheri) is known from fossil records collected between Morocco and Libya, along with numerous accounts of live animals from Roman times up to the latter half of the nineteenth century. Africa’s only bear, the last survivor is thought to have been killed by hunters in 1890. The Syrian brown bear (U. a. syriacus) is a small, light-coloured subspecies that was historically found

across a wide area of the Middle East from the Levant and Anatolia to Turkmenistan. Long since exterminated by hunting over most of this range, it still survives in parts of Iraq, Iran, and Turkey, primarily in mountainous regions. In the early twenty-first century bear tracks were also recorded from the Anti-Lebanon Mountains of Syria. The Balochistan black bear (U. thibetanus gedrosianus) has been driven from its optimal coniferous forest habitats in the mountains to low, arid hills with thickets of acacias and euphorbias, where it has become very rare, apparently owing to paucity of natural food and to conflicts with man, because it is obliged to raid crops to be able to feed. The Barbary macaque (Macaca sylvanus) is the only surviving primate in Africa north of the Sahara Desert, the only native species of primate in Europe and the only member of its genus to be found outside of Asia. It was once an inhabitant of much of Europe and all of North Africa from Libya to Morocco, but is now confined to small, relict patches of high-altitude cedar and oak forests across the Atlas Mountains as well as rocky slopes and coastal scrub in Algeria and Morocco. A famous, semi-wild colony lives on the Rock of Gibraltar, which is a long-established introduced population. The North African population of the African lion (Panthera leo leo), often known as the Barbary lion, was once quite common in North Africa including both the Atlas

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Mountains and the Mediterranean littoral, extending into Egypt where they historically lived along the Nile and as far east as the Sinai Peninsula. In the fourteenth century B . C . the pharaoh Thutmose IV hunted lions in the hills near Memphis. Later, lions for the Roman arenas were constantly shipped to Rome, where according to Pliny hundreds at a time were exhibited. The lions in the menagerie at the Tower of London during the Middle Ages were Barbary lions, and during the nineteenth and early twentieth centuries they were commonly kept in zoos and circuses. In the wild state they still existed from Libya to Morocco up to about 250 years ago, and were even fairly numerous in Tunisia. However, the arrival of firearms, coupled with bounties given for each lion shot, resulted in a very rapid decline. They still occurred along the coast of Libya until the beginning of the eighteenth century. The last Tunisian individual was killed in 1891. In Algeria the history is similar, with the last one also likely killed about 1891 (a reported sighting in 1956 notwithstanding). In Morocco the last one was shot in 1942 in the Middle Atlas, although populations perhaps persisted up to the 1960s in some remote montane areas. There are still a few zoo animals with some Barbary lion ancestry, but whether there is enough genetic material to resurrect the subspecies is unknown. It nevertheless survives in West Africa and the northern parts of Central Africa, as well as in India. The African leopard (Panthera pardus pardus) is the most widespread leopard subspecies, occurring throughout subSaharan Africa and in a variety of habitats from open arid savanna to dense lowland and montane rainforest. Unfortunately, the subspecies has disappeared from many of its former areas due to illegal hunting for its fur (which has a great commercial as well as trophy value), in addition to

poisoning and loss of habitat. This is certainly true north of the Sahara, where the so-called Barbary leopard has been wholly or perhaps entirely eliminated. Up until the latter part of the twentieth century populations were still known to occur in certain Atlas Mountain areas of Morocco, Algeria, and perhaps Tunisia, as well as in parts of southern Egypt, where they continued to be intensively hunted. Leopards were given legal protection in 1948, but those who hunt them invariably claim that they are acting in self-defence or to protect livestock. Apart from human persecution the disappearance or decimation of several species of antelope has also resulted in leopard declines in North Africa. Its last strongholds were in the mountains of Morocco and Algeria, but the animals are now believed to have been extirpated from the former country and may now be entirely restricted to a few survivors in the Ahaggar Mountains. None have actually been seen in the wild for many years. The cheetah (Acinonyx jubatus) is a large spotted cat notable for being the world’s fastest land animal. Historically found across Africa, Asia, and Europe, it has been mercilessly hunted and eliminated from most of its former range. The north-west African cheetah (A. j. hecki) is found patchily in Algeria, Mali, Niger, Benin and Burkina Faso. The serval (Leptailurus serval) is species of spotted cat that is widespread in Africa overall but highly threatened north of the Sahara. Formerly found throughout the Mediterranean coast region in both semi-deserts and cork-oak forests, today it is known to occur only in Morocco and possibly Algeria, with a reintroduced population (from East African stock) in Tunisia. The African wild ass (Equus africanus) is believed to be the ancestor of the domestic donkey, which is usually placed

Figure 2.12 Photo of a live male ‘Barbary lion’ taken in Algeria in 1895. (Credit: Alfred Edward Pease.)

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within the same species (E. a. asinus). Three other subspecies formerly lived throughout the deserts and arid areas of northern Africa. One of these, the Atlas wild ass (E. a. atlanticus) from what is now Algeria, Tunisia, and Morocco, appears to have been driven extinct by Roman sport hunting some time after A . D . 300. The Nubian wild ass (E. a. africanus) is also presumed to be extinct. Historically it lived in the Nubian Desert of north-eastern Sudan from east of the Nile River to the Red Sea, and south to the Atbarah River and northern Eritrea. Long pursued by humans for domestication, it was not until modern firearms came into general use that the decline of this form became serious. It was frequently slaughtered by nomads who accused it of consuming grass needed for their goats (in actual fact, of course, it was the goats that were the real problem). By the mid-twentieth century only a few small herds survived in remote areas of north-eastern Sudan and northern Eritrea. Despite legal protection these too were hunted down and annihilated. The last confirmed sightings were in the Barka Valley and the border area between Eritrea and Sudan during aerial surveys in the 1970s. Two populations outside the historic range potentially survive. A DNA study of feral donkeys on the Caribbean island of Bonaire revealed that these animals are very close to known Nubian wild asses and very different from other forms. The other alleged population, in Gebel Elba National Park on the Egypt/Sudan border, has yet to be scientifically investigated. The bubal hartebeest (Alcelaphus buselaphus buselaphus) was once an abundant antelope across northern Africa from Morocco to Egypt and perhaps historically extending into the

Levant. Large herds were still reported from Morocco north of the Atlas Mountains in 1738, and ‘wild oxen’ which may or may not have been these animals were mentioned as living in the Tassili Mountains of the central Sahara in 1850. In any event the subspecies declined sharply during the nineteenth century, particularly after the French conquest of Algeria when entire herds were massacred by the colonial military. By 1867 it was confined to the mountain ranges of northwestern Africa. It disappeared from the Tunisian Atlas in 1870, with the last individual from that country shot in 1902 near Tataouine. The remaining population was confined to the Western Atlas of Morocco and Algeria, where the last herd, numbering only 15 animals, was wiped out by hunters in 1917. It probably disappeared from Algeria around the same time. The last captive specimen died in the Paris Zoo in 1923, and the last known individual of all was shot in Morocco in 1925. There are reports in the literature of survivors having been shot between 1945 and 1954 in Algeria, and in south-western Morocco in 1945. However, this seems unlikely, as numerous attempts to locate the animals in the 1920s and 1930s had been unsuccessful even in areas where they had once been common. The mountain gazelle (Gazella gazella) occurs sporadically in a range of habitats within Israel, the Palestinian Territories and Jordan from the Golan Heights through central Israel, the Jordan Valley, and the northern Negev Desert. There have been no records from Syria since the 1970s (although a small number was recently seen in Turkey, near the Syrian border), or from the Sinai since 1932. In Lebanon the species was believed to have been extirpated after 1945, until three

Figure 2.13 Nubian wild ass in the Berlin Zoo, 1899. (Credit: Berlin Zoological Gardens.)

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The Palearctic Realm Figure 2.14 Female bubal hartebeest photographed in the London Zoo, 1895. (Credit: Lewis Medland.)

individuals were seen in the Barouk Mountains in 1998. The total number in Israel was estimated at around 10,000 in the late 1990s, but appears to have declined sharply since then. Populations in other areas are negligible. The Persian fallow deer (Dama mesopotamica), the largest of the fallow deer group, formerly occurred throughout the Near and Middle East. Owing to the fact that most of the woodlands in this region have disappeared, this species also vanished nearly everywhere. By 1875 the range had been reduced to just south-western and western Iran. The surviving deer were thought to have been driven extinct by intensive hunting by the 1940s, but a small population of about two dozen was rediscovered in the western foothills of the Zagros Mountains in south-western Iran in 1956, and a number brought to the Opel Zoo in Kronberg, Germany, to form the nucleus of a captive breeding programme. The last seven wild deer were captured in the mid-1960s and shipped to a fenced reserve close to the Caspian Sea in north-eastern Iran. In 1978, during the upheavals of the Iranian Revolution, conservationists transported the latter into Israel for safekeeping. Despite all these efforts the species remains in a precarious state, at least in the wild, inhabiting as it does an island in Lake Urmia (northwestern Iran), parts of northern Israel (where they had been reintroduced in the 1990s), and perhaps two small protected areas in south-western Iran. The Barbary red deer (Cervus elaphus barbarus), also known as the Barbary stag, is the only deer native to Africa. Fossil remains indicate that in prehistoric times it was found as far south as the central Sahara, but retreated along with the

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forests. In Roman times it was still common all over Tunisia, northern Algeria, and parts of Morocco, but declined with further loss of forests and extensive hunting. It suffered grievously in the mid-twentieth century during the Algerian War, when poaching and slaughter were uncontrolled. In recent years it has recovered somewhat owing to better protection, but remains confined to a few areas of the Atlas Mountains in Morocco (where it was reintroduced in the 1990s after having been extirpated), north-eastern Algeria, and western Tunisia. The aoudad (Ammotragus lervia), a type of wild sheep, is divided into six subspecies found patchily across Africa in both mountainous and desert areas. All are threatened by hunting and habitat destruction. The Saharan aoudad (A. l. sahariensis) is the most widespread, occurring as it does in southern Morocco, Western Sahara, southern Algeria, south-western Libya, Sudan, Mali, Niger, Mauritania, and northern Chad. Blaine’s aoudad (A. l. blainei) was historically found from western Sudan to the Red Sea coast, but is now most likely confined to the Red Sea Hills of eastern Sudan and perhaps in south-eastern Libya. The Arabian pipistrelle bat (Pipistrellus arabicus) is known only from a small area in south-eastern Saudi Arabia and north-eastern Oman, and another in southern Iran. Hanaki’s dwarf bat (P. hanaki) is known only from a small area in north-eastern coastal Libya. The lappet-faced vulture (Torgos tracheliotos) is divided into two subspecies. The Asian lappet-faced vulture (T. t. negevensis) is confined to the Negev Desert and to the Sinai and Arabian peninsulas, where it is threatened by habitat

The Saharo-Arabian Region

destruction, human disturbance of its nesting sites and pesticide poisoning. Two subspecies of European turtledove (Streptopelia turtur) are to be found within the Saharo-Arabian Region, where they are threatened by loss of habitat and hunting. Hartert’s turtledove (S. t. arenicola) is widespread from north-western Africa to western Asia, while the Egyptian turtledove (S. t. rufescens) is found in Egypt and northern Sudan. The Basra reed warbler (Acrocephalus griseldis) breeds primarily in the Mesopotamian marshes of south-eastern Iraq, as well as in the Hula Valley of northern Israel and most likely in south-western Iran as well. It winters in Sudan, South Sudan, Ethiopia, Somalia, south-eastern Kenya, eastern Tanzania, southern Malawi, and Mozambique. The neardestruction of its breeding habitat during the latter part of the twentieth century resulted in a massive decline, and the species remains seriously threatened. Jerdon’s babbler (Chrysomma altirostre) is divided into three subspecies found in isolated pockets throughout South Asia. The Sind Jerdon’s babbler (C. a. scindicum) is confined to wetlands within the Indus River drainage of eastern Pakistan. The Egyptian spiny-tailed lizard (Uromastyx aegyptia) is divided into three subspecies. The nominate form (U. a. aegyptia) occurs in north-eastern Africa, Jordan, and extreme north-western Saudi Arabia. The small-scaled Egyptian spiny-tailed lizard (U. a. microlepis) is found in Saudi Arabia, Oman, Yemen, the United Arab Emirates Qatar and Kuwait, while Leptien’s Egyptian spiny-tailed lizard (U. a. leptieni) is found in the Hajar al-Gharbi Mountains of Oman and in north-eastern United Arab Emirates. All are threatened by loss of habitat. Blanc’s spiny-toed lizard (Acanthodactylus blanci) is confined to a few areas of northern Tunisia and north-eastern Algeria, where it is threatened by loss of habitat. Manuel’s cylindrical skink (Chalcides manueli) is confined to a few localities in central coastal Morocco. The small threetoed cylindrical skink (C. minutus) is found in north-eastern Morocco and north-western Algeria. Both are threatened by loss of habitat. Varaldi’s spadefoot toad (Pelobates varaldii) is confined to a few localities on the coastal plains of north-western Morocco.

Mountains and Highlands The major mountain ranges of the Saharo-Arabian Region include the Atlas Mountains of north-western Africa, the Al Hajar Mountains in the Arabian Peninsula, the Hindu Kush, and the Levantine, Kopet Dag, Alborz, and Zagros mountains in the Near and Middle East. Two subspecies of wild goat (Capra aegagrus) are found in the mountains of southern Pakistan, where they are seriously threatened by uncontrolled hunting. The Sindh wild goat (C. a. blythi) is confined to a few areas of south-western Pakistan. The Chiltan wild goat (C. a. chialtanensis) was, by the early 1970s, confined to four or five small populations in southern

Pakistan (Balochistan province). These have since been reduced by uncontrolled hunting to a single surviving population in Hazarganji-Chiltan National Park. The Nubian ibex (Capra nubiana) is a mountain-dwelling goat formerly found across North Africa and the Middle East. Like others of its kind it has been much reduced as a result of poaching, and now survives only in scattered populations in Egypt, Israel, Jordan, Oman, Saudi Arabia, Sudan, Yemen, and possibly Syria. It appears to have been extirpated from Eritrea (where a special reserve had been declared for it in 1959), as well as from Lebanon. Only a few thousand at most remain in the wild. The Aïr aoudad (Ammotragus lervia angusi) is a type of wild sheep confined to the Aïr and Termit massifs of northern Niger. The Queen of Sheba’s gazelle (Gazella bilkis) is known from only five specimens collected in the mountains of Yemen in 1951. A somewhat doubtfully valid taxon, it is nevertheless clear that the gazelle population from this area, which was described as common in the 1950s, is now long extinct. Val’s gundi (Ctenodactylus vali) is a type of rodent confined to two small, widely separated areas in North Africa, the first in north-eastern Morocco and north-western Algeria, and the other in north-western Libya. Zarundny’s jird (Meriones zarudnyi) is confined to semiarid highland areas in north-eastern Iran and northern Afghanistan. Setzer’s mouse-tailed dormouse (Myomimus setzeri) is known only from a few specimens collected in the mountains of eastern Turkey and north-western Iran. The Persian wood mouse (Apodemus avicennicus) is known only from a single locality in the Shirkouh Heights of central Iran. Père David’s mole (Talpa davidiana) is known from two localities on the Iranian Plateau of north-western Iran (Kurdistan province) and south-eastern Turkey. The Persian vole (Microtus irani) is known only from four specimens collected in the 1920s from a city garden in Shiraz (Fars province). Butler’s owl (Strix butleri) is a likely cliff-nesting species known only from a few specimens collected or observed in the mountains of Oman, the United Arab Emirates, and perhaps Iran. The Saharan turtledove (Streptopelia turtur hoggara) is confined to the Aïr Massif of northern Niger and the Hoggar Mountains of southern Algeria. The Persian toad-headed agama (Phrynocephalus persicus) is known from a few fragmented populations in the northwestern and western parts of the Central Plateau in Iran, with an additional report from the highlands of southern Azerbaijan. Schmitz’s agama (Trapelus schmitzi) is known only from three specimens, two of which were collected from the Tassili n’Ajjer Mountains of south-eastern Algeria in around

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1900 and 1952 and the third from the Ennedi Mountains of north-eastern Chad. The thick-tailed rock gecko (Bunopus crassicauda) is known only from a few specimens collected from the northwestern part of the Central Plateau in Iran, although it may occur as well in north-eastern Syria. The Iranian mountain viper (Montivipera albicornuta) is confined to the Zanjan Valley and surrounding mountains in north-western Iran. Two subspecies of Anatolian dwarf racer (Eirenis thospitis) is divided into two subspecies found in the mountains of south-eastern Anatolia. The Van dwarf racer (E. t. thospitis) is confined to a small area near Lake Van, while the Hakkari dwarf racer (E. t. hakkariensis) is known only from two localities near the cities of Siirt and Hakkar. Bury’s blind snake (Myriopholis burii) is known only from a few specimens collected from the mountains of southwestern Saudi Arabia and south-western Yemen. It has not been recorded since 1946. Eichwald’s toad (Bufo eichwaldi) is confined to the Talysh and Alborz mountains of Azerbaijan, as well as parts of northern Iran.

The Atlas Mountains

form a natural barrier between the Mediterranean and the Sahara Desert. The Algerian nuthatch (Sitta ledanti) is known only from four localities in the Tell Atlas of Algeria. Several subspecies of North African fire salamander (Salamandra algira) are found patchily in the Atlas Mountains. The Algerian fire salamander (S. a. algira) is found in a few areas across the Tell Atlas, while the Beni Snassen fire salamander (S. a. spelaea) is confined to the Beni Snassen Massif in north-eastern Morocco. The Edough ribbed newt (Pleurodeles poireti) is confined to the Edough Massif in north-eastern Algeria. The Middle Atlas Range The Middle Atlas is located in north-central Morocco. The Middle Atlas fire salamander (Salamandra algira atlantica) is confined to the northern and central Middle Atlas. The Rif Mountains The Rif Mountains are located in northern Morocco. The Tingitana fire salamander (Salamandra algira tingitana) is confined to the Tingitana Peninsula. The splendid Moroccan fire salamander (S. a. splendens) is confined to the central and western Rif Mountains.

The Atlas Mountains are a series of subranges that stretch across north-western Africa, extending about 2500 km through Algeria, Morocco, and Tunisia. Separating Mediterranean and Atlantic coastlines from the Sahara Desert, these mountains are home to a number of plant and animal species unique in Africa and often more like those found in Europe. Many of them (particularly large mammals) are seriously threatened and a few have already gone extinct. The Barbary sheep or Atlas aoudad (Ammotragus lervia lervia) inhabits the Atlas Mountains of Morocco, northern Algeria, and northern Tunisia, where it is threatened mainly by poaching. Cuvier’s gazelle (Gazella cuvieri) was formerly abundant throughout the Atlas Mountains and neighbouring ranges in Morocco, Algeria, Tunisia, and Western Sahara. Mercilessly hunted during the twentieth century by local tribesmen using modern rifles, often from automobiles, in 1932 it was already said to be one of the rarest of all gazelles. By 1970 only a few small herds were left, scattered among isolated forest fragments and high plateaus. Populations have recovered since then with better protection, but the species remains threatened by loss of habitat. The small-fingered psammodromus (Psammodromus microdactylus) is a type of lizard found in relictual, isolated populations in the Rif, High, and Middle Atlas mountains of northern and central Morocco.

The Levantine Mountains, as here defined, include those ranges found within Israel, Palestine, southern Syria, Lebanon, Jordan, and Egypt’s Sinai Peninsula. The Syrian serin (Serinus syriacus) is a type of passerine bird that breeds in the mountains of Syria, Lebanon, Israel, and Jordan, with some populations making a winter migration to lower-elevation desert areas as far away as the Nile Valley. It has declined due to drought and habitat destruction. The Levantine bent-toed gecko (Mediodactylus amictopholis) appears to be largely restricted to Mount Hermon in southwestern Syria, with a single specimen having been collected from the Mount Lebanon range in central Lebanon. Kulzer’s lizard (Phoenicolacerta kulzeri) is divided into three ill-defined subspecies (P. k. kulzeri, P. k. petraea, and P. k. khasaliensis) found in scattered areas within the Levantine Mountains. All are threatened by loss of habitat. Fraas’ lizard (Parvilacerta fraasii) is known only from the mountains of Lebanon and perhaps adjacent Syria. The Lebanon mountain viper (Montivipera bornmuelleri) is confined to the high mountains of Lebanon and southern Syria.

The Tell Atlas The Tell Atlas runs parallel to the Mediterranean coast in north-eastern Morocco and northern Algeria. The mountains in this system have an average elevation of about 1500 m and

The highest mountains on the eastern Arabian Peninsula, the Al Hajar Mountains are located in north-eastern Oman and eastern United Arab Emirates. The Arabian tahr (Arabitragus jayakari) is a type of ungulate confined to the Al Hajar Mountains, where it prefers

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The Levantine Mountains

The Al Hajar Mountains

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north-facing slopes between 1000 and 1800 m. The estimated world population is less than 5000. The Jebal Akhdar leaf-toed gecko (Asaccus montanus) is confined to the higher elevations of the Jebal Akhdar in Oman.

The Gorgan salamander (Iranodon gorganensis) is confined to the Shir-Abad Cave and the stream flowing from it, approximately 60 km east of Gorgan in the eastern Alborz Mountains.

The Sarawat Mountains

The Zagros Mountains

The Sarawat Mountains (Jibal as-Sarawat in Arabic) are located in western Saudi Arabia and western Yemen, running parallel to the Red Sea.

The Zagros Mountains are a long mountain range in western Iran, north-eastern Iraq, and south-eastern Turkey. The Esfahan mouflon (Ovis gmelini isphahanica) is confined to a very small area of the Zagros Mountains south-west of Esfahan in west-central Iran. Firouz’s jerboa (Allactaga firouzi) is known only from a few areas of montane steppe in west-central Iran (Isfahan province). The Iraqi bent-toed gecko (Mediodactylus heteropholis) is known only from two specimens collected in the foothills of western Iran and north-eastern Iraq. Helen’s bent-toed gecko (Microgecko helenae) is known only from a few, widely scattered records. The spider-tailed horned viper (Pseudocerastes urarachnoides) is known only from the Zagros Mountains of western Iran. Rechinger’s dwarf racer (Eirenis rechingeri) is known only from a single locality within the Zagros Mountains of southwestern Iran. The Luristan newt (Neurergus kaiseri) is an entirely aquatic species confined to the central Zagros Mountains, where it is heavily collected for use in the international pet trade. The Zagros cave garra (Garra typhlops) is a blind, unpigmented freshwater fish confined to caves in the Zagros Mountains of Iran. The Kurdistan mountain catfish (Glyptothorax kurdistanicus) is known only from a single specimen collected in the early twentieth century from the Little Zab River, in the Zagros Mountains of Iran. The Zagros blind loach (Eidenemacheilus smithi) is confined to a single aquifer in the Karun River drainage, in the Zagros Mountains of Iran.

The Asir Range The Asir Range (Jibal al-Asyr in Abrabic) is located in southwestern Saudi Arabia. The Asir magpie (Pica asirensis) is confined to the Asir range, where the total population is less than 300. The Yemen warbler (Sylvia buryi) is confined to the Asir range. Gasperett’s snake (Lytorhynchus gasperetti) is known only from a few specimens collected during the 1970s primarily from the south-eastern Asir Mountains. The Asir garra (Garra buettikeri) is a type of freshwater fish confined to a few small streams in the eastern Asir range. The Hijaz Range The Hijaz Range (Jibal al-Hijaz in Abrabic) is located in western Saudi Arabia. The Arabian himri (Carasobarbus apoensis) is a type of freshwater fish known from six isolated wadis within the Al Hijaz Mountains. An expedition in 2013 failed to find any specimens from three of these sites.

The Kopet Dag Mountains Also known as the Turkmen-Khorasan range, the Kopet Dag Mountains extend about 650 km along the border between Turkmenistan and Iran south-east of the Caspian Sea. The Turkmenean wild goat (Capra aegagrus turcmenica) occurs in scattered populations in the central Kopet Dag Mountains. In 1986 the total population was estimated at around 7000 and decreasing. Steiner’s wall lizard (Darevskia steineri) is known only from a small area of the Kopet Dag Mountains in north-eastern Iran.

The Alborz Mountains The Alborz Mountains (also spelled Alburz, Elburz, or Elborz) are located in northern Iran, where they stretch from the border of Azerbaijan along the western and entire southern coast of the Caspian Sea before finally running north-east and merging with the Aladagh Mountains. The Alborz brush-tailed mouse (Calomyscus grandis) is confined to montane forests of the Alborz Mountains. The Iranian mountain steppe viper (Vipera ebneri) is known only from alpine meadows in the Alborz Mountains. Latif’s mountain viper (Montivipera latifii) is confined to a small area in the upper Lar River drainage of the central Alborz Mountains.

The Hindu Kush The Hindu Kush is an 800-km-long mountain range that stretches near the Afghan–Pakistan border, from central Afghanistan to northern Pakistan. The Balochistan forest dormouse (Dryomys niethammeri) is known only from a single locality in central Pakistan (northeastern Baluchistan province). The Paghman stream salamander (Afghanodon mustersi) is known only from the three glacier-fed, high-elevation tributaries of the Paghman stream drainage in the Hindu Kush of eastern Afghanistan (Kabul province). The total population is thought to be roughly 1000–2000.

The Marrah Mountains The Marrah Mountains (Jebbel Marah in Arabic) are a range of volcanic peaks within a massif located in south-western Sudan.

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Lowe’s gerbil (Gerbillus lowei) and the Darfur gerbil (G. muriculus) are both known only from the Marrah Mountains. The arid woodland thicket rat (Grammomys aridulus) is known only from the Marrah Mountains.

The Nur Mountains The Nur Mountains (Nur Daglari in Turkish, also known as the Amanos Mountains) are located in coastal south-central Turkey (Hatay province). Baran’s kukri snake (Muhtarophis barani) is confined to the Nur Mountains.

Lowland Mediterranean Forests, Woodlands, and Scrub Within the Saharo-Arabian Region Mediterranean forests occur primarily along parts of the North African coast from Morocco to Tunisia, with isolated pockets in Libya, Egypt, and coastal areas of the Levant as well as on the Canary Islands and Cyprus. The vegetation in these areas is, in principle, much like that of the Mediterranean shores of southern Europe except that it has suffered even more from man’s destructive activities. The golden hamster (Mesocricetus auratus), familiar the world over today as a pet, was originally described from a single specimen collected near Aleppo, Syria in 1839 by a British zoologist. It was not recorded again until 1930, when a mother and her young were captured alive. These were bred in Jerusalem as laboratory animals, with some escaping and forming the wild colony we see there today. Descendants of these captive animals were also shipped to Britain beginning in the early 1930s and became the nucleus of all the golden hamsters seen up until the 1970s, when a new stock was finally exported from Syria. In terms of the natural population, it appears to be entirely confined to the Aleppinian Plateau of north-western Syria and south-central Turkey, where it is seriously threatened by loss of habitat and moreover considered to be an agricultural pest. Ebner’s cylindrical skink (Chalcides ebneri) is known only from two localities in northern Morocco, where it was last recorded in 1970. Günther’s cylindrical skink (C. guentheri) is confined to Mediterranean forest fragments in northwestern Jordan, south-western Syria, southern Lebanon, and northern and central Israel. The North African Lataste’s viper (Vipera latastei gaditana) is found sporadically in northern Morocco, northern Algeria and, at least historically, in extreme north-western Tunisia. It is heavily persecuted and has undergone significant declines. Aboubakeur’s false smooth snake (Macroprotodon abubakeri) is known from north-eastern Morocco and north-western Algeria (including the Habibas Islands).

Deserts and Semi-Deserts A large proportion of the Saharo-Arabian Region is, of course, desert and semi-desert. Many of these areas were formerly

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grasslands, rich in perennial grasses, but overgrazing by domestic animals over thousands of years has degraded them. If left in peace by man, however, they would gradually change back, so in a biological sense they are not, in reality, deserts at all. One of the largest of these man-made deserts is in the Middle East, where in the midst of seemingly endless sand and scrub great ruins bear witness to glorious civilizations once based on fertile lands. The biological capital of countries like Jordan, Syria, and Iraq has probably been continuously mistreated by human beings and their goats for longer periods than any other similar area. The vegetation and the fertile soil have disappeared, the erosion gnaws ever deeper, the springs are dried up and many vertebrate species have been exterminated. And every year the mismanagement accelerates. The Persian wild ass (Equus hemionus onager) was historically widely distributed in the arid semi-deserts and plains of the Middle East. Hunting for meat and competition with livestock have steadily reduced it to the point where, today, the natural population of around 600 is confined to two protected areas in north-eastern Iran. A captive breeding programme has been successful, and has resulted in the introduction of this species both to areas of its former range as well as locations once inhabited by the now-extinct Syrian wild ass (E. h. hemippus) in Saudi Arabia, Israel, and Ukraine. During the Middle Ages the scimitar-horned oryx (Oryx dammah) was probably the most numerous larger mammal of the semi-deserts just south of the Sahara, occurring from Mauritania in the west to the Red Sea in the east. It was relentlessly hunted, however, at first by Arab horsemen and later by oil prospectors and soldiers (often using motorized vehicles and automatic weapons). It remained relatively common for a time, with a single herd of 10,000 having been reported in Chad in 1936. By the mid-twentieth century, though, it had been drastically reduced in both range and numbers, and within a few more decades completely wiped out. The species was last reported in Chad in the early 1990s, and was thereafter believed to be extinct in the wild. Small captive populations continue to exist in fenced protected areas in Tunisia, Morocco, and Senegal. As of 2016 efforts were underway to release a large number of oryxes into the Oudi Rimé-Ouadi Achim Faunal Reserve in Chad. The addax (Addax nasomaculatus) is another large, nomadic, desert-adapted antelope historically found throughout almost the whole of the Sahara Desert and Sahelian region west of the Nile Valley, but which has undergone a catastrophic decline over the last century. By the 1970s it was long extinct in North Africa but still present in parts of Western Sahara, Mauritania, northern Mali, Chad, Niger, southern Algeria, southern Libya, and possibly northern Sudan. It was considered very rare everywhere except for an uninhabited and almost waterless area of Mauritania and Mali, where it was still numerous despite being constantly hunted by nomads from motorized vehicles. This intense hunting pressure has continued unabated, and has brought the species to the brink of extinction. Today, the only known remaining population

The Saharo-Arabian Region

thought to be viable lives in the Termit/Tin Toumma region of south-eastern Niger. However, there are sporadic reports of small, isolated groups and individuals from the eastern Aïr Mountains and western Ténéré region of north-eastern and north-central Niger, and from the Djourab region of western Chad. Possible rare vagrants from these areas may be seen elsewhere in northern Niger as well as in southern Algeria and Libya. There have been rumours of addax along the Mali/Mauritania border, and in 2007 fresh tracks from a small herd were observed in central Mauritania. Addax have rarely been reported since. In 2013 it was estimated that probably fewer than 300 still survived in the wild, ranging unevenly along a narrow, 600-km-long band lying between Termit/Tin Toumma and the Djourab sand sea. By 2016 that number had been reduced to less than a hundred, and perhaps considerably less, due to recent poaching. Fortunately, there is a considerable worldwide captive population of around 750, in addition to several thousand on private hunting ranches in the United States and the Middle East. Small numbers have been released as well into large, fenced enclosures within protected areas in Tunisia, Morocco, and Mauritania. The Arabian leopard (Panthera pardus nimr), the smallest of the leopards, still lives in arid areas of the Arabian Peninsula north to Syria, Israel, and Lebanon. Populations from the Sinai Peninsula are now believed to be extirpated. The Arabian grey wolf (Canis lupus arabs) was once found throughout the Arabian Peninsula, but is now confined to small pockets in southern Israel, southern and western Iraq, Oman, Yemen, Jordan, Saudi Arabia, and the Sinai Peninsula of Egypt. It is rare throughout its range and heavily persecuted by humans. The Laristan mouflon (Ovis gmelini laristanica) is confined to a few desert reserves in southern Iran. Hoogstraal’s gerbil (Gerbillus hoogstraali) is confined to a small area of central coastal Morocco between the High Atlas and Anti-Atlas Mountains, where it is threatened by habitat destruction and human disturbance. The Khartoum gerbil (G. stigmonyx) is known only from east-central Sudan. The Dhofar shrew (Crocidura dhofarensis) is known only from two shrubby localities in Oman and Yemen. Katinka’s shrew (C. katinka) is a little-known species sporadically recorded from Israel (where it appears to have been extirpated), Palestine, Syria, and south-western Iran. The Khuzistan shrew (C. susiana) is confined to a small area of south-western Iran (Khuzistan province). The Kerman vole (Microtus kermanensis) is known only from a few specimens collected in south-eastern Iran. Christie’s long-eared bat (Plecotus christii) is known only from a few localities in north-eastern Africa, including Egypt, eastern Libya, and Sudan. It was described as ‘very common’ in Palestine during the late nineteenth century, but appears to have been extirpated from there. Two subspecies of common ostrich (Struthio camelus) have been heavily impacted by hunting and overcollection of their eggs. The Arabian ostrich (S. c. syriacus) was historically

widespread in the arid steppes of the Middle East. Human persecution is believed to have driven it extinct around 1966. The North African ostrich (S. c. camelus), the world’s largest living bird, was once similarly widespread across much of North Africa, but has disappeared from large areas of its former range. Kleinmann’s tortoise (Testudo kleinmanni) was historically widespread along the Mediterranean coast of North Africa (Libya and Egypt), the Sinai Peninsula, and Israel, but is now confined to only a few areas, where it is facing extinction in the wild due to loss of habitat and collection for the international pet trade. Peters’ rock gecko (Pristurus longipes) is known only from historical records originating in Yemen, as well as from a poorly preserved specimen collected from what is now Eritrea. The keel-scaled bent-toed gecko (Mediodactylus aspratilis) is known only from two localities in south-western Iran (Fars province). The Jaz Murian bent-toed gecko (M. sagittifer) is known only from the Jaz Murian Depression of south-eastern Iran (Sistan and Baluchistan provinces). The banded-toed gecko (Saurodactylus fasciatus) is confined to a few scattered localities in north-western Morocco. Werner’s spider gecko (Cyrtopodion gastrophole) is known only from two localities in south-western Iran (Fars province). The Harran spiny-toed lizard (Acanthodactylus harranensis) is known only from the ruins of the ancient city of Harran in south-eastern Turkey, but may occur in adjacent areas of Syria. The south Arabian spiny-toed lizard (A. felicis) is found disjunctly in southern Yemen and southern Oman, where it is threatened by coastal development. Nilson’s spiny-toed lizard (A. nilsoni) is known only from a single, now-degraded locality in western Iran (Kermanshah province). The Red Sea snake-eyed lizard (Ophisops elbaensis) is known only from a few isolated localities in south-eastern Egypt and the south-western Arabian Peninsula (southwestern Saudi Arabia and western Yemen). Werner’s diadem snake (Spalerosophis dolichospilus) is known only from a few records from Algeria, Morocco, and Tunisia. Thomas’ racer (Coluber thomasi) is confined to the Dofar region of southern coastal Oman. The Halfeti blind snake (Letheobia episcopus) is known only from two localities in the Euphrates River Valley of southeastern Turkey. The Iranian worm snake (Xerotyphlops wilsoni) is known only from a single locality in south-western Iran.

The Sahara Desert North Africa is dominated by the Sahara, the largest hot desert in the world. It extends from the Atlantic to the Red Sea and down to the highlands of Ethiopia, but it is not everywhere flat, sandy, and lifeless. Some parts of it do get a little rain, which occasionally or periodically encourages sparse plants to grow. There are mountains at its heart such as those at Tibesti that climb to a height of 3445 m, or extend in a chain, as do the

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cedar-clad and snow-capped Atlas Mountains in the northwestern corner. In the east the mighty Nile creates a strip of vegetation amid the sand. Today, although species richness and endemism in the Sahara are very low, some highly adapted creatures do survive, but as a whole it is barren country, a sea of sand with a few oases. It was not always so. During the last ice age in northern Europe and Asia the climate was significantly wetter in the Sahara, and it was not until geologically recent times that the desert became dominant. There is also evidence that in a nottoo-distant past – 6000–2000 years ago – parts of the Sahara consisted of steppes with perennial grasses supporting an abundance of mammals of a kind found in the African tropics to the south. It is very likely, however, that for long periods there was a desert-like belt, perhaps somewhere in the middle of the present Sahara, but apparently it was not large enough to prevent the dispersal of tropical animals from the tropical savannas. Between 4500 and 3500 B . C ., hippopotamuses, elephants, giraffes, and rhinoceroses were common on the central parts of the Sahara, showing that there was ample grass, shrub, and tree vegetation to satisfy their enormous needs. Hippopotamuses wallowed in the rivers and crocodiles lay along the banks, while buffaloes and several species of antelope lived on the wide expanses of grass. At least part of the Sahara was thus grassy tree savanna in relatively recent times. Around 2000 B . C . the hippopotamuses, elephants, and rhinoceroses began to disappear from the central parts of the Sahara, but as late as 1200 B . C . giraffes were still nibbling the leaves in the crowns of trees. In 1815 there were hippopotamuses in Egypt, and in 1750 both they and the elephant were still found about the lower Senegal River. The drying up of the Sahara at the end of the Pleistocene certainly contributed to the disappearance of many of these forms of wildlife. But man, too, is involved. We know from cave paintings in the western Sahara that herds of domestic stock once lived here along with large grazing and browsing animals. The abuse of the soil and vegetation by domestic livestock through millennia has ruined North Africa and changed to deserts even areas with a moderate rainfall. In the interior of the Sahara scientists have found fossilized profiles of humus-rich soil with pollen, calculated to be 5000–6000 years old, from a cedar forest. As recently as 2000 years ago, there were forests deep in the Sahara where today there is nothing but sand, indicating that the vegetation along the coasts of North Africa once extended into what is now desert. In Morocco alone, forests covering more than one-third of the country have disappeared since Roman times. What was the cause of the widespread desertification of such a vast area? No conclusive answer can be given, but there is good reason to believe that man and his livestock were an accelerating factor in the process. It is a remarkable fact that whenever civilization in North Africa has flourished, the soil has begun to bear plants – before and during the Roman period, and in more recent times under French and Italian control – but during the long periods when the area was dominated by nomadic tribes,

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the soil deteriorated. The climate is, of course, a contributing factor, but it seems to be man, cows, and goats who began the erosion avalanche. The most degradation is found where there is water (oases, etc.). Here, habitats may be heavily altered by human activities. Previously existing tree cover has often been removed for fuel and fodder by nomadic pastoralists and traders. Unfortunately, not all the larger mammals of North Africa could take refuge in the tropics. Many could not escape or were not adapted to survive outside the subtropical region. Persecution by humans left them in isolated pockets, chiefly in mountainous zones. There they were soon reduced in numbers and some were exterminated. Hunting in general was responsible for this decline, but it was only after the use of firearms spread to most tribes that several species were completely wiped out. The African wild dog (Lycaon pictus) was historically distributed throughout sub-Saharan Africa outside the rainforest belt, where it has been divided into a number of subspecies. All have now been reduced to scattered populations that are highly threatened by loss of habitat, human persecution, and infectious disease. The Chadian wild dog (L. p. sharicus) is confined to disjunct areas of south-eastern Algeria and central and southern Chad. The Egyptian aoudad (Ammotragus lervia ornata) is a type of wild sheep that was historically widespread throughout the deserts of Egypt, where it was heavily hunted. Thought to be extinct, evidence for its survival was discovered during the late 1990s within a small area of the Eastern Desert. Fassin’s aoudad (A. l. fassini) is confined to extreme southern Tunisia and adjacent north-western Libya. The dorcas gazelle (Gazella dorcas) historically occurred over virtually all of North Africa from the Mediterranean to the southern Sahel and from the Atlantic to the Red Sea, extending into southern Israel, Syria, Jordan, and the Horn of Africa. By 1960 it was declining at an alarming rate mainly as a result of motorized hunting. Today it has been much reduced and even eliminated in many parts of its former range. Several ill-defined subspecies are sometimes recognized, all of which are considered vulnerable. They include the Egyptian dorcas gazelle (G. d. dorcas), Eritrean dorcas gazelle (G. d. beccarii), Isabelle dorcas gazelle (G. d. isabella), Moroccan dorcas gazelle (G. d. massaesyla), Algerian dorcas gazelle (G. d. osiris), and Pelzeln’s dorcas gazelle (G. d. pelzelnii). The slender-horned or rhim gazelle (G. leptoceros) was once the most common of all the gazelles living in the Sahara Desert. Today, owing to decades of uncontrolled motorized hunting by Arab tribesmen, at most only a few hundred still survive in isolated areas of Algeria, Tunisia, Libya, and Egypt west of the Nile. There are additional, unsupported reports from Chad and Niger, although it is unclear if the species in fact ever occurred on the south side of the Sahara. The dama gazelle (Nanger dama) has become seriously threatened owing to ruthless hunting for their bezoars, which are used in ‘traditional medicine’. The nominate form (N. d. dama) was formerly widespread in both the Sahara and

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Sahel, but has become very rare in the wild. A captive population lives in the Al Ain Zoo, United Arab Emirates. The addra gazelle (N. d. ruficollis) is very rare in the wild but present in captive breeding programmes in Europe, North America, and the Middle East. The mhorr gazelle (N. d. mhorr) historically lived in south-western Morocco and in the north-western Sahara. The last known wild sighting was in 1968, and by 1971 only 12 were known to exist. Fortunately, it still occurs in captive breeding programmes in Europe, North America, North Africa, and the Middle East, and efforts are underway to reintroduce populations to protected areas in the wild. A possible fourth subspecies, the Rio de Oro gazelle (N. d. lozanoi), remains a complete mystery. It was thought that not more than 50 were still in existence in 1968, but there is no information available on its current status. It is worth noting that recent studies have called into question the validity of any dama gazelle subspecies, and suggests that the differences in colour are merely the result of a geographic cline. The African houbara bustard (Chlamydotis undulata) still has a wide range across North Africa, occurring in northernmost Mauritania, Western Sahara, Morocco, Algeria, Tunisia, Libya, and Egypt west of the Nile. There are old records from Sudan as well. It is mainly threatened by hunting. Zoli’s shield-backed lizard (Philochortus zolii) is known only from five, widely scattered semi-desert localities near oases in south-eastern Algeria, north-eastern Egypt, central Mali, north-western Niger, and northern coastal Libya. The Taghit spiny-toed lizard (Acanthodactylus taghitensis) is known only from a few specimens collected in central and northern Mauritania and western Algeria. It possibly occurs in Western Sahara and other areas as well. Doumergue’s spinytoed lizard (A. spinicauda) is confined to the Arba Tahtani and El Abiod-Sidi-Cheikh oases of north-western Algeria. The Mauritanian rock gecko (Pristurus adrarensis) is known only from three specimens collected from two localities in central Mauritania. Atlantic Coastal Desert The Atlantic Coastal Desert lies along the western coast of the Sahara, occupying a narrow strip in Western Sahara and Mauritania. The Tarfaya shrew (Crocidura tarfayensis) is confined to southern Morocco and northern Mauritania. Surveys undertaken since 1980 have failed to find a live specimen, although dead ones have been obtained from owl pellets. The Eastern Desert The Eastern or Nubian Desert covers eastern Egypt and northeastern Sudan between the Nile River and the Red Sea. Heuglin’s gazelle (Eudorcas tilonura) occurs patchily east of the Nile between the southern part of the Red Sea Hills in Sudan and the southern foothills of the Ethiopian highlands in western Eritrea and north-western Ethiopia. The total population is thought to be around 2500–3500.

The Egyptian pygmy shrew (Crocidura religiosa) is known only from two specimens, one collected from the neck of the Nile delta and the other from further down the Nile. A few mummified examples are also known from ancient tombs at Thebes. The Nile Valley The Nile Valley is a lush, fluvial area located on both banks of the Nile River in Egypt and Sudan, adjacent to desert. It is characterized by low, rocky hills, and shallow, sandy wadis. Steudner’s sand gecko (Tropiocolotes nubicus) is known from a few localities in the southern Nile Valley.

The Arabian Desert The Arabian Desert encompasses a vast area of western Asia from Yemen to the Persian Gulf and Oman to Jordan and Iraq, including most of the Arabian Peninsula. The environment is everywhere hostile, no more so than in the Rub’alKhali (‘Empty Quarter’), one of the largest continuous bodies of sand in the world. Nevertheless, a number of interesting species managed to survive quite well there, only to be systematically wiped out by humans over the last century or so. If we consider only the larger animals, the inventory of devastation is horrifying. Lions, cheetahs, bears, and deer are all long gone, but as late as 1917 various species of gazelle as well as bustards (Chlamydotis macqueenii) could still be seen almost everywhere, and there were also many Arabian oryx (Oryx leucoryx) and ostrich (Struthio camelus syriacus). The advent of modern firearms, however, resulted in mass slaughter on a massive scale. Gazelles were even machine-gunned by fighter aircraft in several countries in the Middle East. Such treatment is arguably even more deplorable than the slaughter of bison on the American prairies and of various ungulates on the South African veld during the past centuries. The Arabian oryx (Oryx leucoryx) is a medium-sized antelope that historically occurred throughout most of the Arabian Peninsula, the Sinai, and the Levant north to Kuwait and Iraq. Ruthlessly hunted both by Bedouin tribesmen using modern firearms as well as by motorized hunting parties, the species’ range had already begun to contract by the early years of the twentieth century, with the decline accelerating thereafter. By 1920 the population was separated into areas over 1000 km apart; a northern one in and around the Nafud, and a larger, southern one in the Rub al Khali and the plains of centralsouthern Oman. The northern population was extirpated in the 1950s while, in the south, the range steadily decreased. By the 1960s oryx were restricted to parts of central and southern Oman, where they were continued to be hunted nevertheless. In 1960 and 1961 two parties of poaching sheiks in powerful cars, armed with submachine guns or automatic rifles, slaughtered 60 or 70 of the animals. By 1962, only 11 oryx were believed to remain in Saudi Arabia. The last wild individuals were most likely shot in 1972 on the Jiddat al Harasis.

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Fortunately, the so-called Operation Oryx program, whereby captive oryx were brought together to the Phoenix Zoo, Arizona, in order to create a breeding nucleus, proved to be highly successful. From this, reintroductions to the wild were made first to protected areas in Oman (from 1982), Saudi Arabia (from 1990), Israel (from 1997) the United Arab Emirates (from 2007), and Jordan (from 2014). There is an additional small, introduced population living on Hawar Island, Bahrain, along with larger, semi-managed ones at several sites in Qatar, the United Arab Emirates, and Saudi Arabia. The Saudi gazelle (Gazella saudiya) was historically found in western Saudi Arabia, Kuwait, and Yemen. Heavily hunted, it is thought to have gone extinct in the wild by the 1980s. Purported surviving captive populations have all proven to be either different species or hybrids. Two other species of gazelle from the Arabian Desert region are threatened. The Arabian sand gazelle (G. marica) is found in the Arabian Peninsula and on a few Persian Gulf Islands, where the total population is estimated at less than 10,000. The Arabian gazelle (G. arabica) is found sporadically on the Arabian Peninsula from Israel through coastal areas of Saudi Arabia (including the Farasan Islands), Yemen, Oman, and the United Arab Emirates. There is also a small, introduced population on Farur Island, Iran. The last records from Sinai are from 1932.

Jayakar’s agama (Trapelus jayakari) is a type of lizard known only from a few isolated historical records scattered across the south-eastern Arabian Peninsula (Oman, Bahrain, Yemen, United Arab Emirates, and south-eastern Saudi Arabia). The Omani spiny-tailed lizard (Uromastyx thomasi) was historically known only from a few localities in Oman, where it has not been recorded from the mainland in several decades. The species still survives on Masirah Island, where it is threatened by loss of habitat. The small-scaled spiny-tailed lizard (U. aegyptica microlepis) and Leptien’s spiny-tailed lizard (U. a. leptieni) are confined to northern Oman and parts of the United Arab Emirates and Saudi Arabia. The Masirah spiny-toed lizard (Acanthodactylus masirae) is known only from a few isolated localities in Oman, including Masirah Island. The Mazbah rock gecko (Pristurus mazbah) is known only from a single locality in western Yemen. Wolfgang Boehme’s sand gecko (Tropiocolotes wolfgangboehmei) is known only from two specimens collected from central Saudi Arabia. The Ayun sand lizard (Mesalina ayunensis) is known only from a small area of south-western Oman. Leviton’s cylindrical skink (Chalcides levitoni) is known only from a single locality in south-western Saudi Arabia (Jizan region).

Figure 2.15 Saudi gazelle (Gazella saudiya). (Credit: R. P. Lesson.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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The Arabian worm lizard (Agamodon arabicus) is known only from a single specimen collected in southern Yemen in 1901. The Aden kukri snake (Rhynchocalamus arabicus) is known only from a single specimen collected at an imprecise locality near Aden (Yemen) in 1932. The Yemeni blind snake (Myriopholis yemenicus) is known only from a single specimen collected in the 1930s from an undefined locality in Yemen.

The Levantine Desert The Levantine Desert, as here defined, includes all the lowland desert and semi-desert regions of Egypt’s Sinai Peninsula, Israel, Palestine, and Jordan. Buxton’s jird (Meriones sacramenti) is a type of rodent confined to the coastal dunes and desert areas of northern Egypt, northern Israel, and Palestine, where it is threatened by habitat destruction. Savigny’s agama (Trapelus savignii) is a type of lizard confined to the western Negev sands of Israel and the Gaza Strip west across the northern Sinai to the eastern margins of the Nile delta. The Egyptian populations are highly fragmented and have almost been extirpated, as are those historically found south of Tel Aviv. The Jordanian spiny-toed lizard (Acanthodactylus ahmaddisii) is confined to a small area of central Jordan. The Sinai cat snake (Telescopus hoogstraali) is known only from four scattered localities, two in the Sinai Peninsula, another in the northern Negev Desert of central Israel, and the Petra area of south-western Jordan. The Negev Desert The Negev is a rocky desert that covers more than half of Israel. The Be’er Sheva spiny-toed lizard (Acanthodactylus beershebensis) is confined to a few localities in south-central Israel.

The Syrian Desert The Syrian Desert (Badiyat al-Sham in Arabic) is an area of desert, semi-desert, and steppe located in the Middle East between the Orontes and the Euphrates rivers. It includes parts of south-eastern Syria, north-eastern Jordan, northern Saudi Arabia, and western Iraq. It is typically open and rocky, cut with occasional wadis (valleys or dry riverbeds). The Syrian wild ass (Equus hemionus hemippus), with a shoulder height of just 1 m, was the smallest living horse. It lived in the deserts, semi-deserts, and arid grasslands of Palestine, Israel, Jordan, Turkey, Syria, Saudi Arabia, western Iran, and Iraq. European travellers to the Middle East during the fifteenth and sixteenth centuries reported seeing large herds. However, its numbers began to drop precipitously during the eighteenth and nineteenth centuries due to overhunting, and its existence was further imperilled by the upheavals of the First World War. The last known wild individual was shot in 1927 at al Ghams, near the Azraq Oasis in

Jordan, in 1927, and the last captive specimen died the same year in the Vienna Zoo, Austria. Tsolov’s mouse-like hamster (Calomyscus tsolovi) is known only from a small number of specimens said to have been collected in south-western Syria.

The Dasht-e Kavir Desert The Dasht-e Kavir (also known as the Great Salt Desert) is a large desert located in the middle of the Iranian Plateau in eastcentral Iran (Khorasan, Semnan, Tehran, Isfahan, and Yazd provinces). The Asiatic cheetah (Acinonyx jubatus venaticus) was historically widespread across south-western and central Asia as far as India, but was exterminated by persecution and destruction of its habitat. It was last seen in the Arabian Peninsula in 1950 and in India in 1968, and perhaps held on in Turkmenistan, Afghanistan and Pakistan into the 1970s. Today the subspecies is only known to occur in protected areas of the Dasht-e-Kavir in Iran, a country where it had also very nearly been exterminated but was saved at the last moment by conservation measures. The total population there is believed to be between 60 and 100.

Miscellaneous Deserts The Dasht-e Lut is a large salt desert located in south-eastern Iran (Kerman and Sistan/Baluchestan provinces). Thaler’s jerboa (Jaculus thaleri) is known only from two localities in the Dasht-e-Lut Desert. The Kharan Desert is a sandy and mountainous area located in south-western Pakistan (Balochistan province). The Kharan pygmy jerboa (Salpingotulus michaelis) is known only from the Kharan Desert.

Isolated Caves, Springs, and Pools A large number of species within the Saharo-Arabian Region are confined to caves, springs, and pools. In desert areas wadis – dry (ephemeral) riverbeds that contain water only after heavy rains – are of particular importance to freshwater fishes. The Dhofar house gecko (Hemidactylus lemurinus) is confined to a few areas of southern Yemen and Oman, where it inhabits the bottoms of wadis. The Wadi al Khalili toad (Duttaphrynus scorteccii) appears to be confined to Wadi al Khalili, located on a high plateau near Mafhaq in western Yemen. The Mamilla Pool tree frog (Hyla heinzsteinitzi) is a species of uncertain validity known only from Mamilla Pool, an ancient reservoir located outside the walls of the Old City of Jerusalem, Israel. It possibly went extinct in the late 1990s. Desfontain’s cichlid (Astatotilapia desfontainii) is confined to freshwater springs, irrigated lands, canals, and ditches in Algeria and Tunisia. It is threatened by habitat destruction. The Hadiyah bream (Acanthobrama hadiyahensis) is confined to the Wadi Hadiyah basin of Saudi Arabia.

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The Ksob barbel (Barbus ksibi) is confined to Wadi Ksob in central Morocco. The Tunisian barbel (Luciobarbus antinorii) is known from artesian wells at Chott el Djerid in southern Tunisia, where it was last recorded in 1989. It may already be extinct, a victim of drought. The Sahara toothcarp (Aphanius saourensis) was historically found throughout the Oued Saoura basin in west-central Algeria, but is now reduced to a single remnant population. The Azraq toothcarp (A. sirhani) is confined to the Azraq Oasis in Jordan, where water levels are being artificially maintained to counter illegal extraction. A number of freshwater fish of the genus Garra are confined to small, isolated localities. The Tawi Atair garra (G. dunsirei) is known only from a single sinkhole in the Dhofar region of Oman. The Lorestan garra (G. lorestanensis) is confined to Loven Cave in south-western Iran (Lorestan). The Tashan garra (G. tashanensis) is confined to Tashan Cave in western Iran (Khuzestan province). The Jebel al Akhdar garra (G. longipinnis) is known only from a single wadi in Oman, where it was last reported in 1968. The Wadi Hasik garra (G. smarti) is known only from Wadi Hasik in Oman. Proudlove’s blind cave loach (Eidinemacheilus proudlovei) is a little-known species from the Kurdistan region of northern Iraq.

The Haditha Aquifer System The Haditha aquifer system is a series of limestone karst caves located near Haditha in western Iraq. The Haditha cave garra (Typhlogarra widdowsoni) is type of blind barb also confined the Haditha aquifer system, where it is accessible through two isolated wells. It is extremely rare. The Haditha cavefish (Caecocypris basimi) is a poorly known species confined to the Haditha aquifer system, where it is accessible only through a single well. Not recorded since 1983, it may already be extinct due to water extraction.

Wadi Hadhramaut Wadi Hadhramaut is a large wadi located in Yemen. The Hadhramaut himri (Carasobarbus exulatus) is a type of barb confined to Wadi Hadhramaut. A single record from Wadi Maran appears to be erroneous. The smooth garra (Garra lautior) and the spiny garra (G. mamshuqa) are both confined to the Wadi Hadhramaut drainage.

Lakes, Rivers, and Marshes While notably dry, the Saharo-Arabian Region does have a few larger lakes along with the Tigris/lower Euphrates, lower Nile, lower Indus, Orontes, and Jordan river drainages. Pollution and loss of habitat due to drought and water extraction are a constant threat throughout. The Algerian ribbed newt (Pleurodeles nebulosus) is found patchily in northern Algeria and western Tunisia, where it is threatened by loss of habitat.

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Members of the genus Tristramella are among the few cichlids to be found in western Asia. The shortjaw tristramella (T. simonis) is confined to the Jordan River drainage, including Lake Tiberias, in Israel and Syria. Introduced populations also occur in the Nahr al-Kabir and Orontes drainages of Syria. Two populations, variously described either as full species, subspecies (as here) or synonyms have already gone extinct. The Damascus shortjaw tristramella (T. s. magdelainae) is known only from the vicinity of Damascus, in Syria, where it went extinct at some unknown point after its description in the late nineteenth century. The Syrian spotted bleak (Alburnus qalilus) is confined to parts of three small coastal streams flowing into the Mediterranean, as well as to the reservoir at Nahr al Kabir. The Moroccan barbel (Luciobarbus issenensis) is confined to the Sous and Massa river drainages of north-western Morocco. The Algerian barbel (L. leptopogon) is known only from its original collection in 1834. Hasan’s chub (Squalius spurius) is confined to the Orontes and Nahr al-Kabir river drainages of south-eastern Turkey and northern Syria. The Marqiyah spring minnow (Pseudophoxinus hasani) is confined to Nahr Marqīyah stream in coastal Syria. The Damascus bream (Acanthobrama tricolor) is confined to southern Syria, where it was last recorded from the Barada River in 1908 and from two specimens collected from the Masil al Fawwar River in the late 1980s. Both rivers have largely dried out due to drought, and what remains is heavily polluted. It is most likely extinct, but may survive in the Golan Heights. The Yarkon bream (A. telavivensis) was historically found in the coastal streams of Israel where, in 1999, it became extinct in the wild due to drought. It has since been reintroduced in a number of rehabilitated localities and artificial ponds within the Yarkon and Tut river drainages. The Dead Sea toothcarp (Aphanius richardsoni) was historically found in small springs and streams surrounding the Dead Sea in Israel and Jordan, but has been extirpated from all but three areas due to water extraction. The Moroccan spined loach (Cobitis maroccana) is confined to small tributaries of two rivers (the Loukkos and Sebou) on the Atlantic coast of northern Morocco. Namir’s stone loach (Barbatula namiri) occurs in the Orontes River watershed and a few other coastal rivers in Lebanon, western Syria, and southern Turkey. The Damascus sportive loach (Oxynoemacheilus panthera) is confined to the Barada and Awaj rivers in Syria.

The Dead Sea The Dead Sea is a large, deep, hypersaline lake bordered by Jordan to the east and by Israel and the West Bank to the west. At 430.5 m below sea level, its surface and shores represent the Earth’s lowest elevation by land. The Dead Sea garra (Garra ghorensis) is confined to springs and a single wadi in the southern Dead Sea basin. It

The Saharo-Arabian Region

is now confined to Jordan; the Israel population having been extirpated.

Lake Tiberias Historically known as the Sea of Galilee, Lake Tiberias (Kinneret Lake) is a large lake in north-eastern Israel. It is the lowest-elevation freshwater lake in the world. The longjaw tristramella (Tristramella sacra) was endemic to Lake Tiberias. It has not been seen since 1989/1990, either in the lake or in local markets, and is now considered extinct. It is thought that destruction of marshes, the breeding area for this species, may have been the reason for the extinction.

The Oum Erbiah River The Oum Erbiah River is located in central Morocco. Hartert’s himri (Carasobarbus harterti) is confined to the Oum Erbiah drainage. The Oum Erbiah yellowfish (Pterocapoeta maroccana) is known only from a single specimen collected from the Oum Erbiah River at the beginning of the twentieth century.

The Lower Tigris/Lower Euphrates River Drainage The Tigris and the Euphrates, the two great rivers that define Mesopotamia (‘the land between the rivers’), are located in south-eastern Turkey, Syria, Iran, and Iraq. The two eventually join in the Shatt al-Arab, which empties into the Persian Gulf. The Euphrates softshell turtle (Rafetus euphraticus) is still found throughout much of the Tigris and Euphrates rivers and their tributaries, lakes, ponds, and marshlands in south-eastern Turkey (Anatolia), Syria, Iraq, and south-western Iran (Khuzestan province). Habitat destruction and alteration, and in particular dam construction, are the primary threats. The shabout (Arabibarbus grypus) is a large freshwater carp found throughout the Euphrates and Tigris river drainages, where it is threatened mainly by overfishing. The leopard barbel (Luciobarbus subquincunciatus), yellowfin barbel (L. xanthopterus), and the pike barbel (L. esocinus) were all historically common within the Tigris and Euphrates drainages, but are now seriously threatened by overfishing, pollution, and dams. The binni (Mesopotamichthys sharpeyi) is a type of barbel historically found throughout the lower Euphrates and Tigris drainage. Owing to overfishing and loss of habitat it has been extirpated from most of its former range, and now appears to be confined to a few marshy areas in southern Iraq and Iran, and in Lake Assad, Syria. The kiss-lip himri (Carasobarbus kosswigi) is known only from a few widely separated localities within the Tigris/ Euphrates drainage. The Mesopotamian Marshes The Mesopotamian marshes are a historically extensive wetland area located within the Tigris–Euphrates delta of southern Iraq and south-western Iran. Beginning in the 1950s there was a massive loss of this biologically important habitat due to

large-scale hydrological projects throughout the Euphrates and Tigris river drainages, as well as by deliberate draining to reclaim land for agriculture and oil exploration. The Iran– Iraq War of the 1980s resulted in particularly extensive damage to the reedbeds in southern Iraq. By 2003 this rare aquatic landscape within a desert had been reduced to just 10 per cent of its former size. Since the end of the Iraq War the marshes have been partially re-flooded, although drought and further dam construction upstream in Turkey, Syria, and Iran have hindering recovery. In 2016 they were declared a UNESCO Heritage Site. The Iraq smooth-coated otter (Lutrogale perspicillata maxwelli) is an isolated population confined to the Mesopotamian marshes. Bunn’s short-tailed bandicoot rat (Nesokia bunnii) is confined to the marshes of south-eastern Iraq.

The Lower Indus River The lower Indus River runs the length of western and central Pakistan. Prior to emptying into the Arabian Sea it creates a wide delta of swamps, streams, and mangroves that provide important habitat for migrating waterfowl. The South Asian river dolphin (Platanista gangetica) is divided into two subspecies. The Indus River dolphin (P. g. minor) is one of the world’s rarest mammals and most endangered cetaceans. Only about 1000 of this unique subspecies exist today within the lower reaches of the Indus River between the Jinnah and Kotri barrages, and in a few connecting channels. Historically it extended all the way from the Indus delta through a number of tributaries all the way to the Himalayan foothills, but declined after the late nineteenth century due to dam construction.

The Orontes River The Orontes (Asi in Arabic and Turkish) is a northwardflowing river which begins in Lebanon and continues through western Syria and south-eastern Turkey before finally entering the Mediterranean. The long-spine bream (Acanthobrama centisquama) was historically known from Lake Amik in south-eastern Turkey and from Lake Al-Gab in Syria, within the Orontes River drainage. Lake Amik was drained in the 1940s. The species is most likely extinct but may survive in Lake Golbasi, a small relict of Lake Amik, which has been heavily impacted by pollution and water extraction. Lortet’s barbel (Luciobarbus lorteti) is confined to the lower Orontes River in Syria and Turkey. In 1989 much of its habitat was poisoned and dried out, resulting in the species’ extirpation from Lake Amik and presumably other areas. The Orontes scraper (Capoeta barroisi) is confined to a few areas of the Orontes River drainage in Turkey and Syria. The Levantine nase (Chondrostoma kinzelbachi) was historically widespread in the Orontes River drainage of Syria and Turkey. It has been extirpated from Syria, and in Turkey is likely confined to a couple of localities.

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The Orontes spotted bleak (Alburnus orontis) is confined to the Orontes River drainage in south-eastern Turkey and western Syria. The Orontes spined loach (Cobitis levantina) is confined to the Orontes River drainage. The Orontes sportive loach (Oxynoemacheilus hamwii) was historically known from the Orontes River drainage of southeastern Turkey and northern Syria. Now extirpated from the latter country, it survives only in three Turkish streams. The Litani River The Litani River (Nahr al-Litani in Arabic) is located in central and southern Lebanon. The longest and largest river in that country, it arises in the Beqaa Valley west of Baalbek and empties into the Mediterranean Sea north of Tyre. The Ammiq garra (Garra festai) is confined to the Ammiq Marshes, where it is threatened by water extraction.

The Jordan River Drainage The Jordan River (Nahar ha-Yarden in Hebrew/Narh alUrdunn in Arabic) flows roughly north to south through Lake Tiberias and on to the Dead Sea. It is bordered by Jordan and Syria to the east, while the West Bank and Israel lie to the west. The Jordan cichlid (Haplochromis flaviijosephi) is confined to the Jordan River drainage where it occurs in Lake Muzairib and in some spring lakes in southern Syria, as well as in the vicinity of Lake Kinneret and the Beit She’an Valley of northern Israel. The Jordan barbel (Luciobarbus longiceps) is confined to streams and lakes of the Jordan River watershed in Israel, Syria, and Jordan. The Drusian spring minnow (Pseudophoxinus drusensis) is confined to a small area of the northern Jordan River drainage (Golan Heights and Jebel Druse) in Israel and Syria. Large subpopulations were observed during the 1980s, but the species declined greatly thereafter due to drought and has not recovered. Three stone loaches of the genus Nemacheilus endemic to the Jordan River drainage are threatened by habitat destruction, water extraction, and pollution. Dor’s stone loach (N. dori) is confined to the Beit She’an Valley in northern Israel, where the total population is thought to be less than 250. The Golan Heights stone loach (N. pantheroides) is confined to the Golan Heights and the upper Galilee. The Jordan River stone loach (N. jordanicus) is found patchily in Israel, Jordan, and Syria. The Galilean sportive loach (Oxynoemacheilus galilaeus) was historically confined to Lake Hula in Israel and Lake Muzairib in Syria, within the Jordan River drainage. It has since been extirpated from Lake Hula. Lake Hula Located on the upper course of the Jordan River in the Hula Valley, the once large and swampy Lake Hula and its adjacent

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marshes were deliberately drained in the 1950s and are now greatly diminished. The Hula painted frog (Latonia nigriventer) was long known for certain only from two localities on the eastern shore of Lake Hula. Swamp-drainage schemes were thought to have exterminated the species in the mid-1950s, but in 2011 it was rediscovered in a very small area of the Hula Nature Reserve. The Hula bream (Mirogrex hulensis) was driven to extinction by the 1970s due to loss of habitat. The Hula shortjaw tristramella (Tristramella simonis intermedia) was extinct by the 1970s due to loss of habitat.

Miscellaneous Lakes, Rivers, and Marshes Lake Sidi Ali is located in the mountains of northern Morocco. The Sidi Ali trout (Salmo pallaryi) is known only from two museum specimens collected from the lake. It is thought to have disappeared in the 1930s due to competition with introduced carp. Lake Ifni is a high-elevation lake located in the Atlas Mountains of Morocco. The Ifni trout (Salmo akairos) is endemic to the lake. The Hub River is located in south-western Pakistan (Balochistan). The Hub torrent catfish (Amblyceps macropterus) is a little-known species confined to the Hub River drainage. The Kul River is located in Iran. The Kor sportive loach (Oxynoemacheilus tongiorgii) is known only from a single specimen collected in the Kul River drainage. The Tensift River is located in central Morocco. The Tensift yellowfish (Labeobarbus reinii) is confined to the lower courses of the Tensift River. The Sebou River (Asif en Sbu in Berber) is located in northern Morocco. The Ouerrha combtooth blenny (Salarias atlantica) is confined to the Ouerrha River in the Sebou drainage. The Medjerda River (known historically as the Bagrada River) is located in north-eastern Algeria and northern Tunisia. The Punic spring minnow (Pseudophoxinus punicus) is confined to the Medjerda River drainage. The Baradá River (Nahr Baradá in Arabic) is located in Syria. The Baradá spring minnow (Pseudophoxinus syriacus) is confined to the source of the Baradá River, where it may have been driven to extinction by water extraction. The Ismailia Channel (formerly known as the Sweet Water Channel) is located in north-eastern Egypt. Dug during the late nineteenth century as part of the construction of the Suez Canal, it is now largely dried out and heavily polluted. The Ismailia tilapia (Oreochromis ismailiaensis) is a type of cichlid known only from the Ismailia Channel.

Coasts and Satellite Islands This section includes the north-western African coast including the Canary and Madeira islands, the southern and eastern Mediterranean coasts including Cyprus, the coasts of the Red Sea and Arabian Peninsula, and the northern Arabian Sea.

The Saharo-Arabian Region

The four-toed jerboa (Allactaga tetradactyla) is a hopping rodent confined to northern coast of Egypt and Libya, where it is threatened by habitat destruction. Four species of gerbil (Gerbillus) are threatened by coastal development and human disturbance. The Essaouria gerbil (G. hesperinus) is confined to two sand dune localities north and south of the city of Essaouria, in coastal central Morocco. Grobben’s gerbil (G. grobbeni) is confined to a small area of north-eastern coastal Libya. James’ gerbil (G. jamesi) is confined to a small area of north-eastern coastal Tunisia. The occidental gerbil (G. occiduus) is confined to south-eastern coastal Morocco. Flower’s shrew (Crocidura floweri) is known only from the Nile River delta, where it may have gone extinct in the early 1960s. However, remains in owl pellets indicate that it survived at least until the 1980s. The Macaronesian pipistrelle (Pipistrellus maderensis) is a type of bat confined to the Madeira and western Canary islands, and possibly to the Azores as well. The total population is suspected to be under 1000 individuals. The Socotra cormorant (Phalacrocorax nigrogularis) occurs in two subpopulations in the Persian Gulf and the south-eastern coast of the Arabian Peninsula, where it breeds on a small number of offshore islands. Only 13 colonies are known to be active in total, and the species has been greatly affected by loss of habitat, disturbance, and oil spills. The Red Sea cliff swallow (Petrochelidon perdita) is known only from a single specimen found dead in 1984 at Sanganeb lighthouse, north-east of Port Sudan. Three species of spiny-toed lizard (Acanthodactylus) are threatened by loss of habitat and overcollection for the international pet trade. The Sidi Mechrig spiny-toed lizard (A. mechriguensis) is known only from four localities on the Sidi Mechig beaches of northern Tunisia. The leopard spiny-toed lizard (A. pardalis) is confined to northern Egypt and northeastern Libya. Schreiber’s spiny-toed lizard (A. schreiberi) is known only from a few coastal localities in Israel, Lebanon, south-eastern Turkey and Cyprus, where it is threatened by the development of tourist facilities. The helmethead gecko (Tarentola chazaliae) is confined to the few localities in the coastal lowlands of the western North African coast, from Agadir in Morocco to Cap Blanc in Mauritania. It is threatened by loss of habitat and overcollection for the international pet trade. The two-fingered cylindrical skink (Chalcides mauritanicus) and Doumergue’s cylindrical skink (C. parallelus) are both confined to the coast of north-western Algeria and north-eastern Morocco, with the latter found as well on the Chafarinas Islands. They are threatened by habitat destruction due to coastal development.

The Madeira Islands Situated in the North Atlantic west and slightly south of Portugal, the volcanic Madeira Islands (Ilhas Madeira in Portuguese) consist of Madeira and Porto Santo along with the smaller, uninhabited Desertas and Salvagens Islands.

The Madeiran wood pigeon (Columba palumbus maderensis) was already considered rare during the late nineteenth century, and was most likely extinct by 1924. Madeira Madeira, by far the largest island in the archipelago, was once covered with lush subtropical evergreen forests of a type that was once widespread throughout southern Europe and northwestern Africa. These laurel forest remnants survive intact today only on the steep northern slopes of the island. The Madeiran scops owl (Otus mauli) is known only from bones, but is believed to have been driven extinct by human settlers in the fifteenth century. Zino’s petrel (Pterodroma madeira) breeds only on six cliff ledges in the central mountains of Madeira, where the total population is around 200. During non-breeding seasons the species disperses towards the central Atlantic. The Madeiran laurel pigeon (Columba trocaz) is endemic to Madeira but formerly occurred on Porto Santo as well. At one time abundant, the species declined dramatically to around 2700 birds by 1986, but recovered rapidly soon after thanks to a total ban on hunting. It now occupies all of its former range on the island and is no longer considered to be threatened. The Desertas Islands The Desertas Islands are a small archipelago located roughly between Madeira Island and the Canary Islands. The Desertas petrel (Pterodroma deserta) breeds only on Bugio in the Desertas Islands, ranging throughout the North and South Atlantic at other times.

The Canary Islands The Canary Islands (Islas Canarias in Spanish) are a rugged, volcanic archipelago located approximately 100 km off the southern coast of Morocco. The Canarian shrew (Crocidura canariensis) is confined to the eastern Canary Islands of Lanzarote, Fuerteventura, Lobos, and Montaña Clara. The Canarian long-eared bat (Plecotus teneriffae) is confined to Tenerife, La Palma, El Hierro, and probably also La Gomera. The Canarian Egyptian vulture (Neophron percnopterus majorensis) is confined to the eastern Canary Islands. The Canarian houbara bustard (Chlamydotis undulata fuertaventurae) is confined to the eastern Canary Islands (Fuerteventura, Lanzarote, and La Graciosa), where the total population in the mid-1990s was estimated at between 500 and 1000. The Canarian black oystercatcher (Haematopus ostralegus meadewaldoi) was a type of shorebird that occurred on Fuerteventura, Lanzarote, and their offshore islets in the eastern Canary Islands. It was last collected in 1913 and locally reported to be absent by the 1940s. There were a few reports of sightings both in Tenerife and in Senegal between 1968 and

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The Palearctic Realm Figure 2.16 Canarian black oystercatcher. (Credit: Henrik Grönvold.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

1981, but extensive surveys since then have failed to find any evidence of continued survival. The cause of its extinction is believed to be due mainly to the overharvesting of its invertebrate prey. The Canarian quail (Coturnix gomerae) is known only from bones collected on El Hierro, La Palma, Tenerife, and Fuerteventura, and is believed to have still been present when humans first settled there. Bolle’s pigeon (Columba bollii) occurs on Tenerife, La Palma, La Gomera, and El Hierro, and perhaps formerly on Gran Canaria as well. The species suffered historical declines owing to the clearance of its montane laurel forest habitat, but has stabilized now that this has been slowed or stopped. It is no longer considered threatened. Simony’s cylindrical skink (Chalcides simonyi) is confined to Fuerteventura, Lanzarote, and Lobos. Tenerife Tenerife is the largest and most populous island in the Canaries. It features Mount Teide, the third tallest volcano in the world as measured from its base on the ocean floor. Also notable is the Macizo de Teno, a large massif in the extreme north-west of the island that reputedly has the largest number of endemic species in Europe. The Tenerife blue chaffinch (Fringilla teydea) is confined to the island where it appears to be largely dependent upon the presence of Canary pine (Pinus canariensis). It is frequently captured for use as a cage bird. The Tenerife speckled lizard (Gallotia intermedia) was first discovered in 1996 in the Macizo de Teno. It was later also found on a small area of coastline in the extreme west and from Montana de Guaza in the extreme south, suggesting that

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it was once widespread on the island. It is threatened mainly by feral cats. Fuerteventura The second largest of the Canary Islands, Fuerteventura was declared a UNESCO biosphere reserve in 2009. The Fuerteventura stonechat (Saxicola dacotiae) was historically widespread in the Canary Islands, but now breeds only on Fuerteventura and, occasionally, southern Lanzarote. Gran Canaria Gran Canaria is a roughly round island located south-east of Tenerife and west of Fuerteventura. The Gran Canaria blue chaffinch (Fringilla polatzeki) is confined to just two localities. A forest fire in 2007 cut its total population in half, but the species has since recovered. La Gomera La Gomera is a small, roughly circular and very mountainous island. Remaining areas of laurisilva (laurel rain forest) are protected by Garajonay National Park, although unfortunately nearly 20 per cent of this was lost as a result of a 2012 forest fire. The La Gomera giant lizard (Gallotia bravoana) was long thought to be extinct, but was rediscovered in 1999. It is confined to two separate, inaccessible cliffs some 2 km apart close to the Valle Gran Rey, in the west of the island. The total world population is around 150. Lanzarote Lanzarote is a sandy, rocky island located 11 km north-east of Fuerteventura.

The Saharo-Arabian Region

The Lanzarote chiffchaff (Phylloscopus canariensis exsul), a type of leaf warbler known from Lanzarote and possibly Fuerteventura, has not been recorded in many years. It is likely extinct due to loss of habitat. The Chinijo Archipelago is a group of small islands located north of Lanzarote, only one of which is inhabited. Together with the rocky northern coast of Lanzarote they have been designated as a national park. The Chinijo stonechat (Saxicola dacotiae murielae) formerly occurred on the islands of Alegranza and Montaña Clara, but became extinct during the early twentieth century due to a combination of natural factors and predation by introduced mammals. La Palma La Palma is the most north-westerly of the Canary Islands. Although large areas have been deforested, the upland areas still retain some temperate cloud forest. The La Palma giant lizard (Gallotia auaritae) was long thought extinct until a small, remnant population was rediscovered in northern La Palma in 2007. El Hierro El Hierro is the smallest and most south-westerly of the Canary Islands. The El Hierro chaffinch (Fringilla coelebs ombriosa) is threatened by loss of habitat but is still fairly common. Two subspecies of Simony’s giant lizard (Gallotia simonyi) were historically found on El Hierro. The Roque Chico de Salmor giant lizard (G. s. simonyi) was confined to a small offshore islet, where it went extinct in the 1930s. The El Hierro giant lizard (G. s. machadoi) was formerly present throughout much of the island, but is now restricted to a few small cliff areas.

Cyprus The third largest island in the Mediterranean, Cyprus (Cipros in Greek/Kibris in Turkish) is located south of Turkey and west of Syria and Lebanon. The Cyprian mouflon (Ovis gmelini ophion), a type of wild sheep, was nearly exterminated during the twentieth century but now numbers around 3000. The Cyprian white-throated dipper (Cinclus cinclus olympicus) was an endemic subspecies of passerine bird that formerly occurred in the Troodos Mountains. Last recorded in 1945, it was almost certainly extinct by 1958. The Cyprian whip snake (Hierophis cypriensis) is known from a few localities in the Troodos Mountains and the foothills of western Cyprus.

The Socotra Archipelago The Socotra Archipelago consists of four islands lying between the Horn of Africa and the Arabian Peninsula. Socotra Socotra (Suqutra in Arabic) is by far the largest of the four islands. Described as ‘the most alien-looking place on Earth’,

Socotra’s long geological isolation combined with its fierce heat has resulted in unusual plant and animal life. Unfortunately, 2000 years of human habitation, combined with introduced species, have seriously degraded the environment. The Socotra buzzard (Buteo socotraensis) is estimated to number around 500 individuals. The Socotra bunting (Emberiza socotrana) is confined to a few localities on Socotra Island. The Socotra house gecko (Hemidactylus dracaenoculus) is an interesting species, apparently endemic to a single plateau on Socotra where it is thought to be reliant on the resin of the dragon’s blood tree (Dracaena cinnabari). The short worm lizard (Pachycalamus brevis) is confined to Socotra Island. The Socotra worm snake (Xerotyphlops socotranus) is known only from a few specimens. Boulenger’s blind snake (Leptotyphlops macrurus), Wilson’s blind snake (L. wilsoni), and the Socotra blind snake (L. filiformis) are all confined to the island. Abd al Kuri Abd al Kuri is a small, rocky island located about 105 km south-west of Socotra. The Abd al Kuri sparrow (Passer hemileucus) is confined to this tiny island, where the total population is thought to be less than 1000.

Miscellaneous Islands The Dahlak Archipelago (Arakhbil Dahlak in Arabic) is an island group located near the coast of Eritrea in the Red Sea. It consists of 2 large and 124 small islands, only 4 of which are permanently inhabited. Cherlin’s saw-scaled viper (Echis megalocephalus) is known only from four specimens collected from ‘an island in the southern Red Sea’. It is thought that this may refer to Nokra Island in the Dahlak Archipelago, or perhaps Dissei Island. Astola Island (Jezira Haft Talar in Balochi) is a small, rocky, uninhabited island in the Arabian Sea off the coast of Pakistan. The Astola saw-scaled viper (Echis carinatus astolae) is confined to the island. Sarso Island is located in the Farasan Archipelago, a large coral island group located off south-western Saudi Arabia in the Red Sea. The Sarso racer (Coluber insulanus) is known from only a single specimen collected in 1964 on Sarso Island.

Balance for the Saharo-Arabian Region Modern human beings have inhabited the Saharo-Arabian Region for millennia, where they lived at first as huntergatherers. Western North Africa is believed to have been inhabited by Berbers since at least 10,000 B . C ., while the eastern part (the Nile Valley) was primarily home to the Egyptian civilization. Rock art findings within the Sahara have shown that it hosted various populations before its rapid desertification in 3500 B . C ., and indeed continues to host a small number of nomadic peoples to this day. The history of

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The Palearctic Realm

the Arabian Peninsula, meanwhile, goes back to the very beginnings of human habitation up to 130,000 B . C ., with earlier hominins having lived there very much longer. Its harsh climate, however, prevented much settlement apart from a small number of urban trading settlements such as Mecca and Medina. Throughout the Middle East human impact on the environment was minimal, although in many parts of North Africa and the Middle East large stone corrals were constructed in order to drive herds of gazelle into, making for an easy kill. This method of hunting began in prehistoric times and continued into the early twentieth century. The ancient cities and early civilizations of the Nile Valley, North Africa, the Near East, and the Arabian Peninsula had all been reasonably familiar to Europeans since classical antiquity. In 332 B . C . Alexander the Great was welcomed as a liberator in Persian-occupied Egypt, where he would subsequently found the city of Alexandria. It would become the prosperous capital of the Ptolemaic dynasty after his death. Following the conquest of the North African and eastern Mediterranean coastline by the Romans, the area was fully integrated into the empire. Roman settlement occurred in what is now Tunisia, Libya, and elsewhere along the coast. Unfortunately, during the Roman Era many species and subspecies were driven to extinction or extirpation by hunting and collection for use in gladiatorial spectacles. Notable among these were the North African elephant (Loxodonta africana pharaoensis) and African lion (Panthera

leo). Generally speaking, however, the Greeks, Phoenicians, and Romans who settled along the coast of North Africa improved the environment, stabilized it and allowed it to flourish. All of these efforts would ultimately be cancelled out, however, by the anarchic impact of invading nomads and conquering Arab tribes, who brought on desert-like conditions and a complete disregard for wildlife. The environmental situation improved during the British, French, and Italian colonial period of the early twentieth century, although since that time the habitat destruction, desertification, and drought have continued to this day. Moreover, the region’s significant stocks of crude oil have created new dangers related to pollution. In recent historical time (i.e. since A . D . 1500), the SaharoArabian Region has lost at least 9 species/10 subspecies of vertebrates. Among the extinct forms 3 species/2 subspecies are mammals, 2 species/5 subspecies are birds, 1 species/1 subspecies are reptiles, and 3 species/2 subspecies are freshwater fishes. Another 8 species/2 subspecies are possibly extinct. In addition, there are 230 species/50 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 57 species/30 subspecies are mammals, 21 species/7 subspecies are birds, 82 species/9 subspecies are reptiles, 10 species/4 subspecies are amphibians, and 60 species are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

3 species

~ species

~ species

57 species

2 subspecies

1 subspecies

~ subspecies

30 subspecies

5 taxa

1 taxon

~ taxa

87 taxa

2 species

~ species

~ species

21 species

5 subspecies

1 subspecies

~ subspecies

7 subspecies

7 taxa

1 taxon

~ taxa

28 taxa

1 species

1 species

~ species

82 species

1 subspecies

~ subspecies

~ subspecies

9 subspecies

2 taxa

1 taxon

~ taxa

91 taxa

~ species

1 species

~ species

10 species

~ subspecies

~ subspecies

~ subspecies

4 subspecies

~ taxa

1 taxon

~ taxa

14 taxa

3 species

6 species

~ species

60 species

2 subspecies

~ subspecies

~ subspecies

~ subspecies

5 taxa

6 taxa

~ taxa

60 taxa

9 species

8 species

~ species

230 species

10 subspecies

2 subspecies

subspecies

50 subspecies

19 taxa

10 taxa

~ taxa

280 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

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Balance for the Palearctic Realm

Balance for the Palearctic Realm In recent historical time (i.e. since A . D . 1500), the Palearctic Realm as a whole has lost at least 47 species/25 subspecies of vertebrates. Among the extinct forms 6 species/13 subspecies are mammals, 6 species/5 subspecies are birds, 1 species/4 subspecies are reptiles, 1 species is an amphibian, and 30 species/2 subspecies are freshwater fishes. Another 37 species/4 subspecies are possibly extinct.

In addition, there are 1327 species/206 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 221 species/122 subspecies are mammals, 111 species/22 subspecies are birds, 191 species/45 subspecies are reptiles, 229 species/11 subspecies are amphibians, and 575 species/6 subspecies are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

6 species

2 species

~ species

221 species

13 subspecies

3 subspecies

~ subspecies

122 subspecies

19 taxa

5 taxa

~ taxa

344 taxa

6 species

~ species

~ species

111 species

5 subspecies

1 subspecies

~ subspecies

22 subspecies

11 taxa

1 taxon

~ taxa

133 taxa

1 species

2 species

~ species

191 species

4 subspecies

~ subspecies

~ subspecies

45 subspecies

5 taxa

2 taxa

~ taxa

236 taxa

1 species

2 species

~ species

229 species

~ subspecies

~ subspecies

~ subspecies

11 subspecies

1 taxon

2 taxa

~ taxa

240 taxa

30 species

31 species

~ species

575 species

2 subspecies

~ subspecies

~ subspecies

6 subspecies

32 taxa

31 taxa

~ taxa

581 taxa

47 species

37 species

~ species

1327 species

25 subspecies

4 subspecies

~ subspecies

206 subspecies

76 taxa

41 taxa

~ taxa

1533 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

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Chapter

3

The Afrotropical Realm

The Afrotropical Realm includes all of sub-Saharan Africa along with certain coastal islands such as the Cape Verde Archipelago. Madagascar, although traditionally included within Africa, is treated separately here. Biogeographically the Afrotropical Realm is divided between the African and Guineo-Congolian zoogeographic regions. Africa as a whole has a wide range of climates and habitats, ranging from subtropical to temperate. South of the Saharo-Arabian Region the vast North African desert turns into subdesert, grass, and shrub savannas, and finally to more or less dense wooded savannas. Still further south, forests appear; such growths reach their optimum development in the Congo Basin. In fact, Africa both north and south of the equator can show all types of nature from extreme desert to extreme rainforest. Savannas cover about 40 per cent of Africa, in both subtropical and tropical areas, and they vary greatly from region to region. Africa’s savannas, steppes, arid plains, and subdeserts stretch in an arc across the continent. Beginning in East Africa between the latitudes 20° and 10° N, they fill practically the whole of Africa south of 10° S with the exception of the Cape. Some of these plains have become or are becoming deserts; others are still fertile grass or tree savannas, which in places turn into open forests. No other continent historically possessed such a wealth of fauna as Africa. Indeed, more species of terrestrial vertebrates still live here than anywhere else on Earth, and the density of large mammals on some areas of savanna remains unparalleled. The reason for this is probably that Africa’s fauna is of a very great age; it has had countless eons in which to become adapted to the many types of habitat. Repeated changes of climate, continuously altering the landscape, have caused dislocations in the distribution of plants and animals. These dislocations in turn have favoured speciation and the evolution of species in special directions. To this must be added an evolutionary phenomenon typical of all the Africa fauna – by competition the animals have become fitted to utilize all the possibilities offered by the rich environment. This has enabled numerous species to live side by side in the same habitats and in harmony with the vegetation by utilizing different niches. It is in the tropical areas that we find the largest number of species and most of the spectacular animals which have made Africa famous. The southern part of the continent is not so rich, but there the fauna includes a wealth of vertebrate species.

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The African buffalo (Syncerus caffer) is still found across sub-Saharan Africa, but is now mostly confined to protected areas, as it is a favourite target for trophy hunters. The northwest Africa buffalo (S. c. brachyceros) occurs patchily in the savannas and gallery forests of western and central Africa. The Nile buffalo (S. c. aequinoctialis) is found in the SaheloSudanian savanna zone of central Africa (south-eastern Chad, northern Central African Republic, northern Democratic Republic of Congo, south-eastern Sudan, and western Ethiopia. It has been extirpated from Eritrea. The hippopotamus (Hippopotamus amphibius) is a semiaquatic animal that was historically found throughout subSaharan Africa and the Nile Valley. From the end of the Roman Empire up to the beginning of the eighteenth century at the latest the animals were still present in two disjunct areas in the northern part of their range (the Nile delta and the upper Nile), although the species appears to have become extinct in Egypt sometime during the eighteenth century. It still occupies much of the area that it did in 1960, although population sizes have declined greatly. The main threats are habitat destruction and poaching for their meat and ivory canine teeth. The yellow-backed duiker (Cephalophus silvicultor) is a type of small antelope found widely across western, central, and parts of eastern Africa. The short-headed yellow-backed duiker (C. s. curticeps), long-headed yellow-backed duiker (C. s. longiceps), Bocage’s yellow-backed duiker (C. s. ruficrista), and Afzelius’ yellow-backed duiker (C. s. silvicultor) are all threatened by bushmeat hunting and snaring. The black and white colobus (Colobus angolensis) is a type of leaf-eating monkey divided into a number of subspecies found across Central Africa. All are threatened by loss of habitat and hunting, and will be discussed individually below. The African golden cat (Caracal aurata) is a rare forest inhabitant, with two widely separated populations in central and western equatorial Africa, respectively. It is heavily hunted for its pelt and threatened by loss of habitat. The russet free-tailed bat (Chaerephon russatus) is only known from five widely separated localities in Ivory Coast, Ghana, central Cameroon, the Democratic Republic of Congo, and south-western Kenya. Robbins’ house bat (Scotophilus nucella) is known only from south-eastern Ivory Coast, southern Ghana, western Uganda, and north-eastern Tanzania.

The Guineo-Congolian Region

Beaudouin’s snake-eagle (Circaetus beaudouini) occurs sparsely within a narrow band of savanna and woodlands in sub-Saharan Africa from Senegal, Gambia, and southern Mauritania in the west to South Sudan and the northern Democratic Republic of Congo, with some nomadic movement to outlying countries such as Kenya. It is everywhere threatened by loss of habitat, overgrazing by cattle, pesticides, and hunting pressure. The white-backed vulture (Gyps africanus) was historically the most widespread and common vulture in Africa, being found virtually everywhere south of the Sahara except for the rainforest belt and a few coastal areas. In recent decades it has undergone drastic declines due to habitat destruction, decreases in wild ungulate populations, hunting, human persecution, collisions, and indirect poisoning, and is now considered to be seriously threatened. Rüppell’s vulture (G. rueppelli) was also formerly widespread within the grasslands, mountains, and woodlands of Sahelian and eastern Africa, where it has also undergone similarly drastic declines and is now largely confined to protected areas. The nominate form (G. r. rueppelli) occupies the majority of the range, while the Ethiopian Rüppell’s vulture (G. r. erlangeri) inhabits the highlands of Ethiopia, Eritrea, north-western Somalia, and possibly the southern Arabian Peninsula. Interestingly, this species is thought to be the highest-flying of all birds, with one individual in 1973 having been ingested by a jet engine flying over Ivory Coast at 11,300 m elevation. Since the 1990s there have been increasing records involving small numbers of individuals in Spain and Portugal, although breeding is not known to have taken place. The white-headed vulture (Trigonoceps occipitalis) is found patchily across sub-Saharan Africa but is everywhere threatened by habitat destruction, reduction of prey, indirect poisoning, and collection for use in ‘traditional medicine’. The hooded vulture (Necrosyrtes monachus) is still widespread across sub-Saharan Africa but is undergoing rapid declines almost everywhere due to indiscriminate poisoning, habitat destruction, hunting, human persecution, and collection for use in ‘traditional medicine’. The African lappet-faced vulture (Torgos tracheliotos tracheliotos) occurs patchily throughout sub-Saharan Africa, where it is threatened by habitat destruction, human disturbance of its nesting sites, and pesticide poisoning.

The Guineo-Congolian Region The Guineo-Congolian Region encompasses the tropical rainforest belt of West and Central Africa along with the islands of the Gulf of Guinea. It is bordered to the north, east, and south by the African Region. Two species of gorilla inhabit the Guineo-Congolian Region. The eastern gorilla (Gorilla beringei) is the largest living primate. Two well-defined subspecies are found in remote areas of central Africa. Grauer’s gorilla (G. b. graueri) is often called the eastern lowland gorilla in spite of having the

widest altitudinal range of any gorilla (approximately 600–2900 m). It is endemic to the forests of the Albertine Rift escarpment in eastern Democratic Republic of Congo, where it has a discontinuous distribution from the lowlands east of the Lualaba River to the Mitumba Mountains and the Itombwe Massif. Mount Tshiaberimu in Virunga National Park is the northern limit of the range, while the southern limit is a subpopulation in the Hewa Bora region, Fizi District. Important populations live in the Kahuzi-Biega and Maiko national parks and their adjacent forests, the Tayna Gorilla Reserve, the Usala Forest, and on the Itombwe Massif. The western gorilla (G. gorilla) is generally smaller and lightercoloured than the eastern gorilla (G. beringei). There are two subspecies, which will also be discussed below. The chimpanzee (Pan troglodytes) is, along with the bonobo (P. paniscus – see below), the closest living human relative. Four subspecies are found discontinuously in the tropical rainforests and wet savanna woodlands of Western and Central Africa, where they are highly threatened almost everywhere by loss of habitat, Ebola, and hunting for their meat. The West African or western chimpanzee (P. t. verus) was historically found from southern Senegal all the way east to the Niger River, but has been greatly reduced by forest loss and bushmeat hunting. Today the largest numbers are found in Ivory Coast and Guinea (in particular, the Fouta Djallon Highlands, where some 17,700 survive). Considerable populations still also occur, although patchily, in Liberia, Sierra Leone, and Guinea-Bissau, with relicts in Ghana, Senegal, Mali, and western Nigeria. It is believed to have been extirpated from Gambia, Benin, Burkina Faso, and Togo relatively recently. Less than 65,000 remain. The East African or eastern chimpanzee (P. t. schweinfurthii) ranges from the Oubangui/Congo in south-eastern Central African Republic and Democratic Republic of Congo to Burundi, Rwanda, western Uganda, and western Tanzania, with a small, relict population in South Sudan. The vast majority (about 173,000–248,000) live in the Democratic Republic of Congo, although there are still significant populations within the Albertine Rift. In 2010 the total population was estimated at 181,000–256,000. The other subspecies of chimpanzee will be discussed elsewhere in this book. The red-capped mangabey (Cercocebus torquatus) is a large terrestrial monkey found in the lowland rainforests as well as swamp and mangrove forests of central-western coastal Africa (southern Nigeria, western Cameroon, Equatorial Guinea, and western Gabon), where it is threatened by loss of habitat and bushmeat hunting. L’Hoest’s monkey (Allochrocebus lhoesti) is a semiterrestrial species found patchily in lowland as well as montane forests of the central-eastern Democratic Republic of Congo, north-western Burundi, south-western Rwanda, and southwestern Uganda, where it is threatened by loss of habitat and bushmeat hunting. The guenons are a group of mostly small, often brightly coloured arboreal monkeys from the African tropical forest belt. While most remain common, a few have become threatened due to loss of habitat and bushmeat hunting. The owl-faced guenon

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(C. hamlyni) is a largely terrestrial species of monkey that lives in both lowland as well as montane forest in eastern Democratic Republic of Congo and Rwanda. The crowned guenon (C. pogonias) is divided into three subspieces. The golden-bellied crowned guenon (C. p. pogonias) is found in south-eastern Nigeria, north-western Cameroon, and the island of Bioko. Temminck’s red colobus (Piliocolobus temminckii) is a type of leaf-eating monkey found patchily in both dry and moist forest fragments in southern Senegal, Guinea-Bissau, and north-western Guinea. It has been seriously reduced by habitat destruction and bushmeat hunting.

Figure 3.1 Jentink’s duiker (Cephalophus jentinki). (Credit: Philip Sclater.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

Jentink’s duiker (Cephalophus jentinki) is a type of small antelope found patchily in south-eastern Guinea, Sierra Leone, Liberia, and western Ivory Coast. The banded duiker (C. zebra) is found in southern Sierra Leone, southern Guinea, central and eastern Liberia, and south-western Ivory Coast. Brooke’s duiker (C. brookei) is confined to Sierra Leone, Liberia, Ivory Coast, and south-western Ghana. All are threatened by loss of habitat and heavy hunting pressure. The king genet (Genetta poensis) is known only from a few specimens collected from the Republic of Congo, Liberia, Ghana, Ivory Coast and the island of Bioko. It was last recorded in 1946. The crested genet (G. cristata) is a rare species found in southern Nigeria and western Cameroon, where it is threatened by loss of habitat and hunting. The giant ground pangolin (Smutsia gigantea) is, as its name suggests, the largest species of pangolin or scaly anteater. While still fairly widespread in western and central Africa it is nevertheless under threat from habitat destruction and bushmeat hunting. The black-bellied pangolin (Phataginus tetradactyla) and the white-bellied tree pangolin (P. tricuspis) are both found widely in the forests of western and central Africa, but are everywhere threatened by habitat destruction and hunting. The moon shrew (Crocidura selina) is known only from three swamp forest localities in central Uganda. A record from southern Kenya most likely refers to a different species. Maclaud’s horseshoe bat (Rhinolophus maclaudi) is known only from a few scattered localities in western Guinea. Eisentraut’s horseshoe bat (R. guineensis) is known from fewer than 20 cave and mine roosts in southern Senegal, Guinea, Sierra Leone, Liberia, and Ivory Coast. The Ziama horseshoe bat (R. ziama) is known only from three specimens

Figure 3.2 Giant ground pangolin (Smutsia gigantea). (Credit: Joseph Wolf.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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collected from the Ziama Massif in south-eastern Guinea and two others from Liberia and Sierra Leone. The Lagos serotine bat (Eptesicus platyops) is known only from a few specimens collected in Nigeria, Senegal, and on Bioko. Aellen’s roundleaf bat (Hipposideros marisae) is known only from a few localities in Guinea, Liberia, Sierra Leone, and Ivory Coast. The short-tailed roundleaf bat (H. curtus) is known from Cameroon, Nigeria, and Equatorial Guinea (including the island of Bioko). Both are threatened by loss of habitat and disturbance of their roosting sites. The copper woolly bat (Kerivoula cuprosa) is known only from a few scattered records across western and central Africa. The brown-cheeked hornbill (Bycanistes cylindricus) is found widely but patchily in Ivory Coast, Ghana, Guinea, Liberia, Sierra Leone, and Togo, where it is threatened by hunting and habitat destruction. The African grey parrot (Psittacus erithacus) remains widespread within western and central Africa from Ivory Coast to western Kenya and south to Angola, including islands in the Gulf of Guinea. The species is everywhere declining, however, by habitat destruction, hunting, and collection for the international pet trade. The closely related Timneh grey parrot (P. timneh) is found in Guinea-Bissau, southern Guinea, Sierra Leone, Liberia, and south-western Ivory Coast, where it too is threatened by loss of habitat and heavy trapping pressure. The white-breasted guineafowl (Agelastes meleagrides) is a terrestrial species found patchily in Ivory Coast, Ghana, Guinea, Liberia, and Sierra Leone. It is threatened by loss of habitat and hunting. The grey-necked rockfowl (Picathartes oreas) is confined to isolated rocky areas within the rainforests of south-eastern Nigeria, western and southern Cameroon, Gabon, Equatorial Guinea (including the island of Bioko), and the Republic of Congo. It is everywhere threatened by loss of habitat. The spotted ground thrush (Geokichla guttata) has a wide but very spotty distribution across sub-Saharan Africa, where a number of subspecies are threatened by loss of habitat. The South Sudan spotted ground thrush (G. g. maxis) from southern South Sudan and Lippens’ spotted ground thrush (G. g. lippensi) from south-eastern Democratic Republic of Congo are each known only from a single specimen. The white-headed robin-chat (Cossypha heinrichi) is known from two localities in northern Angola and a third in south-western Democratic Republic of Congo. The African river martin (Pseudochelidon eurystomina) is relatively widespread in central Africa, where it breeds in large but infrequent colonies along forested rivers that are highly vulnerable to exploitation. The Lufira masked weaver (Ploceus ruweti) is confined to swamps and marshes bording the Lufira River and Lake Lufira in south-eastern Democratic Republic of Congo. The Loango weaver (P. subpersonatus) is a rare species confined to a narrow coastal strip in Gabon, Republic of Congo, and the

Democratic Republic of Congo, where it lives in swamps, mangroves, and grassland areas. The Anambra waxbill (Estrilda poliopareia) is found patchily in southern Nigeria and south-eastern Benin. Turner’s eremomela (Eremomela turneri) is a warbler-like bird known from a few scattered localities in east-central Africa. The nominate form (E. t. turneri) is known from the Kakamega and South and North Nandi forests of western Kenya, with further historical records from two other undefined localities. Müller’s worm lizard (Cynisca leonina) is known from the islands of Tumbo and Kassa in the Los Archipelago off the coast of south-western Guinea, as well as in a few gallery forest localities in the north-west. The four-digit toad (Didynamipus sjostedti) was once thought to be the rarest toad in Africa, but in recent years has been found in a few mid-elevation sites in southwestern Cameroon, south-eastern Nigeria, and on the island of Bioko. Bornmüller’s caecilian (Crotaphatrema bornmuelleri) is a type of burrowing, limbless amphibian known only from a single specimen collected from an undefined locality in southwestern Cameroon in the late nineteenth century.

Mountains and Highlands The Guineo-Congolian Region features a number of mountainous areas rich in species endemism. During long periods the animals adapted to life on these high mountains, and were not disturbed. The inaccessibility, cold, damp climate and density of the vegetation were all obstacles to humans and their livestock. This has now changed, and people now penetrate with fire and cattle ever higher up the mountain slopes, violating the forests and threatening the animals living here. This process has created a series of widely separated and biologically important ‘sky islands’. The western klipspringer (Oreotragus oreotragus porteousi) is a type of small antelope known only from the rocky margins of the Jos Plateau in north-central Nigeria and in GashakaGumti National Park in eastern Nigeria, along with two other localities in northern Central African Republic. Fox’s mole-rat (Fukomys foxi) is known only from the Jos Plateau of central Nigeria and from another locality in western Cameroon. The Cameroon soft-furred mouse (Praomys morio) is known only from Mount Cameroon and the island of Bioko. Sladen’s swift (Apus sladeniae) is a mysterious species known from the mountains of Bioko (where six specimens were collected in 1903, although none since) and a few other, disjunct montane areas in south-eastern Nigeria, Cameroon, Angola, and Equatorial Guinea. The Mebebque frog (Arlequinus krebsi) was long known only from a few specimens collected from the Bamileke Plateau along with an additional one taken from the southern slopes of Mount Cameroon. Based on an old photograph the species is

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thought also to occur on Bioko, although this has yet to be confirmed. It appears to be fairly widespread in south-western Cameroon but naturally rare.

The Albertine Rift Mountains The Albertine Rift is the western branch of the East African Rift and covers parts of Uganda, Democratic Republic of Congo, Rwanda, Burundi, and western Tanzania. It extends from the northern end of Lake Albert to the southern end of Lake Tanganyika. The geographical term includes both the valley and the surrounding mountains. The mountain gorilla (Gorilla beringei beringei) is nowadays confined to two surviving populations in montane forests only 25 km apart, but isolated by land that is intensively cultivated and densely settled. One population is in the Virunga Mountains, in three separate national parks straddling the borders between the Democratic Republic of Congo (Virunga National Park), Rwanda (Volcanoes National Park), and Uganda (Mgahinga Gorilla National Park). The other occurs in Bwindi Impenetrable National Park, Uganda, with a small contiguous portion in Sarambwe Nature Reserve in the Democratic Republic of Congo. In 2018 the total population was just over 1000. These populations will be discussed at greater length below. The ashy red colobus (Piliocolobus tephrosceles) is found discontinuously on the eastern border of the Rift Valley in western Uganda and western Tanzania, and possibly in parts of the eastern Democratic Republic of Congo, Rwanda, and Burundi as well. It is everywhere threatened by bushmeat hunting and habitat destruction, and many populations have been extirpated. The Ruwenzori black and white colobus (Colobus angolensis ruwenzorii) is a type of leaf-eating monkey found within a few montane forest pockets in eastern Democratic Republic of Congo, Burundi, Rwanda, southern Uganda, and western Tanzania. Kemp’s thicket rat (Thamnomys kempi) has a fragmented distribution within the Albertine Rift Mountains of eastern Democratic Republic of Congo, Burundi, and Rwanda. The medium-tailed brush-furred rat (Lophuromys medicaudatus) is a rare species known from high elevations on both sides of Lake Kivu in the Democratic Republic of Congo and Rwanda, with a single further record from the Bwindi Forest of Uganda. De Graaff’s soft-furred mouse (Praomys degraaffi) is found patchily in the higher elevations of the Albertine Rift in Burundi, Rwanda, and south-western Uganda. Delany’s swamp mouse (Delanymys brooksi) has a patchy distribution within the Albertine Rift of south-western Uganda, western Rwanda, Burundi, and extreme eastern Democratic Republic of Congo, where it occurs in or very close to high-elevation swamps. The Lemara shrew (Crocidura lanosa) and the Tarella shrew (C. tarella) are both known only from a few localities in the Albertine Rift.

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The Ruwenzori shrew (Ruwenzorisorex suncoides) is a semi-aquatic species endemic to the Albertine Rift. The Ruwenzori horseshoe bat (Rhinolophus ruwenzorii) is confined to the Albertine Rift, where it has been recorded from Democratic Republic of Congo, Uganda, and Rwanda. Hill’s horseshoe bat (R. hilli) is known only from Nyungwe National Park in south-western Rwanda. Rockefeller’s sunbird (Cinnyris rockefelleri) is confined to a small area of the northern Itombwe Mountains and other mountains to the north and west of Lake Kivu in eastern Democratic Republic of Congo, where the total population is thought to be under 1500. Prigogine’s greenbul (Chlorocichla prigoginei) is a type of passerine bird known only from a few high-elevation localities in eastern Democratic Republic of Congo, where it is seriously threatened by habitat destruction. Grauer’s broadbill (Pseudocalyptomena graueri) is confined to the montane forests of the Albertine Rift in eastern Democratic Republic of Congo, and south-western Uganda. At one time an almost legendary bird among ornithologists working in Africa, it remains rare and little-known. The yellow-crested helmet-shrike (Prionops alberti) is known from the Albertine Rift of central-eastern Democratic Republic of Congo, including the Itombwe Plateau and Mount Kabobo. The total population is thought to be small and threatened by loss of habitat. Chapin’s flycatcher (Muscicapa lendu) is a rare species found patchily in eastern Democratic Republic of Congo, south-western Uganda, western Kenya, and possibly Rwanda, where it is threatened by loss of habitat. Grauer’s swamp warbler (Bradypterus graueri) is found patchily in Rwanda, Burundi, eastern Democratic Republic of Congo, and far south-western Uganda, where it is restricted to high-elevation marshy areas. Shelley’s crimsonwing (Cryptospiza shelleyi) is a type of finch known from Burundi, eastern Democratic Republic of Congo, Rwanda, and western Uganda. It is threatened by loss of habitat. Blochmann’s skink (Leptosiaphos blochmanni) is known only from the Nyungwe Forest in south-western Rwanda. Leopold’s snake-eater (Polemon leopoldi) is a type of rearfanged snake known only from a single specimen collected in 1941 from the mountains of Rwanda. The Karissimbi tree frog (Leptopelis karissimbensis) is found patchily in the highlands of far south-western Uganda, north-western and south-western Rwanda, and eastern Democratic Republic of Congo, where it is threatened by loss of habitat. Several species of reed frog (Hyperolius) are threatened by loss of habitat. The Nyungwe reed frog (H. jackie) is known only from Nyungwe National Park in south-western Rwanda. The diaphanous reed frog (H. diaphanous) is known only from a small area of the Itombwe Mountains and the mountains west of Lake Kivu. The white-striped reed frog (H. leucotaenius) is known only from a small area to the west of

The Guineo-Congolian Region

the southern Itombwe Highlands and from the northern slopes of the Kabobo Highlands. The starry reed frog (H. constellatus) is confined to the Itombwe and Kabobo plateaus west of Lake Tanganyika. Raven’s reed frog (H. raveni) is known only from its original collection in the 1930s from an undefined volcanic area north-east of Lake Kivu. The Itombwe screeching frog (Arthroleptis hematogaster) is confined to the Itombwe and Kabobo highlands of eastcentral Democratic Republic of Congo. The Mosso screeching frog (A. mossoensis) is known only from its original collection in the 1950s from southern Burundi. The Mukuzira long-fingered frog (Cardioglossa cyaneospila) is known only from a few ill-defined localities in Rwanda, Burundi, and eastern Democratic Republic of Congo. The yellow-throated puddle frog (Phrynobatrachus sulfureogularis) is known only from two small, disjunct areas of western Burundi and western Tanzania. Dalcq’s puddle frog (P. dalcqi) is known only from its original collection in the 1950s from eastern Democratic Republic of Congo (southern Kivu province). The Albertine Rift bamboo frog (Callixalus pictus) is a rare species known only from the Itombwe and Kabobo highlands of the eastern Democratic Republic of Congo. Fischer’s caecilian (Boulengerula fischeri) is known only from the Cyamudongo Forest in south-western Rwanda, a relict of the larger Nyungwe Forest. The Mitumba Mountains The Mitumba Mountains stretch along the western shore of Lake Tanganyika in eastern Democratic Republic of the Congo (Sud-Kivu and Nord-Kivu provinces). Part of it is protected by the Kahuzi-Biéga National Park, a World Heritage Site. The grey brush-furred rat (Lophuromys cinereus) is known only from a few specimens collected at high elevations in Kahuzi-Biéga National Park. The Kahuzi swamp shrew (Myosorex jejei) is similarly confined to areas of Kahuzi-Biéga National Park. The Mwana screeching frog (Arthroleptis vercammeni) is known only from a single locality in the Mitumba Mountains. Mount Kahuzi is an extinct volcano and the highest peak in the Mitumba Range. The Kahuzi climbing mouse (Dendromus kahuziensis) is known only from two specimens collected in montane forest on Mount Kahuzi in 1969. Mount Nzawa is located in south-eastern Democratic Republic of Congo (Katanga province). The Nzawa chameleon (Kinyongia mulyai) and the Nzawa pygmy chameleon (Rhampholeon hattinghi) are both known only from a small, highly fragmented forest remnant on Mount Nzawa. The Lendu Plateau The Lendu Plateau is located in north-eastern Democratic Republic of Congo, on the western side of Lake Albert.

Chapin’s crombec (Sylvietta chapini) is a very rare (possibly extinct) type of warbler known only from the Lendu Plateau. The circular-scaled chameleon (Kinyongia gyrolepis) is known only from a few specimens collected mainly in the Lendu Plateau’s now mostly cleared forests. The Lendu Plateau clawed frog (Xenopus lenduensis) is an aquatic species confined to the Lendu Plateau. The Ruwenzori Mountains The Ruwenzori Mountains (also known as the Rwenzori Mountains) are located between lakes Edward and Albert on the border between Uganda and north-eastern Democratic Republic of Congo. The mountains are non-volcanic and its highest peaks are permanently snow-capped. The Ruwenzori red duiker (Cephalophus nigrifrons rubidus) is confined to the Ruwenzori Mountains. The montane shaggy rat (Dasymys montanus) is confined to the Ruwenzori Mountains. The moon striped mouse (Hybomys lunaris) is confined to the Ruwenzori Mountains. The montane mouse shrew (Myosorex blarina) is confined to the Ruwenzori Mountains. The Ruwenzori skink (Leptosiaphos meleagris) is confined to the Ruwenzori Mountains. The Ruwenzori yellowfish (Labeobarbus ruwenzori) is confined to the Mubuku and Sibwe rivers on the eastern flank of the Ruwenzori Mountains in Uganda. Mount Teye is located in north-eastern Democratic Republic of Congo. The Teye reed frog (Hyperolius xenorhinus) is known only from a single specimen collected on Mount Teye. The Virunga Mountains The Virunga Mountains are a chain of eight mostly dormant volcanoes located along the border of eastern Democratic Republic of Congo (Nord-Kivu province), northern Rwanda, and western Uganda. The mountain gorilla (Gorilla beringei beringei) has, as indicated earlier, one of its two surviving populations in the Virunga Mountains, where it inhabits montane cloud forests ranging in altitude from 2200 to 4300 m. Most are found on the slopes of three dormant volcanoes (Karisimbi, Mikeno, and Visoke). The vegetation is very dense at the bottom of these mountains, becoming sparser at higher elevations, and the broad-leaved Hagenia forests where the gorillas live are often cloudy, misty, and cold. Since recovering from an all-time low of just 254 in 1981 the population has grown considerably to 480 by 2010, and is completely protected within three contiguous, well-patrolled national parks. Nevertheless, the animals are still threatened by habitat destruction and degradation, disease, and snares set by bushmeat poachers. The golden guenon (Cercopithecus kandti) is confined to the Virunga Mountains of north-western Rwanda and eastern Democratic Republic of Congo.

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The Albertine owlet (Glaucidium albertinum) is known only from a handful of specimens collected within the Itombwe Mountains and from a forest west of Lake Edward. The population is inferred to be very small, fragmented, and under threat from habitat destruction. The Virunga forest tree frog (Leptopelis fenestratus) is known only from the Semliki sector of the Virunga Mountains in Democratic Republic of Congo. The Bwindi Impenetrable Forest The Bwindi Impenetrable Forest is a large area of primeval forest located on the edge of the Albertine Rift, in southwestern Uganda. This forest contains one of the two remaining populations of mountain gorilla (Gorilla beringei beringei). As of 2006 approximately 300–340 gorillas were protected within Bwindi Impenetrable National Park (Uganda), with a small contiguous portion in the Sarambwe Nature Reserve, Democratic Republic of Congo. The Itombwe Mountains The Itombwe Mountains (also know as the Itombwe Massif ) are a range of mountains running along the western shore of Lake Tanganyika in east-central Democratic Republic of Congo (Sud-Kivu province). It contains vast areas of both montane and lowland forest that is home to a variety of wildlife, including populations of Grauer’s gorillas (Gorilla beringei graueri) and African forest elephants (Loxodonta cyclotis). Unfortunately, it is still unprotected and under increasing threat by local people. Grauer’s large-headed shrew (Paracrocidura graueri) is known only from a single specimen collected in the Itombwe Mountains in 1908. Schaller’s mouse shrew (Myosorex schalleri) is known only from a single specimen collected in the Itombwe Mountains in 1966. The Congo bay owl (Phodilus prigoginei) was long known only from a single specimen collected in 1951 at Muusi. However, another was captured in a mist-net in 1996 in the south-east corner of Itombwe Forest. It is possible that the species occurs much more widely. The Itombwe flycatcher (Muscicapa itombwensis) is confined to a narrow band of montane forest within the Itombwe Mountains. Prigogine’s nightjar (Caprimulgus prigoginei) is known only from a single specimen collected in the Itombwe Mountains in 1955, although subsequent sound recordings indicate that it may be much more widespread in equatorial Africa. Parker’s tree toad (Laurentophryne parkeri) is known only from its original collection in montane forest near Mwengwa in 1951. The Itombwe forest tree frog (Leptopelis anebos) is known only from two localities in the Itombwe Mountains. The Luvubu reed frog (Hyperolius leleupi) is confined to an area of high-elevation bamboo forest in the Itombwe

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Mountains, where it has not been seen since its original description in 1951. The Itombwe golden frog (Chrysobatrachus cupreonitens) is confined to montane grassland and marshy areas in the Itombwe Mountains. The Itombwe puddle frog (Phrynobatrachus asper) is confined to high-elevation swamps in the Itombwe Mountains. The Itombwe clawed frog (Xenopus itombwensis) is an aquatic species confined to marshes and pools in the Itombwe Mountains. The Kibara Mountains The Kibara Mountains are located in south-eastern Democratic Republic of Congo (Katanga province). The Shaba reed frog (Hyperolius kibarae) is known only from the highlands of Upemba National Park. Mount Bururi Mount Bururi is located in south-western Burundi. The Bururi forest shrew (Myosorex bururiensis) is a rare species confined to montane forest in the Bururi Nature Reserve. Mount Kabobo Mount Kabobo is located in eastern Democratic Republic of Congo. Prigogine’s black and white colobus (Colobus angolensis prigoginei) is a type of monkey confined to Mount Kabobo. The Marungu Highlands The Marungu Highlands are located west of the southern half of Lake Tanganyika in south-eastern Democratic Republic of the Congo. The Marungu reed frog (Hyperolius atrigularis) is known only from its original collection in the 1940s, in montane grassland of the Marungu Highlands.

The Cameroon Volcanic Line The Cameroon Volcanic Line is a 1600-km chain of volcanoes stretching across the border between western Cameroon and eastern Nigeria. Geologically it also includes the islands of the Gulf of Guinea, which are dealt with separately in this volume. The mountain reedbuck (Redunca fulvorufula) is a type of antelope divided into three subspecies found very disjunctly in eastern, western, and southern Africa, respectively. The western mountain reedbuck (R. f. adamauae) has been reduced to a few small populations in eastern Nigeria and western Cameroon, where it is threatened by hunting and loss of habitat. Two subspecies of Preuss’ monkey (Allochrocebus preussi) are threatened by hunting and habitat destruction. The Cross River Preuss’ monkey (A. p. preussi) is largely confined to higher-elevation forest pockets in western Cameroon and south-eastern Nigeria.

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Cooper’s mountain squirrel (Paraxerus cooperi) is a littleknown species found patchily in the mountains of western Cameroon and eastern Nigeria. The western groove-toothed rat (Otomys occidentalis) is known only from the Gotel Mountains of eastern Nigeria and from Mount Oku and another nearby volcano in western Cameroon. Hartwig’s soft-furred mouse (Praomys hartwigi) is known only from a few localities in western Cameroon. The Bamenda shrew (Crocidura picea) is known only from the Bamenda Highlands and Mount Oku in western Cameroon. The Heim de Balsac’s forest shrew (Sylvisorex camerunensis) is found widely but patchily in the mountains of eastern Nigeria and western Cameroon. Eisentraut’s pipistrelle bat (Hypsugo eisentrauti) is likely confined to the montane forests of south-western Cameroon. There are records from four other countries, but it seems likely that these are misidentifications. Bannerman’s turaco (Tauraco bannermani) is largely confined to montane forest fragments within the Bamenda Highlands of western Cameroon, although small populations exist on nearby Mount Mbam as well as in two localities to the south-west. The Kupe bush-shrike (Chlorophoneus kupeensis) was long thought to be confined to a small area of forest on Mount Kupe in western Cameroon, but has since been discovered at a few other localities in both western Cameroon and eastern Nigeria. The small remaining population is highly threatened by habitat destruction. The green-breasted bush-shrike (Malaconotus gladiator) is a rare species found patchily in the highlands of western Cameroon and south-eastern Nigeria. The white-throated mountain babbler (Kupeornis gilberti) is confined to a few localities in western Cameroon and eastern Nigeria. Bannerman’s weaver (Ploceus bannermani) is a type of passerine bird found patchily in western Cameroon and south-eastern Nigeria. Several chameleons of the genus Trioceros are threatened by habitat destruction and overcollection for the international pet trade. Perret’s montane chameleon (T. perreti) is confined to the Manengouba and Bakossi mountains of western Cameroon. Pfeffer’s two-horned chameleon (T. pfefferi) is confined to a few montane forest remnants in western Cameroon. The serrated chameleon (T. serratus) is known from a few localities in western Cameroon. Wiedersheim’s chameleon (T. wiedersheimi) is known only from two small areas in western Cameroon and another in eastern Nigeria. The four-horned chameleon (T. quadricornis) is found in a few scattered localities in western Cameroon and south-eastern Nigeria. The southern four-horned chamelon (T. q. quadricornis) occurs on Mount Kupe, Mount Manengouba, and the Bakossi Mountains. The northern four-horned chameleon (T. q. gracilior) is known from the Bamboutos Mountains, Bamenda Highlands and Mbulu Hills

of western Cameroon, and from the Obudu Plateau in southeastern Nigeria. Paulian’s skink (Leptosiaphos pauliani) is known only from two localities in western Cameroon (Mount Bamboutos and Mount Kupe). Villiers’ toad (Sclerophrys villiersi) is confined to the mountains of western Cameroon, where it is threatened by habitat destruction and degradation. The Cameroon Wolterstorff toad (Wolterstorffina parvipalmata) is found widely but patchily in western Cameroon and south-eastern Nigeria, where it is threatened by habitat destruction and most likely chytridiomycosis. Several species of smalltongue toad (Werneria) are endemic to small areas of western Cameroon, where they are threatened by loss of habitat. Preuss’ smalltongue toad (W. preussi) is known only from Mount Cameroon and Mount Kupe. The submontane smalltongue toad (W. submontana) is known only from Mount Kupe and the Bakossi Mountains. Tandy’s smalltongue toad (W. tandyi) is known only from Mount Manengouba and the Rumpi Hills. Mertens’ smalltongue toad (W. mertensiana) is known from a few scattered localities. The Cameroon leaf-folding frog (Afrixalus lacteus) is known only from a few mountaintops in western Cameroon. Steindachner’s puddle frog (Phrynobatrachus steindachneri) and Jim Zimkus’ puddle frog (P. jimzimkusi) are both confined to a small area of western Cameroon and southeastern Nigeria, where they are threatened by loss of habitat and possibly chytridiomycosis. Perret’s water frog (Petropedetes perreti) is confined to a few localities in south-western Cameroon. Mertens’ egg frog (Leptodactylodon mertensi) is known only from four localities within the mountains of western Cameroon (the Bamileke Plateau, Mount Nlonako, and Mount Manengouba). Perret’s egg frog (L. perreti) is known from the higher peaks of western Cameroon. The Buea egg frog (L. bueanus) is known only from the eastern slopes of Mount Cameroon and from nearby Bimbia Hill in southwestern Cameroon. The ornate egg frog (L. ornatus) is divided into two subspecies, with the nominate form (L. o. ornatus) found on Mount Manengouba, Mount Nlonako, Mount Kupe, and from the Bonandam Hills and Ebonji Hills. All are threatened by loss of habitat and possibly chytridiomycosis. Perret’s screeching frog (Arthroleptis perreti) is known for certain only from Mount Manengouba in south-western Cameroon, but may occur as well on nearby peaks and in the Rumpi Hills. The Nkongsamba white-lipped frog (Amnirana asperrima) is known from a few highland areas of south-eastern Nigeria and south-western Cameroon. The decorated running frog (Kassina decorata) is known only from two widely separated areas of western Cameroon (the Bamileke Highlands and Mount Manengouba). Perret’s night frog (Astylosternus perreti) and Amiet’s night frog (A. ranoides) are both known only from a few localities in western Cameroon.

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Several species of long-fingered frog (Cardioglossa) are threatened by loss of habitat. The black long-fingered frog (Cardioglossa pulchra), Schiotz’s long-fingered frog (C. schioetzi), lovely long-fingered frog (C. venusta), Alsco longfingered frog (C. alsco), and Amiet’s long-fingered frog (C. melanogaster) are all found patchily in the mountains of western Cameroon and south-eastern Nigeria, where they are threatened by loss of habitat. The Western High Plateau The Western High Plateau (also known as the Western Highlands or the Bamenda Highlands) lies along the Cameroon Volcanic Line in western Cameroon (North-West, West, and South-West regions). It is ill-defined area of mountain ranges and ancient volcanoes characterized by high relief, cool temperatures, heavy rainfall, and savanna vegetation. The Bamenda forest shrew (Sylvisorex silvanorum) is known only from a few specimens collected from the Western High Plateau. The Manenguba shrew (Crocidura manengubae) is known from two or three mountaintop localities on the Western High Plateau. The Oku mouse shrew (Myosorex okuensis) is known from a few scattered localities in the Western High Plateau, including Mount Manenguba, Mount Oku, and Mount Lefo. The banded wattle-eye (Platysteira laticincta) is a type of passerine bird confined to forest fragments within the Bamenda Highlands. The Western High Plateau smalltongue toad (Werneria bambutensis) is confined to a few localities on the Western High Plateau. The Oku long-fingered frog (Cardioglossa oreas) is confined to the Western High Plateau between Mount Oku and the Bamboutos Mountains. The Oku Massif is a volcanic field located in centralwestern Cameroon (north-west region). It contains four major stratovolcanoes (Mount Oku, Mount Babanki, Mount Nyos, and Mount Nkambe) and three crater lakes (Lake Oku, Lake Nyos, and Lake Monoun). The Oku rat (Lamottemys okuensis) is confined to the montane rainforest atop Mount Oku, where it is threatened by deforestation. Dieterlen’s brush-furred mouse (Lophuromys dieterleni) is known only from a few specimens collected on Mount Oku. The Oku wood mouse (Hylomyscus grandis) is known only from a few specimens collected on the upper slopes of Mount Oku. Mittendorf’s striped grass mouse (Lemniscomys mittendorfi) is known only from a few specimens collected from a small area of grassland on Mount Oku. Chirio’s Wolterstorff toad (Wolterstorffina chirioi) is known only from the summit of Mount Oku. The Lake Oku puddle frog (Phrynobatrachus njiomock) is known only from a single forest fragment on Mount Oku near Lake Oku, a shallow crater lake, where it was last recorded in

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2010. It may possibly be extinct due to chytridiomycosis. The spiny puddle frog (P. chukuchuku) is known only from a few high-elevation localities in and around Mount Oku. The Oku clawed frog (Xenopus longipes) is confined to Lake Oku. The Oku caecilian (Crotaphatrema lamottei) is confined to Mount Oku. The Bamileke Plateau is located in central-western Cameroon (North-west region). Densely populated, most of its original forest cover has been lost. The Lefo brush-furred rat (Lophuromys eisentrauti) is known only from its original collection in montane forest on Mount Lefo in 1978. The Bamileke ornate egg frog (Leptodactylodon ornatus permaculatus) is confined to the southern and western slopes of the Bamileke Plateau. The Bambouto Massif is located in western Cameroon (West region). It is dominated by Mount Bambouto. Mire’s Wolterstorff toad (Wolterstorffina mirei) appears to be confined to the Bambouto Massif. The Bamboutos egg frog (Leptodactylodon axillaris) is known only from Mount Bamboutos. The Bakossi Mountains are located in western coastal Cameroon (south-west region). The highest peak in the range is Mount Kupe, which has been heavily deforested. Fortunately, other areas of cloud forest remain in the range, some of which are protected within the Bakossi Forest Reserve. Weiler’s gecko (Urocotyledon weileri) is a little-known species possibly confined to Mount Kupe. Dintelmann’s reed frog (Hyperolius dintelmanni) is known only from Mount Kupe and the Edib Hills. Wild’s egg frog (Leptodactylodon wildi) is known only from a small area of the Bakossi Mountains. The Adamawa Plateau The Adamawa Plateau (Massif de l’Adamaoua in French) is located in south-eastern Nigeria (Adamawa and Borno states), north-central Cameroon (Adamawa and North Cameroon regions), and western parts of the Central African Republic. The Djohong toad (Sclerophrys djohongensis) is known only from the Adamawa Plateau in north-central Cameroon, although it may also occur in adjacent areas of southeastern Nigeria. The Nganha night frog (Astylosternus nganhanus) is known only from five specimens collected in 1978 on Mount Nganha, Cameroon (Adamawa region). The Tchabal Mbabo caecilian (Crotaphatrema tchabalmbaboensis) is known only from three specimens collected in 1997 from Mount Tchabal Mbabo, Cameroon (Adamawa region). The Gotel Mountains The Gotel Mountains are located in south-eastern Nigeria (Taraba state) and north-central Cameroon (north-west region). They are partly protected by Gashaka Gumti National Park, Nigeria’s largest.

The Guineo-Congolian Region

The Gotel soft-furred mouse (Praomys obscurus) is confined to the Gotel Mountains in south-eastern Nigeria. The Rumpi Hills The Rumpi Hills are located about 80 km north of Mount Cameroon in western Cameroon (south-west region). It has suffered severe forest loss due to smallholder farming activities, logging, and human settlement. The Rumpi mouse shrew (Myosorex rumpii) is known only from a single specimen collected in 1967. Eisentraut’s four-horned chameleon (Trioceros quadricornis eisentrauti) is confined to the Rumpi Hills. The Jynx midwife frog (Alexteroon jynx) is known only from two localities on the eastern slopes of the Rumpi Hills. Mount Manengouba Mount Manengouba is located in south-western Cameroon (littoral region). The Manengouba puddle frog (Phrynobatrachus manengoubensis) is known only from a shallow crater lake at 2000 m on Mount Manengouba. The Nsoung long-fingered frog (Cardioglossa trifasciata) and the Manengouba Peak long-fingered frog (C. manengouba) are both confined to Mount Manengouba. The redbelly egg frog (Leptodactylodon erythrogaster) is confined to the south-eastern slopes of Mount Manengouba. Mount Cameroon Mount Cameroon is an active volcano that sits near the coast in western Cameroon (south-west region). Its forested slopes are fairly intact and partially protected within Mount Cameroon National Park. Burton’s groove-toothed rat (Otomys burtoni) is known only from Mount Cameroon. Eisentraut’s striped mouse (Hybomys badius) is confined to Mount Cameroon. Eisentraut’s shrew (Crocidura eisentrauti) is confined to the summit of Mount Cameroon. The Mount Cameroon forest shrew (Sylvisorex morio) is confined to the higher elevations of Mount Cameroon. The Mount Cameroon francolin (Pternistis camerunensis) is a partridge-like bird confined to the south-eastern and north-eastern slopes, where it has become quite scarce in recent years. The Mount Cameroon white-eye (Zosterops melanocephalus) is a small, warbler-like bird is confined to a narrow altitudinal belt around Mount Cameroon, within which it is common. Poliak’s killifish (Aphyosemion poliaki) is confined to brooks on the eastern slopes of Mount Cameroon.

The Nimba otter shrew (Micropotamogale lamottei) is a tiny, semi-aquatic species endemic to the small upland forest streams of the Nimba Range in Liberia, Guinea and Ivory Coast, and the Putu Range in Liberia. It appears to be generally uncommon. The white-eyed prinia (Schistolais leontica) is a type of passerine bird found patchily in submontane areas of the Guinea Highlands of Ivory Coast, Guinea, Liberia, and Sierra Leone. It is threatened by habitat destruction due to mining activity. The Guinea Highlands day gecko (Cnemaspis occidentalis) is confined to a few disjunct areas of montane forest, including the Nimba Range on the Liberia/Guinea border, the Loma Mountains of Sierra Leone, the Ziama region of Guinea, and Mount Tonkoui in Ivory Coast. It is threatened by logging and mining activities. The Nimba Range The Nimba Range is a narrow ridge extending for approximately 40 km on the border between Liberia, Guinea, and Ivory Coast. One of the most interesting of the West African equatorial massifs, its isolation has produced hundreds of endemic plants and animals. Whether this remarkable number is due to the fact that the massif served as a refuge for species that had a wider distribution during previous pluvial periods or because they, through isolation, have evolved from ancestral forms, it is very important for research into evolution. Unfortuately, its forests have been severely reduced by logging, fires, clearing for agriculture, and large-scale mining activities, and its wildlife intensively hunted. The Nimba viviparous toad (Nimbaphrynoides occidentalis) is known only from three small, non-connected montane grassland localities in the Nimba Range. The sites are only a few kilometres apart but are divided by unsuitable habitat. Mount Richard-Molard (also known simply as Mount Nimba) is located on the south-eastern Guinea/Ivory Coast border. It is the highest peak in the Nimba Range. Lamotte’s roundleaf bat (Hipposideros lamottei) is known only from Mount Richard-Molard. The Nimba screeching frog (Arthroleptis nimbaensis) is known only from its original collection on Mount RichardMolard in the mid-twentieth century. The Fouta Djallon Highlands The Fouta Djallon Highlands are a plateau region located in north-central Guinea and (marginally) south-eastern Senegal. The Fouta Djallon aquatic skink (Cophoscincopus senegalensis) is known only from small, fast-flowing streams in the Fouta Djallon Highlands.

The Guinea Highlands The Guinea Highlands is a densely forested mountainous plateau extending from central Guinea through northern Sierra Leone and Liberia to western Ivory Coast. It includes a number of distinct mountain ranges and plateaus.

Idanre Hill Idanre Hill is located in south-western Nigeria (Ondo state). The Ondo Forest day gecko (Cnemaspis petrodroma) is confined to Idanre Hill.

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Perret’s toad (Sclerophrys perreti) is confined to Idanre Hill, where it was long believed to be extinct until rediscovered in 2013. The Jos Plateau The Jos Plateau is located in central Nigeria (Plateau state). Fox’s shaggy rat (Dasymys foxi) is confined to the central Jos Plateau. The giant day gecko (Cnemaspis gigas) is confined to the Jos Plateau. Dunger’s skink (Leptosiaphos dungeri) is known for certain only from high-altitude savanna on the Jos Plateau, although another population on the Adamawa Plateau in Cameroon may also be attributable to this species. The Kigomo worm lizard (Cynisca kigomensis) is known only from a few specimens collected in the 1960s from the Jos Plateau. The Simandou Massif The Simandou Massif is a range of hills located in southeastern Guinea. The Simandou white-lipped frog (Amnirana fonensis) is known only from the Simandou Massif.

Miscellaneous Mountains and Highlands The South Cameroon Plateau is Cameroon’s dominant geographical features, covering as it does more than half of the country’s total area. It lies south of the Adamawa Plateau and south-east of the Cameroon Volcanic Line. The whitebelly egg frog (Leptodactylodon albiventris) is known for certain only from the western edge of the South Cameroon Plateau, where it is uncommon. The Crystal Mountains (Monts de Cristal in French) are a group of low mountains (or high hills) inland of the Atlantic coast of Equatorial Guinea, Gabon, the Republic of Congo, Democratic Republic of Congo, and Angola. They are actually the edge of the Woleu-Ntem Plateau against the coastal lowlands, and are characterized by a steep descent deeply cut by streams. The Crystal Mountains forest tree frog (Leptopelis crystallinoron) was long known only from a single specimen, until five more were collected in 2009 from Crystal Mountains National Park. The Chaillu Mountains are located on the border between southern Gabon and the Republic of Congo. The highest peaks are Mount Iboudji and Mount Mimongo. Wuendsch’s killifish (Aphyosemion wuendschi) is confined to a single, swampy rainforest creek of the Ngounie River, in the central Chaillu Mountains of Gabon. Mount Iboundji (Mont Iboundji in French) is located in south-central Gabon. The Iboundji smalltongue toad (Werneria iboundji) is known only from two specimens collected from a rocky waterfall.

Lowland Tropical Rainforests and Swamp Forests Most of Africa’s lowland rainforests are located within the Congo drainage basin between the Albertine Rift Valley in

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the east to the Atlantic Ocean, with additional patches occurring along coastal West Africa as far as Guinea. As a result of the damage done by man, the rainforests of Africa today cover less than a third of their former area. Nevertheless, about onethird of all the tropical rainforest in the world is in Africa. Western and Central African rainforests share many of the same plants and animals, suggesting that they may have once been connected in the past. The African forest elephant (Loxodonta cyclotis) is found discontinuously in the lowland rainforests of western Africa in Ivory Coast and Ghana, and in central Africa from Cameroon to the Democratic Republic of Congo. The largest stable population lives in Gabon. The species is threatened in many areas by bushmeat hunting and ivory poaching. The greater roundleaf bat (Hipposideros camerunensis) is known only from three widely spaced localities (western Cameroon, the eastern Democratic Republic of Congo, and the Kakamega Forest of western Kenya). The eastern wattled cuckoo-shrike (Lobotos oriolinus) is widely but patchily found in the lowland rainforests of central Africa, where it nevertheless appears to be very rare for reasons as yet unknown. Home’s hinge-back tortoise (Kinixys homeana) is fairly widespread in the lowland rainforests of western and central Africa, but has undergone considerable declines due to loss of habitat and overcollection for food, ‘traditional medicine’, and the international pet trade.

Lowland Guinean Rainforests and Swamp Forests The lowland Guinean rainforests and swamp forests are a discontinuous expanse of coastal tropical rainforest in West Africa. The Dahomey Gap, an area of savanna and dry forest in Togo and Benin, divides the Lower Guinean Forests from the Upper Guinean Forests to the west. The yellow-casqued hornbill (Ceratogymna elata) is found widely but patchily in West Africa from Senegal to Cameroon, where it is threatened mainly by hunting. Upper Guinean Lowland Rainforests and Swamp Forests The Upper Guinean lowland rainforests and swamp forests extend from central and southern Guinea and Sierra Leone through Liberia, southern Ivory Coast, and south-western Ghana, with isolated patches in Togo and north-western Benin. The region has suffered massive deforestation. The crescent-naped mangabey (Cercocebus lunulatus) is found in parts of Ivory Coast, south-western Burkina Faso, and south-western Ghana. It is threatened mainly by hunting and habitat destruction, and persecuted in some areas for crop-raiding. Two closely related and relatively large species of guenon (Cercopithecus) are endemic to Upper Guinean lowland forests, where they are threatened by hunting and habitat destruction. The Diana guenon (C. diana) is found in coastal south-eastern Guinea to Sierra Leone, Liberia, and southern Ivory Coast west of the Sassandra River. The Roloway guenon

The Guineo-Congolian Region

(C. roloway) was historically found across southern Ivory Coast and southern Ghana, roughly from the Sassandra River to the Pra River. Today the species survives only in a few scattered forest patches in central coastal and south-eastern Ivory Coast and south-western Ghana. Two colobus monkeys of the genus Colobus were historically widespread within the Upper Guinean lowland rainforests and gallery forests. During the nineteenth century these leafeating monkeys were nearly hunted into extinction for their beautiful fur, although today the main threat comes from loss of habitat and bushmeat hunting. The king colobus (C. polykomos) occurs in far western Africa from southern Senegal, Guinea-Bissau, Guinea, Sierra Leone, and Liberia east to the Sassandra River in Ivory Coast. The ursine colobus (C. vellerosus) is found in eastern Ivory Coast through southern Ghana, Togo, and Benin to south-western Nigeria. Miss Waldron’s red colobus (Piliocolobus waldronae) was originally found in south-eastern Ivory Coast and southwestern Ghana. Habitat destruction and hunting have long since eliminated it from most of this area, and there have been no confirmed sightings since 1978. After the announcement of the species’ probable extinction in 2000, however, new evidence (i.e. skins and a photograph) gave hope that a small number may yet survive between the Ehy Forest and Tanoé River in extreme south-eastern Ivory Coast. The Upper Guinea red colobus (P. badius) occurs in fragmented populations in south-western Ivory Coast, southern Guinea, Liberia, and Sierra Leone, where it is threatened by bushmeat hunting. The pygmy hippopotamus (Choeropsis liberiensis) historically ranged far more widely in equatorial Africa than it does today, although it appears to have always been rather rare. Considerably less aquatic than its more familiar cousin the common hippopotamus (Hippopotamus amphibius), it is a secretive inhabitant of lowland forests near rivers and swamps where, in recent years, whole populations have disappeared and become isolated. The Upper Guinea pygmy hippopotamus (C. l. liberiensis) is found in Ivory Coast, Guinea, Liberia, and Sierra Leone. Fortunately, it breeds fairly well in captivity, a guarantee that it will not become extinct. Bourlon’s genet (Genetta bourloni) is known only from a small number of specimens collected in southern Guinea, Liberia, Sierra Leone, and western Ivory Coast. The West African linsang (Poiana leightoni) is a weasellike arboreal carnivore known only from a few localities in south-western Ivory Coast and eastern Liberia. The Liberian mongoose (Liberiictis kuhni) occurs in Liberia, western Ivory Coast, and possibly southern Guinea, where it is heavily hunted. Baer’s wood mouse (Hylomyscus baeri) is known only from a few scattered localities in Ivory Coast, Ghana, Guinea, and Sierra Leone. The groove-toothed forest mouse (Leimacomys buettneri) is known only from two specimens collected in west-central Togo in 1890. Two expeditions in the 1990s failed to

rediscover the species, although local people were said to be familiar with it. Wimmer’s shrew (Crocidura wimmeri) is confined to a small area of south-western Ivory Coast. The rufous fishing-owl (Scotopelia ussheri) is found widely but patchily in Ivory Coast, Ghana, Guinea, northern Liberia, and Sierra Leone, where it is threatened by loss of habitat. The white-necked rockfowl (Picathartes gymnocephalus) has a highly fragmented distribution in Guinea, Sierra Leone, Liberia, Ivory Coast, and Ghana, where most of its breeding colonies are small and isolated. The yellow-bearded greenbul (Criniger olivaceus) is found widely but patchily in Sierra Leone, Liberia, south-eastern Guinea, southern Ivory Coast, and south-western Ghana, but is everywhere threatened by loss of habitat. The western wattled cuckoo-shrike (Lobotos lobatus) is found widely across eastern Sierra Leone, Liberia, southeastern Guinea, southern Ivory Coast, and south-western Ghana, but is everywhere threatened by loss of habitat. The Upper Guinean flycatcher (Melaenornis annamarulae) is found patchily in eastern Sierra Leone, Liberia, southeastern Guinea, south-western Ivory Coast, and south-central Ghana. Highly dependant upon lowland closed-canopy rainforest with large emergent trees, it is threatened by loss of habitat. Ballmann’s malimbe (Malimbus ballmanni) is a rare type of weaverbird known only from a few disjunct populations in eastern Sierra Leone, Liberia, southern Guinea, and western Ivory Coast. Necas’ chameleon (Chamaeleo necasi) is known only from a few localities in south-western Togo and southern Benin. Williams’ worm lizard (Cynisca williamsi) is known only from a single specimen collected in the early 1960s from westcentral Ghana. Mueller’s worm lizard (C. muelleri) is known only from its original collection in the late nineteenth century in south-western Ghana. Degrys’ worm lizard (C. degrysi) is known only from a single specimen collected in the 1940s from the unspecified locality ‘Sierra Leone’. Mann’s worm snake (Afrotyphlops manni) is known only from two specimens collected from two, widely separated localities (Mount Nimba, Guinea, and south-western Liberia). It is possible that one or both localities are incorrect. The Liberian blind snake (Letheobia leucostictus) is known only from a single specimen collected in the late nineteenth century from the unspecified locality ‘Liberia’. Broadley’s blind snake (Tricheilostoma broadleyi) is known only from two localities in south-central Ivory Coast. The Monogaga toad (Sclerophrys danielae) is known only from a small area of south-western coastal Ivory Coast, where it was last recorded in the late 1970s. The Taï toad (S. taiensis) is known only from two disjunct localities (Taï National Park in south-western Ivory Coast and the Gola Forest in southeastern Sierra Leone). Perret’s running frog (Kassina arboricola) is known only from a few localities over a relatively wide area of southern

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Ivory Coast and south-western Ghana. Lamotte’s running frog (K. lamottei) is known only from south-western Ivory Coast, but may also be found in eastern Liberia and southern Guinea. Both are threatened by loss of habitat. Several reed frogs of the genus Hyperolius are threatened by loss of habitat. The Nimba reed frog (H. nimbae) is known only from a small area of western Ivory Coast. It went unrecorded from 1967 until its rediscovery in 2010. The Ukami reed frog (H. torrentis) is confined to a few localities in eastern Ghana and western Togo. The Bobiri reed frog (H. bobirensis) is confined to three widely separated protected areas in southern Ghana. Laurent’s reed frog (H. laurenti) is known only from a few localities in south-eastern Ivory Coast and southwestern Ghana. The Ivory Coast reed frog (Morerella cyanophthalma) is known only from three widely scattered lowland rainforest localities in southern Ivory Coast. Langer’s screeching frog (Arthroleptis langeri) is known only from two small, disjunct localities, one on the northern Liberia/southern Guinea border and the other in eastern Liberia. Several species of puddle frog (Phrynobatrachus) are threatened by loss of habitat. Vogt’s puddle frog (P. vogti) is known only from its original collection in the 1920s in southwestern Ghana. The Accra puddle frog (P. albolabris) is known only from its original collection in 1938 from what is now the city of Accra, Ghana. Schiotz’s puddle frog (P. ghanensis) is confined to forest fragments in south-western Ghana and south-eastern Ivory Coast. The Ankasa puddle frog (P. intermedius) is known only from the Ankasa National Reserve in south-western Ghana. The ringed puddle frog (P. annulatus) is known only from three small, disjunct localities (the northern Liberia/southern Guinea border, south-western Ivory Coast, and south-western Ghana). The Taï Forest in south-western Ivory Coast is the single largest tract of undisturbed tropical rainforest in West Africa. While officially protected as Taï National Park, the survival of the forest has been seriously threatened by the activities of slash-and-burn farmers, poachers, timber companies, and illegal gold miners. The Taï hairy bush viper (Atheris hirsuta) is known only from its original collection in Taï National Park. The Taï reed frog (Hyperolius nienokouensis) is known only from Taï National Park and another locality 30 km to the north. The Taï puddle frog (Phrynobatrachus taiensis) is known only from a single locality within Taï National Park. Lower Guinean Lowland Rainforests and Swamp Forests The Lower Guinean lowland rainforests and swamp forests, as here defined, extend along the eastern coast of the Gulf of Guinea from south-eastern Benin through southern Nigeria and western Cameroon. The white-throated guenon (Cercopithecus erythrogaster) occurs in scattered populations in isolated lowland forest

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patches, where they have been intensively hunted for their beautiful fur. There are two subspecies. The red-bellied white-throated guenon (C. e. erythrogaster) is found in southern Benin, including the Lama Classified Forest, and possibly Togo as well. What remains of its habitat is generally protected by both legal and religious strictures. The black colobus (Colobus satanas) is divided into two subspecies. Leconte’s black colobus (C. s. anthracinus) is found in Equatorial Guinea, south-western Cameroon, and central and north-western Gabon, with possibly some additional small populations in northern Republic of Congo. It is threatened by hunting and habitat destruction. Wilson’s palm squirrel (Epixerus ebii wilsoni) from Cameroon, Congo, Equatorial Guinea, and Gabon is threatened by habitat destruction. The Campo-Ma’an fruit bat (Casinycteris campomaanensis) is known only from two specimens collected in the vicinity of Campo-Ma’an National Park, Cameroon (south region) and a possible third in neighbouring Nigeria. Bates’ weaver (Ploceus batesi) is a very rare type of passerine bird from southern Cameroon. The Lama Forest house gecko (Hemidactylus lamaensis) is known only from a single locality in southern Benin. Greenwell’s blind snake (Tricheilostoma greenwelli) is known only from specimens collected in the Ibadan University Zoological Gardens of western Nigeria (Oyo state). The Victoria caecilian (Herpele multiplicata) is known only from a now-lost type specimen collected in south-western Cameroon in 1912. Nigerian Lowland Rainforests The Nigerian lowland rainforests are located in south-western Nigeria. Very little of the original forest cover now remains. The Ibadan malimbe (Malimbus ibadanensis) is a type of small passerine bird that was originally discovered in 1951, when it was considered to be fairly common. Today it is confined to a few remnant forest localities in southwestern Nigeria. Niger Delta Swamp Forests The Niger delta lies directly on the Gulf of Guinea in southwestern Nigeria. Densely populated, virtually all of the original swamp forest is now gone. The Niger delta pygmy hippopotamus (Choeropsis liberiensis heslopi) was an isolated population that lived from the Niger delta east to the Cross River in Nigeria. There have been no reliable reports of it since 1945, and it appears to be extinct. The Nigerian white-throated guenon (Cercopithecus erythrogaster pococki) was long considered extinct until a small group was rediscovered in 1988. Although other populations have since been located within the Niger River delta, it remains highly threatened. The Niger delta red colobus (Piliocolobus epieni) is endemic to the swamp forests of the Niger delta, where surviving populations are now highly fragmented and concentrated

The Guineo-Congolian Region

Figure 3.3 Niger delta red colobus. (Credit: Daniel Giraud Elliot.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

in a small area in the south-eastern part of their original range. It is not found in any protected areas. Cross-Niger Transition Forests The Cross-Niger transition forests are located in the low hills of south-eastern Nigeria (Abia, Akwa Ibom, Anambra, Ebonyi, and Imo states), between the Niger River on the west and the Cross River on the east. Historically covered by a mixture of tropical forest and savanna woodland, the region is today one of the most densely populated in Africa and most of the forest is gone. Sclater’s guenon (Cercopithecus sclateri) is confined to fragmented and degraded forest remants in southern Nigeria from the eastern Niger delta east to the Cross River. Gans’ worm lizard (Cynisca gansi) is known only from a single abandoned forest clearing in south-eastern Nigeria, where it may already be extinct. Cross-Sanaga Coastal Forests The Cross-Sanaga coastal forests are located in south-eastern Nigeria (Cross River state) and western Cameroon, between the Cross and Sanaga rivers and up to 300 km inland from the

Atlantic coast. Important protected areas include Cross River National Park and Korup National Park. The Cross River gorilla (Gorilla gorilla diehli) is confined to remote forests on rough terrain along the Nigeria–Cameroon border, where it is separated from other gorilla populations by the Sanaga River. Although the subspecies has been known to science since the early twentieth century, little attention was paid to its conservation status until the late 1980s. Surveys undertaken between 1990 and 2005 estimated that only 250–300 still survived. The Nigeria–Cameroon chimpanzee (Pan troglodytes ellioti) is the most threatened subspecies of chimpanzee, being found, like the Cross River gorilla, only in the rainforests of the Cameroon/Nigeria border area west of the Sanaga River. Historically it was probably widespread in southern Nigeria as far as the Niger River, although today it is confined to a few isolated forest fragments where, in 2011, the total population was estimated at no more than 6000–9000. Monkeys of the genus Mandrillus are short-tailed and notable for the adult males, which attain enormous sizes and are brightly coloured. Both species are threatened by hunting and loss of habitat. The drill (M. leucophaeus), which is divided into two subspecies, is one of the most endangered monkeys in Africa. The mainland drill (M. l. leucophaeus) is found only between the Cross and Sanaga river valleys in south-eastern Nigeria and western and south-western Cameroon. The total population is thought to be between 3000 and 8000. Animals recovered from illegal capture are held in semi-captivity at the Drill Rehabilitation and Breeding Centre in Nigeria, and there is an additional international captive population. The rusty-mantled crested mangabey (Lophocebus osmani) is a found disjunctly in south-eastern Nigeria and west-central Cameroon. The russet-eared guenon (Cercopithecus erythrotis) is divided into two subspecies. The Cross River russet-eared guenon (C. e. camerunensis) is found from south-eastern Nigeria to just south of the Sanaga River in Cameroon, where it is threatened by hunting and habitat destruction. Preuss’ red colobus (Piliocolobus preussi) is a highly threatened species confined to the Cross River region. The largest and most important population occurs in Korup National Park and surrounding forests of south-western Cameroon, and in Cross River National Park in adjacent south-eastern Nigeria. Historically it was also found in the area of the Ebo Forest, although recent surveys have failed to detect it. It is heavily hunted for its meat, even within protected areas. Schneider’s leaf-folding frog (Afrixalus schneideri) is known only from its original collection in the late nineteenth century in what is now the city of Douala, western Cameroon. Lindholm’s leaf-folding frog (A. lindholmi) is similarly known only from a single specimen collected in the early twentieth century from coastal western Cameroon. The Nlonako screeching frog (Arthroleptis nlonakoensis) is known only from a few areas of rainforest in western Cameroon (littoral region), including the Ebo Forest.

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The Makamunu Assumbo caecilian (Idiocranium russeli), one of the smallest of all caecilians, is known only from two localities in western Cameroon, but may occur more widely.

Lowland Congolian Rainforests and Swamp Forests The Congolian forests are a broad belt of lowland rainforests and swamp forests extending across the basin of the Congo River and its tributaries in Central Africa in south-eastern Cameroon, Gabon, Equatorial Guinea, north-western Angola (Cabinda), the Republic of the Congo, northern and central Democratic Republic of Congo, and portions of southern and south-western Central African Republic. All told, they represent the world’s second largest tropical forest (after the Amazon), roughly a quarter of what remains in the world. The okapi (Okapia johnstoni), a large but secretive forest-dwelling relative of the giraffe, went unknown to science until 1901. It is still found over a wide area of central, northern and eastern Democratic Republic of Congo north and east of the Congo River, as well as in a smaller region west and to the south of it. Long hunted by local pygmies, since about 1980 expansion of settlement and deforestation have eliminated the species from many of its former strongholds, as for instance the southern and eastern parts of the Ituri Forest. The western lowland gorilla (Gorilla gorilla gorilla) is the most widespread and abundant gorilla subspecies, being found over a large area of tropical forest and swamps that includes parts of Cameroon, Equatorial Guinea, Gabon, Nigeria, the Republic of Congo, the Democratic Republic of Congo, and the Cabinda exclave of Angola. The actual range can be defined as extending from the Sanaga River in the north-west and along the northern forest–savanna boundary to the Oubangui River, south to the Congo River, and west to the Atlantic coast. The total population is still relatively high (roughly between 300,000 and 350,000) but threatened by a number of factors including loss of habitat, poaching and disease. With very few historical exceptions this form is the only type of gorilla to be found in captivity. The Central African chimpanzee (Pan troglodytes troglodytes) lives in areas that are the least disturbed by humans overall, mainly in Gabon, Cameroon, and the Republic of Congo, but with smaller populations in the Central African Republic, Equatorial Guinea, the Cabinda exclave of Angola, south-eastern Nigeria, and possibly the coastal extension of the Democratic Republic of the Congo. Its range can be broadly defined as the area between the Sanaga River and the Congo/ Oubangui. The total population is likely between 47,000 and 78,000, although possibly as high as 140,000. The bonobo or pygmy chimpanzee (Pan paniscus) can be distinguished by its relatively long legs, pink lips, dark face, and certain features of the hair. It has a discontinuous range in the low-lying basin of the Democratic Republic of the Congo south of the Congo River, where it has long been isolated from other chimpanzee populations. While still relatively abundant it, too, is threatened by poaching both for meat as well as for

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supposed ‘medicinal’ purposes, as well as by habitat destruction and infectious disease. Several species of crested mangabey (Lophocebus) are threatened by habitat destruction and hunting. The Ugandan crested mangabey (L. ugandae) is confined to a few areas in Uganda and north-western Tanzania. The black crested mangabey (L. aterrimus) is found in the Democratic Republic of Congo south and west of the Congo–Lualaba drainage. Opdenbosch’s black crested mangabey (L. opdenoschi) is found in south-western Democratic Republic of Congo and northern Angola. The grey-cheeked crested mangabey (L. albigena) is found in southern Cameroon, south-western Central African Republic, Equatorial Guinea, Gabon, and the Republic of Congo. Johnston’s crested mangabey (L. johnstoni) is found in the Democratic Republic of Congo, western Rwanda, western Burundi, and possibly South Sudan. The golden-bellied mangabey (Cercocebus chrysogaster) is only known from a few localities in south-central and western Democratic Republic of Congo, where it is threatened by bushmeat hunting. The sun-tailed monkey (Allochrocebus solatus) is confined to central Gabon, primarily within Lopé National Park and the unprotected Forêt des Abeilles. It is threatened mainly by bushmeat hunting. Several species and subspecies of guenon (Cercopithecus) are threatened by bushmeat hunting. The Lesula guenon (C. lomamiensis) is found in central Democratic Republic of Congo between the Lomami and Tshuapa rivers. The Dundo diademed guenon (C. mitis mitis) is confined to western coastal Angola. The black-footed crowned guenon (C. pogonias nigripes) is found in Equatorial Guinea, western Gabon, and the Republic of Congo. Gray’s crowned guenon (C. p. grayi) occurs in southern Cameroon, south-western Central African Republic, north-western Democratic Republic of Congo, north-eastern Gabon, northern Republic of Congo, and possibly northern Angola (Cabinda). The white-nosed moustached guenon (C. cephus ngottoensis) is found in south-western Central African Republic, northern Republic of Congo, and possibly north-western Democratic Republic of Congo. The grey-tailed moustached guenon (C. c. cephodes) is found in western Gabon and western Republic of Congo. Curry’s bat (Glauconycteris curryae) is a little-known species from southern Cameroon and north-western Democratic Republic of Congo. Stevart’s egg frog (Leptodactylodon stevarti) is known for certain only from a small area of north-western Gabon, but may also occur in adjacent areas of Equatorial Guinea. The speckled egg frog (L. ventrimarmoratus) is found in southwestern Cameroon, south of the Sanaga River. Both are threatened by loss of habitat. Lowland Congolian Terra Firma Forests Terra firma literally means ‘firm earth’, and refers to rainforest that is not inundated by flooded rivers.

The Guineo-Congolian Region

The mandrill (Mandrillus sphinx), the world’s largest monkey, lives in the rainforests of west-central Africa south of the Sanaga River in Cameroon, Congo, Equatorial Guinea, and Gabon. Current numbers are unknown, but it has undoubtedly declined in recent years. It is generally rare and has been locally exterminated in places. The Tshuapa red colobus (Piliocolobus tholloni) is found patchily in the forests of the Democratic Republic of Congo south of the Congo River and west of the Lomami. Like other members of its genus, it has been greatly impacted by bushmeat hunting. Du Chaillu’s striped squirrel (Funisciurus duchaillui) is known only from a small area of central Gabon between the Ogooué River and Massif du Chaillu. Schouteden’s thicket rat (Thamnomys schoutedeni) is known only from two widely separated localities in the eastern Democratic Republic of Congo. The velvet African climbing mouse (Dendroprionomys rousseloti) is known only from a few specimens collected in the Zoological Garden of Brazzaville, in south-eastern Republic of Congo. The greater Congo shrew (Congosorex polli) is known only from a single specimen collected in the 1950s at Lubondai, in southern Democratic Republic of Congo. The lesser forest shrew (Sylvisorex oriundus) is known only from two localities in north-eastern Democratic Republic of Congo. Dieter’s mouse-eared bat (Myotis dieteri) is known only from a single cave in southern Congo. Two species of horseshoe bat (Rhinolophus) are threatened mainly by disturbance of their cave-roosts. The forest horseshoe bat (R. silvestris) is known only from three localities in eastern Gabon and another in the Republic of Congo. Adam’s horseshoe bat (R. adami) is known only from a few specimens collected from Kimanika Cave, southern Republic of Congo (Kouilou department), in 1968. Glen’s wattled bat (Glauconycteris gleni) is known only from two widely spaced localities (Cameroon and western Uganda). Machado’s butterfly bat (G. machadoi) is known only from a single locality in east-central Angola. The Congo peafowl (Afropavo congensis) still occurs over a wide area of central Democratic Republic of Congo, where it went unknown to science until 1936. It seems to be everywhere rare, with much of its range unpopulated. The Angolan greenbul (Phyllastrephus albigularis viridiceps) is known only from two small areas in north-western Angola (Cuanza Norte and Cuanza Sul provinces). The golden-naped weaver (Ploceus aureonucha) is confined to north-western Democratic Republic of Congo and adjacent areas of western Uganda. The Congolian eremomela (Eremomela turneri kalindei) is a type of passerine bird known from a few scattered localities in east-central Democratic Republic of Congo. The Urungu toad (Sclerophrys urunguensis) is known only from an area of remnant rainforest near the southern end of Lake Tanganyika, in the Tanzania/Zambia border area.

Blanc’s egg frog (Leptodactylodon blanci) is known only from specimens collected in Lopé National Park, central Gabon, and from Moukalaba-Doudou National Park in south-western Gabon. It is likely to also be found in a few intervening localities. The Lomami screeching frog (Arthroleptis phrynoides) is known only from a single locality in central Democratic Republic of Congo. The Cambondo screeching frog (A. carquejai) is known only from a small area of north-western Angola (Cuanza Norte province). The Ituri Rainforest is an area of lowland rainforest located in north-eastern Democratic Republic of Congo (Ituri province). First traversed by Henry Morton Stanley in 1887, it is notable for being home to the Mbuti pygmies, a huntergatherer people. About one-fifth of the forest is protected within the Okapi Wildlife Reserve, a World Heritage Site. The yellow-legged weaver (Ploceus flavipes) is a rare bird species known only from the central and eastern Ituri Forest. The Ituri reed frog (Hyperolius ferrugineus) is known only from a single locality in the Ituri Forest. Loveridge’s screeching frog (Arthroleptis loveridgei) is known only from two specimens collected in the Ituri Forest in 1925. The Kakamega Forest is located in south-western Kenya near the border with Uganda. Kenya’s only area of tropical rainforest, it is a last remnant of the ancient Guineo-Congolian rainforest that once spanned the continent. The Kakamega blind snake (Afrotyphlops kaimosae) is known only from a single specimen collected in the Kakamega Forest in 1935. Mackay’s forest tree frog (Leptopelis mackayi) is known only from Kakamega Forest and its vicinity. Lowland Congolian Swamp Forests Located in both the Republic of the Congo and in the Democratic Republic of the Congo, the lowland Congolian swamp forests form one of the largest continuous areas of freshwater swamp forest in the world. They consist of flooded forest with a high canopy, dense undergrowth, and a muddy floor. Still little-disturbed by human interference due to their impenetrability, they remain almost unstudied. The dryas guenon (Cercopithecus dryas) is an extremely rare species known only from a few locations in the Lomela and Wamba regions in central Democratic Republic of Congo. The remaining population (less than 200) is somewhat protected within the community-managed Kokolopori Reserve. Bouvier’s red colobus (Piliocolobus bouvieri) is known from an area of swampy forest in west-central Republic of Congo. The species had not been reported since the 1970s, and was thought to be extinct prior to its rediscovery in Ntokou-Pikounda National Park in 2015. Nahan’s partridge (Ptilopachus nahani) is known only from a few localities in eastern Democratic Republic of Congo. The Tumba tree frog (Cryptothylax minutus) is known only from the Lake Tumba area of west-central Democratic Republic of Congo.

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The Afrotropical Realm

Forest–Savanna Mosaic Forest–savanna mosaic is a transitionary bioregion between the moist lowland rainforests of Equatorial Africa and the drier savannas and open woodlands to the north, south, and east. Trevor’s free-tailed bat (Mops trevori) is known only from a few specimens collected over a very wide area of West and Central Africa.

Guinean Forest–Savanna Mosaic The Guinean forest–savanna mosaic includes the transitional areas between the lowland Guinean rainforests of West Africa and of the West Sudanian savanna. It extends from GuineaBissau and Guinea in the west, through central Ivory Coast, south-eastern Ghana, southern Togo, southern Benin, and south-central Nigeria to the Cameroon Highlands of northern Cameroon. In the Dahomey Gap the forest–savanna mosaic extends to the coast, thereby separating the Upper and Lower Guinean forests. Walter’s duiker (Philantomba walteri) is known from a few dozen skulls and carcasses taken from bushmeat markets in Benin, Togo, and southern Nigeria. They are believed to have originated from the Dahomey Gap. Jackson’s fat mouse (Steatomys jacksoni) is known only from two localities (west-central Ghana and south-western Nigeria). While likely to be found in intervening areas, in particular the Penjari Biosphere Reserve in Benin, the species is hunted and appears on occasion in ‘medicinal’ markets. Bouet’s spiny-toed lizard (Acanthodactylus boueti) is known only from a few specimens collected over a relatively wide area of West Africa (south-western Ghana, northern Togo, central Benin, and west-central Nigeria). The ornate long-tailed lizard (Latastia ornata) is known only from a single specimen collected in 1938 from northwestern Guinea-Bissau. The Kérouané mabuya skink (Trachylepis keroanensis) is known only from a small area of eastern Guinea. The Cassine worm lizard (Cynisca oligopholis) is known only from a small area of north-western Guinea and southern GuineaBissau. The Ivory Coast worm lizard (C. rouxae) is known only from two widely separated localities in north-eastern and central Ivory Coast. Krauss’ worm lizard (C. kraussi) is known only from a single locality in south-central Ghana. The Mokwa worm lizard (Baikia africana) is known only from a few specimens collected from a single locality in western Nigeria (Niger state). Dunger’s file snake (Mehelya egbensis) is known only from a single specimen collected in the 1960s from westcentral Nigeria. The beautiful screeching frog (Arthroleptis formosus) is known only from a small area of western Guinea.

Congolian Forest–Savanna Mosaic The Congolian forest–savanna mosaic includes the transitional areas between the lowland Congolian rainforests of Central

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Africa and the East Sudanian savanna to the north, as well as to the drier areas to the east and south. It encompasses portions of Cameroon, Democratic Republic of Congo, the Central African Republic, southern Gabon, southern Republic of Congo, north-western Angola, and south-western South Sudan. Pousargues’ mongoose (Dologale dybowskii) is known only from a small number of specimens collected in eastern Central Africa Republic, northern Democratic Republic of Congo, Uganda, and southern South Sudan. The Lunda rope squirrel (Funisciurus bayonii) is known only from south-western Democratic Republic of Congo and north-eastern Angola. Dollman’s tree mouse (Prionomys batesi) is known only from a few specimens collected from south-central Cameroon, south-western Central African Republic, and north-western Republic of Congo. The Gounda mouse (Mus goundae) and the Oubangui mouse (M. oubanguii) are each known only from a single locality in the Central African Republic. The fuscous shrew (Crocidura polia) is known only from its original collection in the north-eastern Democratic Republic of Congo in the early twentieth century. Hayman’s dwarf epauletted fruit bat (Micropteropus intermedius) is a rare species known only from four localities in northern Angola and southern Democratic Republic of Congo. The lesser Angolan epauletted fruit bat (Epomophorus grandis) is known only from three specimens collected in the mid-twentienth century from north-eastern Angola and southwestern Republic of Congo. The Niangara free-tailed bat (Mops niangarae) is known only from a single specimen collected in 1917 from northeastern Democratic Republic of Congo. The black-lored waxbill (Estrilda nigriloris) is a type of finch restricted to the area of the Lualaba River and Lake Upemba in the southern Democratic Republic of Congo. It was last recorded in 1950. Makolowode’s house gecko (Hemidactylus makolowodei) is known only from a single locality in western Cameroon. Jacobsen’s reed frog (Hyperolius jacobseni) is known only from a single locality in southern Central African Republic.

Lakes, Rivers, and Marshes Among the more important lakes, rivers, and marshes within the Guineo-Congolian Region are Lake Tanganyika and Lake Albert, along with the Congo, Volta, and Niger river systems. The Congo clawless otter (Aonyx congicus) is found in the wetland areas of Central Africa, where it is heavily hunted for its valuable pelt. The spotted-necked otter (Hydrictis maculicollis) is widespread and relatively common in the larger lakes and rivers of sub-Saharan Africa, but for some reason is often absent from what appear to be suitable habitats, particularly in East Africa and in the Zambezi below Victoria Falls. No doubt habitat

The Guineo-Congolian Region

destruction, pollution, and hunting have taken a toll on the species. The aquatic genet (Genetta piscivora) from the forests of the central and eastern Congo has long been considered among the rarest of African carnivores. Known mainly from museum specimens, it has not been reported since the late 1970s. The savanna swamp shrew (Crocidura longipes) is known only from two small and isolated swamps near Lake Kainji, in north-western Nigeria (Kebbi state). The West African slender-snouted crocodile (Mecistops cataphractus) occurs discontinuously from southern Senegal to Cameroon. The Central African slender-snouted crocodile (M. leptorhynchus) is found from Cameroon through the Congo Basin to South Sudan and south to northern Angola and western Tanzania. Both are seriously threatened by loss of habitat and bushmeat hunting. The African dwarf crocodile (Osteolaemus tetraspis) is the smallest living crocodile species. Two rather ill-defined subspecies are found in wetland areas across the Guineo-Congolian Region. The nominate form (O. t. tetraspis) historically ranged from Senegal to Uganda, while the Congo dwarf crocodile (O. t. osborni) is found in the Congo Basin south to northern Angola. Both have disappeared in many areas due to hunting and habitat destruction. The Nubian flapshell turtle (Cyclanorbis elegans) is a large and distinctive species that formerly occurred across Western and Central Africa, but which had not been reported in recent decades. Populations have recently been discovered, however, along small tributaries of the White Nile in South Sudan, where they are seriously threatened by collection for food. The Senegal flapshell turtle (C. senegalensis) is found in three main disjunct populations across northern-central Africa: one extending from Senegal to western Nigeria; another from Lake Chad to northern Central African Republic; and a third in Sudan, South Sudan, and western Ethiopia. The species is declining everywhere but particularly in West Africa due to overcollection and loss of habitat. The ocellated water snake (Hydraethiops laevis) is known only from a few specimens collected in Cameroon and Gabon. The banded toad (Sclerophrys vittata) is a little-known species confined to marshy areas in Uganda. The Gbanga forest tree frog (Leptopelis bequaerti) is known only from two widely separated localities in Liberia. The Omaniundu reed frog (Hyperolius sankuruensis) is known only from swamps and marshes in south-central Democratic Republic of Congo. First discovered in 1979, it was not seen again until 2010. Rochbrune’s reed frog (H. rhizophilus) and the Cabinda reed frog (H. maestus) are both known only from the Cabinda enclave of northwestern Angola. The slippery frogs (Conraua) inhabit swift-moving rivers within the rainforests of West Africa. The goliath or giant slippery frog (C. goliath) is the largest frog on Earth, being about the size of housecat. Found only in south-western

Cameroon and Equatorial Guinea, its numbers have dwindled due to overcollection for food and the pet trade. The Cameroon slippery frog (C. robusta) from Cameroon and Nigeria, Togo slippery frog (C. derooi) from the Togo-Volta Highlands. and Allen’s slippery frog (C. alleni) from northern Sierra Leone, southern Guinea, Liberia. and south-western Ivory Coast are all similarly threatened. The Efulen water frog (Petropedetes palmipes) is known only from a few localities in south-western Cameroon, Equatorial Guinea. and north-western Gabon. Ahl’s puddle frog (Phrynobatrachus brevipalmatus) is known only from its original collection in the early twentieth century in what is now the city of Luanda, north-western Angola. The Yambata puddle frog (P. giorgii) is known only from its original collection in the early twentieth century from somewhere in north-western Democratic Republic of Congo. The Kakamega puddle frog (P. kakamikro) is known only from a single locality in south-western Kenya. Amiet’s river frog (Amietia chapini) is known only from a small area of east-central Democratic Republic of Congo. The Fopouanga night frog (Astylosternus fallax), Apouh night frog (A. schioetzi). and Laurent’s night frog (A. laurenti) are all confined to a few small areas of western and southwestern Cameroon, where they are threatened by loss of habitat and overcollection for food. Fea’s dwarf clawed frog (Hymenochirus feae) is known only from its original collection from the central coast of Gabon in the early twentieth century. Boulenger’s dwarf clawed frog (H. boulengeri) is known only from two localities in north-eastern Democratic Republic of Congo. The Ugandan clawed frog (Xenopus ruwenzoriensis) is known only from two localities in western Uganda. Two species of stingray (Fontitrygon) are threatened by overfishing and accidental by-catch. The pincushion stingray (F. ukpam) is found discontinuously in rivers and lakes from Cameroon and Nigeria to Gabon and the Congo River drainage. The smooth stingray (F. garouaensis) is confined to the Niger/Benue, Cross. and Sanaga rivers, where it has declined or disappeared in many areas. The redbelly tilapia (Tilapia busumana) is a type of cichlid confined to a few river drainages and lakes in south-western Ghana and south-eastern Ivory Coast, where it is threatened by habitat degradation and pollution. The clown tilapia (Coelotilapia joka) is known only from the lower courses of the Jong and Waanje rivers in coastal Sierra Leone, but may occur in western Liberia as well. Holly’s tilapia (Coptodon camerunensis) is confined to the Meme, Mungo, and Wouri rivers of western Cameroon, where it is threatened by sedimentation and pollution. The discolor tilapia (C. discolor) is confined to a few lakes and rivers in southern Ghana and south-eastern Ivory Coast. Linke’s cichlid (Chromidotilapia linkei) is confined to the Mungo and Wouri rivers of western Cameroon. The elongated cichlid (C. elongata) is known only from a few rivers and

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creeks within the Dimonika Biosphere Reserve, southern Republic of Congo. The short-beaked cichlid (Parananochromis brevirostris) is known only from a few scattered localities in the Ivindo and Ntem river drainages of eastern Gabon and south-eastern Cameroon and from the Ogooué River drainage of western Gabon. Several cichlids of the genus Benitochromis are threatened by sedimentation and pollution. Lamboj’s cichlid (B. conjunctus) is confined to rivers and creeks around Mount Cameroon and the middle Mungo drainage in western Cameroon. Finley’s cichlid (B. finleyi) is confined to the Mungo and Meme river drainages of western Cameroon. The black-bellied cichlid (B. nigrodorsalis) is confined to an ill-defined area of western Cameroon. Ufermann’s cichlid (B. ufermanni) is known only from a few rivers and creeks in Korup National Park, western Cameroon. The Rio Muni cichlid (B. riomuniensis) is confined to south-western Cameroon and Equatoria Guinea. Bates’ cichlid (B. batesii) is found in western Cameroon, Equatorial Guinea, and Gabon. The graceful barb (Barboides gracilis) occurs patchily in small, slow-moving forest rivers in the coastal lowlands of West Africa, where it is threatened by loss of habitat. The red-eyed cichlid (Haplochromis erythromaculatus) is confined to Lake Burera, Lake Ruhondo, and the Mukungwa River in north-western Rwanda, where it is threatened by habitat destruction and overfishing. A great many barbs of the genus Enteromius are threatened by overfishing, pollution, and habitat destruction. The Nimba barb (E. eburneensis) is confined to streams and smaller rivers within the Nimba Range of western Ivory Coast, southern Guinea, and northern Liberia. Daget’s barb (E. dialonensis) is known from the upper Gambia, upper Niger, and upper Senegal River drainages of Guinea and eastern Senegal. Hopson’s barb (E. bawkuensis) is known only from the White Volta on the Ghana/Burkina Faso border and from the Sokoto River, a tributary of the Niger in north-western Nigeria. The Niokolo barb (E. niokoloensis) is known from the upper and middle Gambia River in Senegal and Guinea, as well as from the Bafing River in the upper Senegal River drainage of Mali. Raimbault’s barb (E. raimbaulti) is known from the Bafing River (a tributary of the Senegal River) and from the Niandan River (a tributary of the upper Niger River) in Guinea. Steindachner’s barb (E. liberiensis) is known only from three suspected localities in Liberia and Sierra Leone. Alicia’s barb (E. aliciae) is known only from the Saint John River in Liberia and from the Saint Paul River in Guinea. Bourdarie’s barb (E. bourdariei) is known only from Lake Monoun and the Noun River in western Cameroon. Stauch’s barb (E. stauchi) is known only from a few localities in the Loeme and Kouilou-Niari rivers drainages of south-western Congo. The silvan barb (E. sylvaticus) is known from the Iguidi River in southern Benin and from the lower Niger River delta in southern Nigeria. Hugueny’s barb (E. huguenyi) is known only from the Saint Paul River drainage in south-

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eastern Guinea and from the Lofa River drainage in northwestern Liberia. Salesse’s barb (E. salessei) is confined to rivers ands streams in northern Guinea and northern Sierra Leone. Boulenger’s barb (E. taeniurus) is confined to the Dja, Kribi, and Ntem rivers of southern Cameroon. The Kribi barb (E. tegulifer) is known only from its original collection in the 1930s from south-western Cameroon. Walker’s barb (E. walkeri) is known only from a few rivers in southern Ghana and south-eastern Ivory Coast. Thy’s barb (E. thysi) is known from coastal rivers in western Cameroon and Bioko. Tiekoro’s barb (E. tiekoroi) is known from the Konkouré, Kolenté, and Little Scarcies rivers of Guinea and Sierra Leone. Collart’s barb (E. collarti) is confined to the Chiloango and Kouilou-Niari rivers of south-western Republic of Congo. Several species of yellowfish (Labeobarbus) are threatened by sedimentation, pollution, overfishing, and introduced species. The Mungo yellowfish (L. mungoensis) is confined to the rivers of west-central Cameroon. Royli’s yellowfish (L. roylii) is confined to the Chiloango and Kouilou rivers of northern Angola and south-western Republic of Congo. Petitjean’s yellowfish (L. petitjeani) is confined to rivers in Guinea. The Tanganyika yellowfish (L. leleupanus) is confined to the northern part of Lake Tanganyika and to the Ruzizi River. Claudina’s yellowfish (L. claudinae) is confined to a few rivers in Rwanda and Burundi. Alluaud’s labeo (Labeo alluaudi) is confined to a few rivers in Liberia, Ivory Coast ,and possibly Sierra Leone, where it is threatened by overfishing and habitat degradation. The Sierra Leone trout barb (Raiamas scarciensis) is known only from the Little Scarcies River and the Waanje River of Sierra Leone and Guinea. The Sierra Leone minnow (Leptocypris taiaensis) is known from the Taia, Little Scarcies, and Waanje rivers of Sierra Leone. The denticle herring (Denticeps clupeoides) is confined to a few rivers in eastern Benin, southern Nigeria, and northwestern Cameroon. Gerlach’s rasbora (Rasbora gerlachi) is known only from its initial collection in the 1920s from a single, undefined locality in Cameroon. The species is so poorly known that even its generic assignment is in question. Ashby Smith’s climbing gourami (Ctenopoma ashbysmithi) is known only from an unnamed marsh near Kindu in east-central Democratic Republic of Congo. Nichols’ climbing gourami (Microctenopoma lineatum) is known only from a few localities in the Central African Republic and northern Democratic Republic of Congo. Axelrod’s tetra (Neolebias axelrodi) is known only from a few specimens collected from southern Nigeria and an unknown locality in Benin. Roloff’s dwarf characin (Ladigesia roloffi) is known for certain only from a single rainforest brook in south-central Sierra Leone. The Kribi citharine (Nannocharax micros) is known only from a single locality in south-western Cameroon.

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The La Passa tetra (Brachypetersius notospilus) is known only from a single locality in the Ogooué River in southwestern Congo and another in the Kouilou River of southern Gabon. Boulenger’s tetra (Neolebias spilotaenia) is known only from the Kouilou, Chiloango, and Loeme rivers in southeastern Congo. Smykala’s tetra (Alestopetersius smykalai) is confined to the lower Niger and Ogun rivers in southern Nigeria. The Niger tetra (Arnoldichthys spilopterus) is confined to the lower Ogun and lower Niger rivers in southern Niger, where it is threatened by habitat destruction. The Adonis tetra (Lepidarchus adonis) is confined to a few small coastal basins in southern Ivory Coast and south-western Ghana, where it is mainly threatened by pollution. Derham’s robber tetra (Brycinus derhami) is confined to a few small river drainages in south-western Ivory Coast. The short robber tetra (B. brevis) is known only from southern Ghana and southern Nigeria. Good’s goby (Ebomegobius goodi) is known only from a single specimen collected from a brackish water stream near Kribi, in coastal Cameroon, in the 1940s. A number of species and subspecies of panchax (Epiplatys) are threatened by habitat destruction and mining pollution. Many are popular aquarium fish. The Belefuanai panchax (E. coccinatus) is known only from a single rainforest creek in northern Liberia. Etzel’s panchax (E. etzeli) is confined to a small area in and around Ono Lagoon and the Hébé River of south-eastern Ivory Coast. The Port Loko panchax (E. lokoensis) is known for certain only from swamps and streams of western Sierra Leone. Hildegard’s panchax (E. hildegardae) is known only from a small area of south-eastern Guinea within the upper drainage of the Saint John/Mani and Saint Paul/Oulé rivers. The Njala panchax (E. njalaensis) is confined to small rainforest rivers and creeks in south-eastern Sierra Leone. Roloff’s panchax (E. roloffi) is known only from two small areas of north-western Liberia. The Biafra panchax (E. biafranus) is found in small brooks and swamps within two disjunct areas of southern coastal Nigeria. The red-spotted panchax (E. lamottei) occurs in a number of river systems within southeastern Guinea and north-western Liberia. Olbrechts’ panchax (E. olbrechtsi) is divided into a number of subspecies. The nominate form (E. o. olbrechtsi) occurs in southern Liberia, Guinea, and Ivory Coast. Daures’ panchax (E. o. dauresi) is confined to forest streams within the Bandama River drainage, Ivory Coast. The Kassiapleu panchax (E. o. kassiapleuensis) is confined to western Ivory Coast. The azure panchax (E. o. azureus) is confined to the upper Gwen River, a tributary of the Nipoué (Cess) River in east-central Liberia. Puetz’s panchax (E. o. puetzi) is confined to a small area within the St. John River drainage of central Liberia. Josiana’s striped panchax (E. fasciolatus josianae) is known only from a few localities within the rainforests of eastern Sierra Leone. The Totota striped panchax (E. f. tototaensis) is confined to rainforest brooks and rivers in coastal Liberia. Spillman’s toothed

panchax (E. chaperi spillmanni) is confined to marshes, creeks, and small rivers in central and south-eastern Ivory Coast. Sheljuzhko’s toothed panchax (E. c. sheljuzhkoi) is found in the river systems of southern Ivory Coast and western Ghana. Peters’ panchax (Nimbapanchax petersi) is confined to south-eastern Ivory Coast and south-western Ghana. The green panchax (N. viridis) is confined to the upper Saint Paul and upper Saint John rivers of south-eastern Guinea and north-western Liberia. Todd’s panchax (Callopanchax toddi) is known from a few localities in Guinea and Sierra Leone. Brachet’s lampeye (Micropanchax bracheti) is known only from two savanna brook localities in northern Togo. Keilhack’s lampeye (M. keilhacki) is confined to small rivers and brooks in southern Togo. Three lampeyes of the genus Rhexipanchax are potentially threatened by deforestation and mining pollution. Lambert’s lampeye (R. lamberti) is confined to small forest streams and brooks in north-western Guinea. Daget’s lampeye (R. kabae) is known from small streams and rivers in south-central Guinea. The Nimba lampeye (R. nimbaensis) is confined to small rivers, brooks, and swamps in the montane forests of southern Guinea and northern Liberia. Bustamante’s goby (Awaous bustamantei) is confined to rivers on the Gulf of Guinea islands of Bioko, São Tomé, Príncipe, and Annobón. The Gulf of Guinea sleeper goby (Eleotris annobonensis) is confined to Bioko, São Tomé, Príncipe, and Annobón. The Sierra Leone sleeper goby (Kribia leonensis) is known only from three localities in eastern Sierra Leone. Several killifish of the genus Scriptaphyosemion are threatened by habitat destruction and degradation. Berthold’s killifish (S. bertholdi) occurs in shallow pools and small streams within the rainforests of eastern Sierra Leone. Bruening’s killifish (S. brueningi) is confined to the Bomi Hills area of western Liberia and eastern Sierra Leone. Schmitt’s killifish (S. schmitti) is known only from brooks and rainforest streams within a small area of eastern coastal Liberia, between the Cavally and Cess (Nipoué) rivers. Etzel’s killifish (S. etzeli) is confined to brooks and marshes within a small area of western Sierra Leone. Roloff’s killifish (S. roloffi) is confined to Sierra Leone and western Liberia. Several killifish of the genus Aphyosemion are threatened by habitat degradation and overcollection for the international aquarium trade. The goby killifish (A. franzwerneri), Kake killifish (A. volcanum), Amoes’ killifish (A. amoenum), and the Edea killifish (A. edeanum) are all known only from western Cameroon. The sky-blue killifish (A. coeleste) is confined to southern Gabon and western Republic of Congo. The mourning killifish (A. lugens) is confined to south-western Cameroon and northern Equatorial Guinea. Seegers’ killifish (A. seegersi) is known only from north-eastern Gabon and north-western Democratic Republic of Congo. Pellegrin’s killifish (A. louessense) is known only from southern Gabon and south-western Republic of Congo. Schlupp’s killifish (A.

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schluppi) is confined to south-western Republic of Congo. Schouteden’s killifish (A. schoutedeni) is known only from its original collection in the 1920s from north-eastern Democratic Republic of Congo. Ahl’s killifish (A. congicum) is known only from a few brooks between the Cuango and Gamba rivers south-east of Kinshasa, in western Democratic Republic of Congo. The primitive killifish (A. primigenium) is confined to southern Gabon. Wildekamp’s killifish (A. wildekampi) is known from south-eastern Cameroon, northeastern Gabon, south-western Central African Republic, and north-western Republic of Congo. Haas’ killifish (A. cameronense haasi) is known only from its original collection from a mountain stream in northern Gabon. The obscure killifish (A. c. obscurum) is confined to south-central Cameroon. Amiet’s killifish (Fundulopanchax amieti), Kribi killifish (F. fallax), marbled killifish (F. marmoratus), Spoorenberg’s killifish (F. spoorenbergi), Robertson’s killifish (F. robertsoni), ruby-lipped killifish (F. rubrolabialis), and the cinnamon killifish (F. cinnamomeus) are all confined to western Cameroon, where they are threatened by habitat destruction and degradation. The Bandama elephant-snout fish (Mormyrus subundulatus) is confined to the Bandama River in central Ivory Coast and to the Tano River in western Ghana. Felix’s elephantsnout fish (M. felixi) is known only from its original collection in the 1930s from an undefined locality in Cameroon. Thomas’ elephant-snout fish (M. thomasi) is known only from its original collection in the 1930s from the imprecise locality ‘French Congo’. The Sierra Leone mormyrid (Marcusenius meronai) is known only from the Bagbé and Rokel rivers of northern Sierra Leone, where it is threatened by habitat destruction and drought. The Ntem mormyrid (M. ntemensis) is known from the Ntem River of southern Cameroon and the Ivindo River of northern Gabon, where it is threatened by habitat destruction and pollution. Several electric mormyrids of the genus Paramormyrops are threatened by habitat destruction and pollution. The Makokou mormyrid (P. gabonensis) is known from a few scattered localities in Cameroon, Gabon, and north-central Democratic Republic of Congo. Hopkins’ mormyrid (P. hopkinsi) is known from the Ivindo River of north-eastern Gabon and from the Ntem River of south-western Cameroon. The long-tailed mormyrid (P. longicaudatus) is confined to the Ivindo and Ogooué rivers of Gabon. The Ivory Coast mormyrid (Pollimyrus eburneensis) is confined to the Agnébi, San Pedro, and Banco river drainages of southern Ivory Coast. Opdenbosch’s mormyrid (Ivindomyrus opdenboschi) is known only from a few localities in the Ivindo and Ntem rivers of south-western Cameroon, Equatorial Guinea, and northern Gabon. Thomas’ catfish (Notoglanidium thomasi) is known only from three localities in southern Sierra Leone. Walker’s catfish (N. walkeri) is known only from the Ibi River in southern

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Ghana and the Agnébi River in southern Ivory Coast, where it is threatened by invasive aquatic weeds. The Korup catfish (Amphilius korupi) is known only from a few coastal streams in western Cameroon. Laman’s catfish (A. lamani) is confined to a small area of western Democratic Republic of Congo. Risch’s catfish (Chrysichthys aluuensis) is known from south-eastern Nigeria and south-western Cameroon. Teugels’ catfish (C. teugelsi) is confined to south-eastern Liberia and south-western Ivory Coast. The blotched airbreathing catfish (Clarias submarginatus) is confined to a few river drainages in southern Cameroon, northern Gabon, and western Republic of Congo. The asymmetrical-tailed suckermouth catfish (Chiloglanis asymetricaudalis) is a rare species confined to the Rusizi and Luiche rivers of north-eastern Burundi, south-western Rwanda, and eastern Democratic Republic of Congo. The Kogon squeaker catfish (Synodontis kogonensis) is confined to the Kogon and Fatala rivers of western Guinea. The marbled squeaker catfish (S. marmoratus) is known only from a few specimens collected from western Cameroon. The Woleu squeaker catfish (S. woleuensis) is known only from a few rivers in Equatorial Guinea and north-eastern Gabon. Teugels’ eel catfish (Channallabes teugelsi) is known only from the Ivindo River of eastern Gabon and the upper Ogooué River drainage in south-western Republic of Congo. Skelton’s loach catfish (Doumea gracila) is known only from a few localities in south-western Cameroon, northern Equatorial Guinea and north-western Gabon. Chappuis’ loach catfish (D. chappuisi) is known only from a few rivers in Liberia and Guinea-Bissau. Firestone’s loach catfish (Paramphilius firestonei) is known only from three localities in the coastal rivers of western Liberia. Teugels’ loach catfish (P. teugelsi) is confined to western Guinea. The Taï spiny eel (Mastacembelus taiaensis) is known only from the Forecariah River drainage in Guinea and from four other localities in Sierra Leone. The Liberian swamp eel (Monopterus boueti) is known only from a single specimen collected from a small freshwater rivulet flowing into a swamp near Monrovia, Liberia in 1922. The Guinea-Bissau swamp eel (Ophisternon afrum) is known only from two specimens collected from a coastal river at Mansoa Point, Guinea-Bissau in the early twentieth century.

The Upemba Wetlands The Upemba wetlands are an area of flooded grassland located along the Lualaba River in south-eastern Democratic Republic of Congo (Katanga province). The Upemba lechwe (Kobus leche anselli) is a type of antelope restricted to the Upemba wetlands. Now seriously threatened, its population has declined from about 20,000 in the 1980s to less than 1000 today, mainly as a result of poaching. It is at least nominally protected in the western part of Upemba National Park.

The Guineo-Congolian Region

Zimmer’s shrew (Crocidura zimmeri) is known only from two specimens collected from Upemba National Park in 1936. The Upemba masked weaver (Ploceus upembae) is known only from the Upemba wetlands. Schmidt’s toad (Mertensophryne schmidti) is known only from Upemba National Park. The Kanole forest tree frog (Leptopelis parvus) and the Nyonga forest tree frog (L. lebeaui) are both known only from Upemba National Park. The Upemba leaf-folding frog (Afrixalus upembae) is known only from a small area of south-eastern Democratic Republic of Congo.

Lake Tanganyika Located at the intersection of Tanzania, Democratic Republic of Congo, Burundi, and Zambia, Lake Tanganyika is one of the most interesting of the great lakes of Africa. It is the longest lake in the world (673 km) and 1470 m at its greatest depth; only Lake Baikal in Siberia is deeper or larger by volume. Its enormous age is reflected in its animal life. The degree of specialization of the animals to their environment, the pronounced speciation, and the astonishing number of unique species must be seen as the result of the long periods of time the lake has been isolated and without outlet. For example, the number of described species of the family Cichlidae alone in Lake Tanganyika is at least 250, virtually all of which are endemic. The various species have evolved during millions of years toward an effective utilization of every existing niche in the lake. This has resulted in an exceptionally high energy flow through various food chains – algae–plankton–fishes – resulting in an immense bioproductivity. Many African lakes show similar features, but for Lake Tanganyika it is remarkable, because below a depth of about 400 m there is no oxygen. This means that about two-thirds of the lake is a freshwater desert without life. Yamaoka’s cichlid (Petrochromis macrognathus) is confined to Lake Tanganyika. The white-spotted cichlid (Tropheus duboisi) is confined to three subpopulations within the northern part of Lake Tanganyika. It is in heavy demand by the international aquarium trade. The zebra cichlid (Trematocara zebra) is confined to the rocky north-western shore of Lake Tanganyika. Kendall’s cichlid (Lepidiolamprologus kendalli) is confined to the southern part of Lake Tanganyika. The eccentric cichlid (Perissodus eccentricus) is a deepwater species known only from a single locality in Lake Tanganyika. The Uvira cichlid (Lamprologus finalimus) is known only from a single specimen collected at the northern tip of Lake Tanganyika in 1931. The Kungwe Bay cichlid (L. kungweensis) is a rare species confined to Kungwe Bay, on the north-eastern shore of Lake Tanganyika. Several cichlids of the genus Neolamprologus are endemic to Lake Tanganyika, where they are threatened by

sedimentation. Christy’s cichlid (N. christyi) is known only from a strip along the central-eastern Tanzanian shoreline. Schreyen’s cichlid (N. schreyeni) is known only from the eastern shores of Burundi. Wauthion’s cichlid (N. wauthioni) is known only from a small area near Kalemi, Democratic Republic of Congo. The olive cichlid (N. olivaceous) is known only from the area of Ruhanga Bay, Democratic Republic of Congo. Margaret’s cichlid (Pseudosimochromis margaretae) is confined to shallow waters off Kigoma, Tanzania. The margined cichlid (P. marginatus) is confined to shallow waters off the Ubwari Peninsula on the north-western shore of Lake Tanganyika. Hecqu’s cichlid (Xenochromis hecqui) is confined to the deeper waters of Lake Tanganyika. Poll’s herring cichlid (Cyprichromis microlepidotus) is confined to the northern part of Lake Tanganyika. The Burton Bay sand cichlid (Xenotilapia burtoni) is confined to Burton Bay, where it is threatened by habitat disturbance and pollution. The Tanganyika clown (Eretmodus cyanostictus) is widespread in southern Lake Tanganyika but confined to a very narrow band along the shoreline. The bigeye lates (Lates mariae), Tanganyika lates (L. angustifrons), and forktail lates (L. microlepis) are perch-like fish endemic to Lake Tanganyika and the Ruzizi and Malagarasi river deltas, where they are heavily fished. The Lake Tanganyika tetra (Micralestes vittatus) is confined to rivers and lagoons within the Lake Tanganyika basin.

Lake Albert Lake Albert is located in the center of the African continent, on the border between Uganda and the Democratic Republic of Congo. It is the northernmost of the chain of lakes in the Albertine Rift, the western branch of the East African Rift. The Lake Albert lates (Lates macrophthalmus) is confined to waters deeper than 18 m in a lake that is becoming increasingly eutrophic, and which is likely to become anoxic unless organic pollution is reduced. It is also subject to heavy fishing. Hulot’s yellowfish (Labeobarbus huloti) is known only from a single locality on the Vuda River in the Lake Albert basin.

Lake Rukwa Lake Rukwa is an endorheic lake located in the Rift Valley of south-western Tanzania, midway between Lake Tanganyika and Lake Malawi. The lake has seen large fluctuations in its size over the years, due to the varying inflow of streams. The Rukwa sardine (Chelaethiops rukwaensis) is confined to Lake Rukwa, where it is heavily fished. The Katavi cichlid (Haplochromis katavi) is confined to the Katuma River in the Lake Rukwa basin. The Rukwa tilapia (Oreochromis rukwaensis) is a type of cichlid confined to the Lake Rukwa basin.

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The Rukwa barb (Enteromius pseudotoppini) is known only from the Lake Rukwa basin. The Mbozi suckermouth catfish (Chiloglanis mbozi) is known only from a small stream in the Lake Rukwa basin. Similarly, the Rukwa suckermouth catfish (C. rukwaensis) is known only from two small rivers.

Lake George Lake George is located north-east of Lake Edward in Uganda. Two species of mouthbrooding cichlid (Haplochromis) are endemic to Lake George, where they are threatened by water pollution. The Kashaka Crater Bay cichlid (H. petronius) is almost entirely confined to rocky bays in the vicinity of the fishing village of Kashaka. The bronze-coloured cichlid (H. aeneocolor) is confined to Lake Edward and the Kazinga Channel.

Lake Barombi-Mbo Lake Barombi-Mbo is a relatively large and ancient volcanic crater lake located in south-western Cameroon (south-west region). Only the upper 40 m contains enough dissolved oxygen to sustain vertebrate life. Nevertheless, it is home to a large number of endemic species, all of which are threatened by pollution and sedimentation due to human activities and water extraction. Commercial fishing is forbidden in the lake, but extensive subsistence fishing has caused serious declines in cichlid stocks. The Dikume cichlid (Konia dikume) and the Konye cichlid (K. eisentrauti) are both confined to Lake Barombi-Mbo. The Myaka cichlid (Myaka myaka) is confined to open, deeper waters in Lake Barombi-Mbo. The Pungu cichlid (Pungu maclareni) is confined to the shallow, nearshore waters of Lake Barombi-Mbo. The Nsess cichlid (Stomatepia mariae), Mongo cichlid (S. mongo) and Pindu cichlid (S. pindu) are all confined to Lake Barombi-Mbo. The Fissi tilapia (Sarotherodon caroli), blackfin tilapia (S. linnellii), Keppi tilapia (S. lohbergeri), and Kululu tilapia (S. steinbachi) are all confined to Lake Barombi-Mbo. Maclaren’s airbreathing catfish (Clarias maclareni) is confined to Lake Barombi-Mbo.

Lake Bermin Lake Bermin is a small, isolated volcanic crater lake located in south-western Cameroon (south-west region). Nine small species of tilapia cichlid (Coptodon) are found only in this one small lake, representing one of the highest rates of endemism to area in the world. The Bakossi tilapia (C. bakossiorum), Bemin tilapia (C. bemini), Snyder’s tilapia (C. snyderae), Thys’ tilapia (C. thysi), goitered tilapia (C. gutturosa), bottom-walking tilapia (C. bythobates), spongotroktis tilapia (C. spongotroktis), blonde tilapia (C. flava), and imbriferna tilapia (C. imbriferna) are all highly threatened by pollution, sedimentation, and occasional volcanic carbon dioxide emissions.

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Lake Ejagham Lake Ejagham is a small non-volcanic lake located in southwestern Cameroon (south-west region). Like other isolated lakes in the area it has a high number of endemic cichlid species, all of which are threatened by pollution, sedimentation, and an introduced species of catfish. The Ejagham tilapia (Coptodon ejagham), Deckert’s tilapia (C. deckerti), slender tilapia (C. fusiforme), and black tilapia (C. nigrans) are all confined to Lake Ejagham. Lamprecht’s tilapia (Sarotherodon lamprechti) and Knauer’s tilapia (S. knauerae) are confined to Lake Ejagham. The Ejagham blue killifish (Fundulopanchax gardneri lacustris) is confined to Lake Ejagham.

The Congo River Drainage The Congo is the second longest river in Africa after the Nile and the second largest in the world by discharged volume of water (after the Amazon). It is also the world’s deepest river, with measured depths in excess of 220 m. The river and its many tributaries flow through the Congo Rainforest and provide habitat for a large number of endemic species. Poll’s cichlid (Ctenochromis polli) is known only from a few localities within the lower and middle Congo, where it is threatened by pollution and urbanization. Teugels’ cichlid (Lamprologus teugelsi) and Werner’s cichlid (L. werneri) are both known only from Stanley Pool and parts of the lower Congo. The Dungu sprat (Microthrissa minuta) is known from a small number of localities spread throughout the Congo River drainage, but is threatened by overfishing and dam construction. The Boma mormyrid (Stomatorhinus microps) is known for certain only from the lower Congo, although there are scattered unconfirmed records throughout the Congo Basin. Teugels’ snake catfish (Clariallabes teugelsi) is known from Stanley Pool (Pool Malebo) and the lower Congo. The Upper Congo River Upstream of Stanley Falls (Boyoma Falls), near Kisangani (Stanleyville), the Congo River is known as the Lualaba River. It is the primary source (headstream) of the Congo River drainage. The elongated cichlid (Tylochromis elongatus) is confined to the upper Congo (Lualaba) River and Wagenia Falls. The Kisangani cichlid (Congochromis pugnatus) is known only from a single locality near Kisangani, in the upper Congo (Lualaba) River. Nichols’ barb (Enteromius nigrifilis) is known only from a single specimen collected in the 1920s from the junction of the Lualaba and Luvua rivers. Leleup’s salmon (Opsaridium leleupi) is known only from the upper Congo (Lualaba) River in southern Democratic Republic of Congo. Whitehead’s sawtooth pellonuline (Potamothrissa whiteheadi) is a type of herring known only from a few specimens collected within the Hombo River (a tributary of the Luhoko

The Guineo-Congolian Region

River) and the Lindi/Tshopo tributary, both in north-eastern Democratic Republic of Congo. The Lukinda mormyrid (Petrocephalus squalostoma) is known only from the Lukinda and Luapula rivers in southeastern Democratic Republic of Congo. Aelsbroeck’s mormyrid (Hippopotamyrus aelsbroecki) is known only from its original collection in the 1940s from south-eastern Democratic Republic of Congo. The spotted-backed squeaker catfish (Synodontis dorsomaculatus) is confined to the upper Congo (Lualaba) River. It is threatened by overfishing and pollution. The Katanga suckermouth catfish (Chiloglanis elisabethianus) is confined to a few tributaries of the upper Congo (Lualaba) in south-eastern Democratic Republic of Congo and northern Zambia. Lake Mweru is located on the border between northern Zambia and south-eastern Democratic Republic of Congo. The second largest lake in the Congo River drainage basin, it is fed mainly by the Luapula River which comes in through swamps from the south, and by the Kalungwishi River from the east. The Mweru cichlid (Thoracochromis moeruensis) is confined to Lake Mweru, where it is threatened by overfishing. The Mweru sprat (Poecilothrissa moeruensis) is confined to Lake Mweru. Bredo’s mormyrid (Campylomormyrus bredoi) is known only from Lake Mweru. The Lukinda River is located within the Lake Mweru basin of south-eastern Democratic Republic of Congo. The line-spotted barb (Enteromius brachygramma) and the Lukinda barb (E. lukindae) are both known only from the Lukinda River drainage. The Luvua River (also known as the Lowa River) is located in south-eastern Democratic Republic of Congo. It flows from the northern end of Lake Mweru in a north-westerly direction to is confluence with the upper Congo (Lualaba). The Chambeshi River of north-eastern Zambia is the most remote headwater of the Congo River. The Chambeshi barb (Enteromius lornae) is known only from the Chambeshi River drainage. The Luapula River is a transitional river forming, for nearly all of its length, part of the border between Zambia and the Democratic Republic of Congo. It joins Lake Bangweulu to Lake Mweru. Bleher’ cichlid (Steatocranus bleheri) is known only from the Kafubu River, a tributary of the Luapula River. The mola cichlid (Chetia mola) is confined to the Luongo River, a tributary of the Luapula River located in northern Zambia. The Luongo tilapia (Orthochromis luongoensis) is a type of cichlid confined to the Luongo River, a tributary of the Luapula River in northern Zambia. The Upemba yellowfish (Labeobarbus upembensis) and Gestetner’s yellowfish (L. gestetneri) are both known only from their original collection within the Kalumengonga River, a tributary of the Lualaba River, during the 1970s.

Myer’s barbel (Barbus myersi) is known only from its original collection in the 1930s from the Muniengashsi River, a tributary of the Luapula River. Three species of killifish (Nothobranchius) within the Luapula River drainage are threatened by loss of their seasonal wetland habitats. Rosenstock’s killifish (N. rosenstocki) is confined to north-eastern Zambia. Malaisse’s killifish (N. malaissei) is confined to temporary pools and marshes within the Luapula River drainage of south-eastern Democratic Republic of the Congo. Symoens’ killifish (N. symoensi) is confined to small temporary pools, swamps, and flooded grassy areas adjacent to rivers in northern Zambia and south-eastern Democratic Republic of Congo. Malaisse’s shellear (Parakneria malaissei) is known only from the Luanza River, a tributary of the Luapula River in south-eastern Democratic Republic of Congo. The Luapula mormyrid (Campylomormyrus luapulaensis) is known only from two localities within the Luapula River. The Luongo suckermouth catfish (Chiloglanis macropterus) is confined to the Luongo River, a tributary of the Luapula River located in northern Zambia. The Lufira River is located in south-central and southeastern Democratic Republic of Congo. The Lofoi Falls reed frog (Hyperolius polystictus) is known only from a single locality within the Lofoi River, a tributary of the Lufira River in south-eastern Democratic Republic of Congo. Symoens’ cichlid (Lamprologus symoensi) is known only from two effluents of the Lufira River. The Mwashia tilapia (Oreochromis salinicola) is confined to a few saline springs within the Lufira River drainage of south-eastern Democratic Republic of Congo. Witte’s yellowfish (Labeobarbus wittei) and the Lufira yellowfish (L. iphthimostoma) are both confined to the Lufira River drainage. Hasson’s killifish (Nothobranchius hassoni) and Poll’s killifish (N. polli) are both confined to temporary pools and swamps within the Lufira River drainage. The Lufira shellear (Parakneria lufirae) is known only from a few specimens collected from the Lufira River and its effluents. Thys’ shellear (P. thysi) is known only from its original collection in the 1960s from the rapids above Kiubo Falls, in the Lufira River drainage. The Lufira suckermouth catfish (Chiloglanis lufirae) is known only from the Muye River, an effluent of the Lufira River located within Upemba National Park. The Lufira squeaker catfish (Synodontis lufirae) is confined to the Lufira River system and Lake Koni. The Luhoho River is located in central-eastern Democratic Republic of Congo. The Luhoho yellowfish (Labeobarbus longifilis), longfinned yellowfish (L. longidorsalis), and Braun’s yellowfish (L. brauni) are all known only from the Luhoho River drainage. The Luhoho shellear (Parakneria kissi) is confined to the Luhoho River.

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The Afrotropical Realm

Marlier’s suckermouth catfish (Chiloglanis marlieri) is known only from the Ndakirwa River, an effluent of the Luhoho River. The Kasuku River is located in east-central Democratic Republic of Congo. The Kasuku barb (Enteromius papilio) is known only from an unnamed tributary of the Kasuku River. Stanley Falls (now known as Boyoma Falls) is a series of cataracts, each no more than 5 m high, extending over more than 100 km along a curve of the Lualaba River between Ubundu and Kisangani. The seven cataracts have a total drop of 61 m, and form the largest waterfall by volume in the world. Bentley’s mormyrid (Marcusenius bentleyi), Wagenia Falls whiptailed catfish (Phractura fasciata), Wagenia Rapids loach catfish (Dolichamphilius longiceps), and Wagenia Falls claroteid catfish (Amarginops platus) are all known only from Wagenia Falls, the last in the Stanley Falls cataract system. The Middle Congo River The middle or central Congo stretches from Kinshasa to Stanley Falls (Boyoma Falls). The Monsembé cichlid (Pelmatochromis ocellifer) is known only from a single specimen collected from the central Congo in 1899. Brichard’s cichlid (Teleogramma brichardi) is known only from the rapids near Kinsuku. The monogrammed cichlid (T. monogramma) is confined to a few localities within the middle Congo drainage. The small-toothed cichlid (Tylochromis microdon) is known only from Lake Tumba and possibly Lake MaiN’Dombe, within the middle Congo River drainage. The pygmy barb (Enteromius pygmaeus) is known only from a single rainforest stream within the middle Congo River drainage. The Lulua labeo (Labeo luluae) is known from two specimens collected from the Lulua and Aruwimi rivers in the early twentieth century. The smooth-belly pellonuline (Congothrissa gossei) is a type of herring known only from a few specimens collected from two localities within the middle Congo River drainage. Schouteden’s pufferfish (Tetraodon schoutedeni) is known from Stanley Pool and the middle Sangha River. Poll’s tetra (Alestopetersius nigropterus) is only known for certain from Lake Mai Ndombe, although it may occur in the Lengoué River as well. Schouteden’s tetra (Clupeocharax schoutedeni) is found widely but patchily in the middle Congo drainage. Known localities include Lake Tumba, Lake Yandja (Yangambi), and the area in and around Lake Mai-Ndombe. Philippe’s tetra (Neolebias philippei) is known only from two widely spaced localities within the middle Congo drainage (the Itimbiri and Tshuapa rivers). The Kutu characin (Paraphago rostratus) is only known from a single locality in the central Congo River drainage. Ladiges’ shellear (Parakneria ladigesi) is known only from the Cuango and Lulua rivers.

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Two species of killifish (Aphyosemion) are threatened by overcollection for the international aquarium trade. The Léfini killifish (A. lefiniense) is known only from the Léfini River and from a small effluent of the Louna River, both within the Lesio-Louna Gorilla Reserve of south-western Republic of Congo. Chauche’s killifish (A. chauchei) is known only from the Likona River drainage of central Republic of Congo. The New Antwerp mormyrid (Mormyrops microstoma) is known only from its original collection during the nineteenth century. Simeons’ catfish (Clariallabes simeonsi) is known only from the Congo River drainage in north-central Democratic Republic of Congo. Lambert’s upside-down catfish (Microsynodontis lamberti) is known only from the Lilanda River in north-central Democratic Republic of Congo. Boulenger’s loach catfish (Tetracamphilius angustifrons) is known from the Oubangui River (upstream of Bangui) and from another specimen collected from Stanley Pool (Pool Malebo). The minute loach catfish (Trachyglanis minutus) is known for certain only from the Oubangui River, but may occur as well within the Lossi River (a tributary of the Sangha River). The Chobe loach catfish (Zaireichthys dorae) is known only from the Luachimo River in north-eastern Angola. Lake Mai-Ndombe (formerly known as Lake Leopold II) is a large freshwater lake in western Democratic Republic of Congo. It is of irregular shape and is known to double in size during the rainy season. It drains to the south through the Fimi River into the Kwah and Congo. The Mai-Ndombe jewel cichlid (Hemichromis cerasogaster) is confined to Lake Mai-Ndombe and some smaller satellite lakes. The transvestite cichlid (Nanochromis transvestitus) is confined to Lake Mai-Ndombe and a few satellite lakes. The Mai-Ndombe dwarf sprat (Nannothrissa stewarti) is confined to Lake Mai-Ndombe. The Mai-Ndombe mountain catfish (Amphilius opisthophthalmus) is known only from a few specimens collected from Lake Mai-Ndombe. Lake Tumba is a shallow lake in the middle Congo River drainage, north-western Democratic Republic of the Congo. All of the bigger fish in the lake have long disappeared due to overfishing. The Tumba cichlid (Lamprologus tumbanus) is confined to Lake Tumba. The Tumba robber tetra (Rhabdalestes aerates) is confined to Lake Tumba. Stanley Pool (Pool Malebo) is a shallow, lake-like widening of the middle Congo River in south-eastern Congo and southwestern Democratic Republic of Congo. General threats include overfishing, urbanization, and pollution. Dubois’ pufferfish (Tetraodon duboisi) is known only from a single locality in Stanley Pool, where it may already be extinct. There are a few specimens owned by private aquarists in the United States.

The Guineo-Congolian Region

The Stanley Pool labeo (Labeo maleboensis) is known only from a single specimen collected from Stanley Pool in the 1990s. The hummingbird lampeye (Poropanchax myersi) is known only from Stanley Pool, including its islands. Chabanaud’s upside-down catfish (Atopochilus chabanaudi) is known only from an area of Stanley Pool near Brazzaville. The Stanley Pool squeaker catfish (Synodontis pulcher) is known only from two specimens collected from Stanley Pool. Brien’s loach catfish (Belonoglanis brieni) is known only from a single channel within Stanley Pool. The Stanley pool loach catfish (Dolichamphilius brieni) is known only from a single channel in Stanley Pool. The delicate loach catfish (Psammphiletria delicata) is known only from Stanley Pool. The Itimbiri River is located in north-central Democratic Republic of Congo. The Rubi mormyrid (Mormyrops intermedius) is known only from its original collection within the Rubi River, a tributary of the Itimbiri River. The Kagola electric catfish (Paradoxoglanis cryptus) is known only from the Kagola River, a tributary of the Itimbiri River. The Sangha River is formed at the confluence of the Mambéré River and the Kadéi River at Nola, in western Central African Republic. From there it flows along the border with Cameroon and then of the Republic of Congo before finally joining the Congo River. The Sangha tetra (Phenacogrammus taeniatus) is known only from a single locality on the middle Sangha River.The Sangha squeaker catfish (Synodontis longispinis) is known only from the Sangha River. The Mbessy eel catfish (Channallabes sanghaensis) is known only from the Mbessy River, a tributary of the Sangha River. The Sangha loach catfish (Trachyglanis sanghensis) is known only from a single locality on the Sangha River. The Dja River (also known as the Ngoko River) forms part of the Cameroon/Republic of Congo border. It has a course of roughly 720 km. General threats include cobalt mining and overfishing. The Bumba mormyrid (Mormyrops batesianus) is known only from the Bumba River, a small tributary of the Dja River in south-eastern Cameroon. The leopard squeaker catfish (Synodontis pardalis) is a type of upside-down catfish known only from a few localities in the Dja River and its tributary, the Libi River, in southern Cameroon. The Kasai River is a major left-bank (southern) tributary of the central Congo River, located in north-eastern Angola and south-western Democratic Republic of the Congo. Intense diamond mining and the resulting sedimentation and pollution are a major threat to small rivers within the Kasai drainage.

The Kasai cichlid (Paratilapia toddi) is known only from the Kasai River. Thys’ cichlid (Sargochromis thysi) is known only from the Luembe River, a tributary of the Kasai River in north-eastern Angola. Roux’s cichlid (Steatocranus rouxi) is known only from the Lulua River, a tributary of the Kasai River. The Lulua cichlid (Haplochromis luluae) is known only from the Lulua River, a tributary of the Kasai River. Schwetz’s cichlid (H. schwetzi) is confined to the Cuango River drainage, a tributary of the Kasai River. The middle-scaled barb (Enteromius mediosquamatus) is known from the Luachimo, Lucoge, and Lulua rivers, tributaries of the upper Kasai River. Machado’s barb (E. machadoi) is known only from a single pool within the Luachimo River drainage of north-eastern Angola. Petchkovsky’s barb (E. petchkovskyi) and the tail-signed barb (E. caudosignatus) are both known only from their original collection from the Cuilo River of north-eastern Angola. Jubb’s yellowfish (Labeobarbus jubbi) is known from the Luembe, Luachimo, and Canhicuna rivers of northeastern Angola. Bowen’s salmon (Opsaridium boweni) is known only from the Lulua River drainage of south-central Democratic Republic of Congo. Pellegrin’s salmon (O. maculicauda) is known only from its original collection in the 1920s from north-eastern Angola. Poll’s citharine (Hemigrammocharax lineostriatus) is known only from the Cavuemba River, a tributary of the Kasai River in north-eastern Angola. The Chicapa characin (Dundocharax bidentatus) is confined to the Chicapa River, a small tributary of the Lucoge River in north-eastern Angola. The Dundo shellear (Parakneria vilhenae) is known only from the Dundo River, a tributary of the Kasai River in northeastern Angola. The Tchimenji lampeye (Aplocheilichthys nigrolateralis) is known only from the Tchimenji River, a tributary of the Kasai River in north-eastern Angola. The Kananga mormyrid (Pollimyrus maculipinnis) is known only from its original collection at Kananga (Luluabourg), south-central Democratic Republic of Congo. The donkey-faced mormyrid (Campylomormyrus cassaicus) is known only from two specimens collected from the Luachimo River and another from the Kasai River main channel. Poll’s mormyrid (Gnathonemus barbatus) is known only from a single locality within the Kasai River of northeastern Angola. The Tshikapa mormyrid (Marcusenius macrophthalmus) is known only from its original collection in the 1920s from north-eastern Angola. The Zovo mormyrid (M. cuangoanus) is known only from a small area of the Cuango River. The Dundo mormyrid (M. dundoensis) is known only from a small area of the Kasai and Cuango rivers of northeastern Angola.

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Machado’s airbreathing catfish (Platyclarias machadoi) is known only from a single locality within the Cuango River of north-eastern Angola. Pellegrin’s upside-down catfish (Atopochilus pachychilus) is known only from the Lubilonji River, a tributary of the Kasai River. The Cuango upside-down catfish (A. macrocephalus) is known only from the Cuango River in the vicinity of Fort Don Carlos I, Angola. Boulenger’s upside-down catfish (Euchilichthys boulengeri) is known only from the Lulua River, a tributary of the Kasai River. The Kasai whiptailed catfish (Phractura macrura) is known only from the Uamba and Cuilo rivers in northeastern Angola. Roberts’ squeaker catfish (Synodontis robertsi) is a type of upside-down catfish known only from specimens collected in the Lukenie River, east-central Democratic Republic of Congo, in 1974. The Cuango squeaker catfish (S. cuangoanus) is known only from a small area of the Cuango River. The Lufume loach catfish (Zaireichthys flavomaculatus) is known only from the Lufume River, a tributary of the Kasai River. The Lulua loach catfish (Trachyglanis intermedius) is known only from the Lulua River, a tributary of the Kasai River. The Oubangui River (also called the Ubangui) is the largest right-bank (northern) tributary of the Congo River. The Uele River, which arises in the mountains near Lake Albert, is in turn the longest tributary of the Oubangui River. The Mbomou cichlid (Steatocranus ubanguiensis) is known only from the Mbomou River, a tributary of the Oubangui River in the Central African Republic. Schouteden’s barb (Clypeobarbus schoutedeni) is known only from the Dungu River, a tributary of the Uele River in north-eastern Democratic Republic of Congo. The Fort Sibut barb (Enteromius castrasibutum) and the Tomi barb (E. tomiensis) are both known only from their original collection in the 1930s from south-western Central African Republic. The lemon barb (E. citrinus) is known only from a few specimens collected from the Oubangui and Uele rivers of northern Democratic Republic of Congo. The Oubangui trout barb (Raiamas longirostris) is known only from its original collection from north-western Democratic Republic of Congo in the early twentieth century. The Dungu yellowfish (Labeobarbus urotaenia) is known only from the Dungu River, a tributary of the Uele River. The ambiguous tetra (Micralestes ambiguus) is known only from the Bari River, a tributary of the Oubangui River in Democratic Republic of Congo (Équateur province). Bleher’s tetra (Phenacogrammus bleheri) is confined to a small tributary of the Bari River, itself a tributary of the Oubangui River. Abel’s robber tetra (Brycinus abeli) is known only from Fort Sibut on the Oubangui River. Holly’s citharine (Nannocharax hollyi) and the Fort Sibut citharine (N. pteron) are both known only from their original

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collection in the 1930s from south-western Central African Republic. Lang’s disichodus (Distichodus langi) is known only from Faradje in the Uele River drainage. Decorse’s killifish (Aphyosemion decorsei) is known only from its original collection in south-western Central African Republic in the early twentieth century. The Dungu mormyrid (Petrocephalus hutereaui) is known only from the Dungu River, a tributary of the Uele River. The Bomekendi mormyrid (Hippopotamyrus macroterops) is known only from the Bomekendi River, an effluent of the Uele River. The Bima mormyrid (H. retrodorsalis) is known only from its original collection in the early twentieth century from a small rainforest brook (tributary of the Bima River), in the Uele River drainage. The Uele catfish (Chrysichthys uniformis) is known only from a single specimen collected in the Uele River. The dotted squeaker catfish (Synodontis multimaculatus) is known only from a single specimen collected in the early twentieth century from northern Democratic Republic of Congo. Vaillant’s squeaker catfish (S. vaillanti) is known only from its original collection in the late nineteenth century from the upper Oubangui River drainage. The Oubangui loach catfish (Psammphiletria nasuta) is known only from the Oubangui River upstream of Bangui, Central African Republic. The clandistine loach catfish (Tetracamphilius clandestinus) is known only from the Oubangui River upstream of Bangui, Central African Republic. The Tshuapa River (also known as the Ruki or Ruka River) is located in north-central and north-western Democratic Republic of Congo. Deguide’s barb (Enteromius deguidei) is known only from marshes along the Boteli River, a tributary of the Tshuapa River. Fowler’s barb (E. syntrechalepis) and the Oka barb (E. okae) are both known only from the Oka River, a tributary of the Tshuapa River. The Boteli tetra (Neolebias gracilis) is known only from marshes along the Boteli River. The Tshuapa squeaker catfish (Synodontis camelopardalis) is known only from its original collection in 1971. The Ituri/Aruwimi River is located in north-eastern and north-central Democratic Republic of Congo. It arises near Lake Albert as the Ituri River and runs generally south-west before becoming the Aruwimi River where it is joined by the Nepoko River. From there it continues westward before finally joining the Congo River. The Kavalli barb (Enteromius tetraspilus) is known only from the Kavalli and upper Ituri rivers. The Epulu yellowfish (Labeobarbus macroceps) is known only from the Epulu River, a tributary of the Ituri River. The Aruwimi catfish (Gephyroglanis gymnorhynchus) is confined to the Aruwimi River. The Ituri squeaker catfish (Synodontis iturii) is known only from its original collection in the early twentieth century.

The Guineo-Congolian Region

The Lower Congo River The lower Congo comprises the lower section of the main river from its mouth at the Atlantic Ocean to the twin capitals of Brazzaville and Kinshasha. The Grand Inga Dam project is a series of seven proposed hydroelectric power stations located at Inga Falls. The first two existing dams have had minimal environmental impact, though the proposed dams, if built, will divert a significant amount of water flow and block a great part of the Congo mainstream. The tidal cichlid (Tylochromis praecox) is known only from two localities in the tidal stretches of the lower Congo. The Inga Dam cichlid (Steatocranus glaber) is confined to a small area of the lower Congo River. The Inga Rapids cichlid (Nanochromis consortus) is confined to the Inga Rapids. The lesser cichlid (N. minor) is known from two localities within the lower Congo rapids, above the Inga 1 and 2 dams. The splendent cichlid (N. splendens) is confined to the lower Congo down to below the Inga 1 and 2 dams. The gracile cichlid (Teleogramma gracile) is confined to the main channel of the lower Congo River. The lower Congo tilapia (Oreochromis lepidurus) is a type of cichlid known only from the mouth of the Congo upstream as far as Boma. The banded cichlid (Thorachromis fasciatus) is known only from a single locality within the lower Congo River. Demeus’ cichlid (T. demeusii) is confined to the Lower Congo and effluent streams. The blind cichlid (Lamprologus lethops) is a nonpigmented and blind species known only from a single locality within the lower Congo. It is believed to live as deep as 160 m below the surface. The embroidered cichlid (L. tigripictilis) is confined to a small stretch of the lower Congo between Luozi and Matadi. Kheel’s trout barb (Raiamas kheeli) is known only from a few specimens collected from two widely separated localities in the lower Congo River, although it likely extends more widely. Laman’s barb (Enteromius lamani) is known only from its original collection from a small effluent of the lower Congo in the early twentieth century. The Congo blind barb (Caecobarbus geertsii) occurs only in the Mbanzagungu Caves near Thysville in the lower Congo River drainage, where it has been protected. It is, however, extremely vulnerable to disturbance and can easily be wiped out. Tshibwabwa’s labeo (Labeo rectipinnis) is known only from two localities in the lower Congo River. Dartevelle’s yellowfish (Labeobarbus dartevellei) is known only from a single specimen collected in the 1940s from the lower Congo River. Miller’s climbing gourami (Microctenopoma milleri) is known only from the lower Congo and its tributary, the Lunyonzo River. The Schelly tetra (Micralestes schelly) and Günther’s tetra (M. holargyreus) are each known only from a single locality within the lower Congo River.

The Inga elephant-snout fish (Mormyrus iriodes) and the dark blue elephant-snout fish (M. cyaneus) are both known only from the lower Congo River drainage. The small mormyrid (Mormyrops parvus) is confined to the lower Congo River. Schouteden’s mormyrid (Stomatorhinus schoutedeni) is known only from its original collection from the lower Congo River in the 1940s. The Kidada mormyrid (S. ater) is known only from an effluent of the lower Congo River. The pallid catfish (Notoglanidium pallidum) is known only from the lower Congo River. The spiral-feeding catfish (Chrysichthys helicophagus), Poll’s catfish (C. polli), and the carpenter catfish (C. dendrophorus) are all confined to the lower Congo River. The Lubozi catfish (Bagrus lubosicus) is known only from the Lubozi River, a tributary of the Luala River within the lower Congo River drainage. The sky-blue catfish (B. caeruleus) is known only from the lower Congo River. Bove’s whiptailed catfish (Phractura bovei) is known only from its original collection from a single locality within the lower Congo River in the late nineteenth century. Brichard’s squeaker catfish (Synodontis brichardi) is confined to the lower Congo River rapids. The blind eel catfish (Gymnallabes nops) is known only from a few specimens collected from the lower Congo River. The Kinsuka loach catfish (Zaireichthys zonatus) is known only from two specimens collected from the rapids just below Stanley Pool in the 1960s. The bird-headed spiny eel (Mastacembelus aviceps), Crassus spiny eel (M. crassus), and the late spiny eel (M. latens) are all confined to a few areas of the lower Congo River. The Inkisi River is the closest large tributary to the Congo River’s mouth. It flows in a northerly direction from Angola to its confluence with the Congo some 80 km downstream from Kinshasha/Brazzaville. Roberts’ yellowfish (Labeobarbus robertsi) is known only from the Inkisi River. The Inkisi butter catfish (Schilbe zairensis) is known only from the Inkisi River. The M’pozo River is located in north-western Angola and south-western Democratic Republic of Congo. The M’pozo cichlid (Steatocranus mpozoensis) is known only from the mouth of the M’pozo River and from the opposite bank in the main Congo River. The Foulakari River is located in southern Republic of Congo. The Foulakari barb (Enteromius erythrozonus) is known only from the Foulakari River.

The Kagera River The Kagera River (also known as the Akagera River) is located in Burundi and Rwanda. It forms part of the upper headwaters of the Nile. The Kagera squeaker catfish (Synodontis ruandae) is known only from the middle and upper Kagera River drainage.

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The Nyabarongo River The Nyabarongo River is located in the upper Kagera River drainage. The Mukungwa River is an effluent of the Nyabarongo River located in northern Rwanda. The Mukungwa yellowfish (Labeobarbus ruasae) and the broadmouth yellowfish (L. platystoma) are both known only from the Mukungwa River.

The Volta River Drainage The Volta River is located in Burkina Faso, Togo, and Ghana. It includes Lake Volta, an impoundment of the Akosombo Dam on the lower Volta River, which is one of the largest man-made reservoirs in the world. The Upper Volta robber tetra (Brycinus luteus) is known from a few rivers in the upper Volta drainage of Burkina Faso, where it is threatened by pollution and drought. Two species of squeaker catfish (Synodontis) are endemic to the Volta River drainage, where they are threatened by habitat destruction and degradation. Poll’s squeaker catfish (S. macrophthalmus) is known only from a single locality in Ghana. Arnoult’s squeaker catfish (S. arnoulti) is known only from two localities in northern Togo and western Burkina Faso. The Oti River The Oti River (also known as the Pendjari River) rises on the Benin/Burkina Faso border and flows through northern Togo before finally joining the Volta River in Ghana. The naked shellear (Cromeria occidentalis) is a sandburrowing freshwater fish known for certain only from the Oti River. The Lower Volta River The lower Volta empties into the Gulf of Guinea. The Hedjo barb (Enteromius guildi) is known only from the Hedjo River, a tributary of the lower Volta located on the border between south-western Togo and south-eastern Ghana.

The Niger River The Niger River (fleuve Niger in French) is the principal river of West Africa, extending as it does some 4180 km from the Guinea Highlands to the Gulf of Guinea. The silver catfish (Bagrus filamentosus) is known only from a single specimen collected in the 1920s, most likely (but not certainly) from somewhere in the Niger River drainage. The Upper Niger The Upper Niger exhibits an unusual crescent-shaped course that baffled geographers for centuries. From its source in the highlands of south-eastern Guinea, just 240 km from the Atlantic Ocean, it runs directly away from the sea into the Sahara Desert before taking a sharp turn near the ancient city of Timbuktu.

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Caroline’s robber tetra (Brycinus carolinae) is known only from the Niandan River and two other small rivers at the headwaters of the Niger in Guinea. Oudot’s mormyrid (Mormyrops oudoti) is known only from a single specimen collected from the Niger River at Bamako, south-western Mali. The Kissidougou River is located in the highlands of southeastern Guinea. The Kissidougou barb (Enteromius kissiensis) is known only from the Kissidougou River. The Benue River The Benue River (la Bénoué in French) is the main tributary of the Niger. It originates in the Adamawa Plateau of northern Cameroon and flows west into eastern Nigeria. The Bamenda garra (Garra allostoma) is confined to the mountains of north-western Cameroon, within the Benue River drainage. The Benue killifish (Aphyosemion bualanum) is known only from the upper reaches of the Benue River drainage and the Mayo-Kébbi River drainage of the Central African Republic. The Menchum River is a tributary of the Benue River located in western Cameroon. The Bamenda suckermouth catfish (Chiloglanis niger) is known only from the Menchum River. The Lower Niger and the Niger River Delta The lower Niger is located in southern Mali, south-western Niger, northern Benin, and western Nigeria, with the delta itself covering a wide area of southern Nigeria. The latter, an important global oil terminal, is threatened by pollution and civil unrest. The Niger delta climbing gourami (Ctenopoma nebulosum) is known only from a few localities within the Niger River delta. Powell’s tetra (Neolebias powelli) is known only from a few rivers within the Niger River delta. The long-finned panchax (Epiplatys longiventralis) is known from a few swampy areas and streams within the lower Niger River drainage, where it is threatened by habitat destruction and degradation. Powell’s killifish (Fundulopanchax powelli) is known only from a single temporary pool on an unnamed island in the western part of the Niger delta. Arnold’s killifish (F. arnoldi) is known only from a few swampy areas and brooks within the Niger delta. The Niger delta catfish (Notoglanidium akiri) is known only from a few rivers within the Niger River delta. Buettikofer’s catfish (Parauchenoglanis buettikoferi) is known only from its original collection in the late nineteenth century in the Warri River, Niger River delta. The spotted squeaker catfish (Synodontis guttatus) is known only from its original collection in 1865 from an undefined locality within the Niger River delta.

The Guineo-Congolian Region

The Ogooué River The Ogooué (or Ogowe) River runs some 1200 km in WestCentral Africa. Its watershed drains nearly the entire country of Gabon, with some tributaries reaching into the Republic of Congo, Cameroon, and Equatorial Guinea. The Ogooué cichlid (Parananochromis ornatus) is confined to a few localities within the Ogooué/Ivindo drainage. Regan’s cichlid (Chromidotilapia regani) is known only from three localities with the Ogooué River drainage of Gabon. The Franceville citharine (Nannocharax ogoensis) is known only from its original collection in the early twentieth century from south-eastern Gabon, within the Ogooué River drainage. The Ogooué tetra (Neolebias kerguennae) is known only from three localities within the Ogooué/Ivindo drainage, where it is threatened by habitat destruction and degradation. The flame lampeye (Plataplochilus chalcopyrus) is confined to a few localities in the lower Ogooué River of north-western Gabon and southern Equatorial Guinea. It is threatened by loss of habitat. The chain lampeye (Hypsopanchax catenatus) is known only from a single rainforest brook within the Ogooué River drainage in south-eastern Gabon. The red-striped lampeye (Plataplochilus miltotaenia) is known from a few localities within the lower Ogooué and Komo rivers of north-western Gabon, where it is threatened by the expansion of Libreville. Terver’s lampeye (P. terveri) is confined to the upper Ogooué River drainage of southeastern Gabon, where it is threatened by pollution and sedimentation. Several species of killifish (Aphyosemion) are threatened by pollution and sedimentation resulting from deforestation. The caudal-stripe killifish (A. caudofasciatum) is known only from a single locality within the upper Ogooué River drainage of south-central Republic of Congo. The alpha killifish (A. alpha) is confined to a few coastal creeks between Libreville and Cap Esterias in north-western Gabon. The golden killifish (A. aureum) is confined to the upper Lolo and Yaou river drainages on the Du Chaillu Massif of southern Gabon. The Monts de Cristal killifish (A. etsamense) is known only from a few rainforest creeks within the Ogooué River drainage of northwestern Gabon. Hera’s killifish (A. hera) is known only from a single rainforest creek within the Ogooué River drainage of western Gabon. The Bateke killifish (A. rectogoense) is confined to the Bateke Highlands within the upper Lékoni-Djouya and upper M’passa River drainages of south-eastern Gabon. The Gabon shellear (Grasseichthys gabonensis) is known only from the Ogooué/Ivindo drainage of Gabon. The Ogooué mountain catfish (Amphilius caudosignatus) is confined to the south-eastern tributaries of the Ogooué River. The scissortail squeaker catfish (Synodontis acanthoperca) is known only from the Massoukou Rapids on the Ogooue River and another locality on the Louétsi River, both in southcentral Gabon.

The Ngomo snake catfish (Clariallabes brevibarbis) is known only from a single locality on the Ogooué River in central coastal Gabon. The Ogooué eel catfish (Channallabes ogooensis) is known only from specimens collected in the Franceville area of Gabon. The Ivindo River The Ivindo River is located in north-eastern Gabon. General threats include iron mining, commerical logging and slashand-burn agriculture. Axelrod’s cichlid (Parananochromis axelrodi) is known only from the Ivindo River drainage. The Ivindo barb (Enteromius condei) is known only from a single locality in the Ivindo River drainage. The Ivindo killifish (Aphyosemion fulgens) and the mosaic killifish (A. abacinum) are both known only from rainforest brooks and swamps within in the Ivindo River drainage, where they are threatened by habitat destruction and degradation. The Biale Creek mormyrid (Stomatorhinus ivindoensis) is known only from small forest streams in the Ivindo River drainage. The Ivindo eel catfish (Channallabes longicaudatus) is known only from the Ivindo River. The Lekoudi River The Lekoudi River is located in south-eastern Gabon. Tirbak’s killifish (Aphyosemion tirbaki) is known only from two rainforest brooks within the Lekoudi River drainage. The Ofoué River The Ofoué River is located in south-central Gabon. Passaro’s killifish (Aphyosemion passaroi) is known only from three small rainforest creeks flowing to the Onoy River, a tributary of the Ofoué River. The Okano River The Okano River is located in northern Gabon. The Okano upside-down catfish (Microsynodontis nasutus) is known only from rapids in the Okano River. The Ezanga River The Ezanga River is located in western Gabon. The Ezanga upside-down catfish (Microsynodontis notata) is known only from the Ezanga River. The Ngounié River The Ngounié River is located in western Gabon. Barton’s robber tetra (Brycinus bartoni) is known only from a few specimens collected mainly (or entirely) from the Ngounié River.

The Konkouré River The Konkouré River arises in the Fouta Djallon Highlands of west-central Guinea and flows into the Atlantic.

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The Konkouré tilapia (Tilapia rheophile) is a type of cichlid confined to the Konkouré River drainage. The Konkouré barb (Enteromius guineensis) and Cadenat’s barb (E. cadenati) are both confined to the upper Konkouré River drainage within the Fouta Djallon Highlands of western Guinea. Leveque’s trout barb (Raiamas levequei) is confined to the middle reaches of the Konkouré River and the Kakrima River. The Konkouré panchax (Epiplatys guineensis) is known only from a few streams within the middle Konkouré River drainage on the southern part of the Fouta Djallon Highlands. It is threatened by loss of habitat and pollution. Dekimpe’s squeaker catfish (Synodontis dekimpei) is confined to the Konkouré River drainage. The Kakrima River The Kakrima River is a major tributary of the Konkouré River located in western Guinea. Rouanet’s labeo (Labeo rouaneti) is confined to the Kakrima River. The Kakrima minnow (Leptocypris konkourensis) is confined to the Kakrima River. The Kakrima mountain catfish (Amphilius kakrimensis) is confined to the Kakrima River. The Samou River The Samou River is located in western Guinea. Leveque’s catfish (Chrysichthys levequei) is confined to the Samou River.

The Pra River The Pra River is located in south-western Ghana. Notable for its many cataracts, for most of its length it is not navigable even by canoe. Robert’s cichlid (Limbochromis robertsi) is confined to the upper tributaries of the Pra River drainage, where it is threatened by habitat degradation due to mining activities. Walker’s catfish (Chrysichthys walkeri) is confined to the Pra River drainage. The Subin River The Subin River is located in south-central Ghana. The Subin barb (Enteromius subinensis) is confined to the Subin River.

The Cavalla River The Cavalla River (also known as the Cavally River) originates in the mountains of Guinea and runs through Ivory Coast and Liberia before finally emptying into the Gulf of Guinea. The Cavalla cichlid (Chromidotilapia cavalliensis) is confined to the upper and middle reaches of the Cavalla River in Ivory Coast. The Cavalla barb (Enteromius traorei) is confined to the Cavalla River.

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Lamotte’s suckermouth catfish (Chiloglanis lamottei) is known only from its original collection in 1948.

The Cross River The Cross River (native name Oyono) is located in southeastern Nigeria and Cameroon. General threats include overfishing and deforestation. The Cross River cichlid (Gobiocichla ethelwynnae) is known only from two localities within the upper Cross River drainage. The redline pufferfish (Tetraodon pustulatus) is confined to two localities within the Cross River drainage (Korup National Park and the lower Cross River). Reid’s citharine (Nannocharax reidi) is known only from one or two localities in the upper Cross River drainage, but may be more widespread. Several species of killifish (Fundulopanchax) endemic to the Cross River drainage are threatened by deforestation and overcollection for the international aquarium trade. The miraculous killifish (F. mirabilis) is known only from a single locality within the upper Cross River drainage of western Cameroon. The Numba killifish (F. moensis) is confined to the Mo, Me, and Man rivers, tributaries of the upper Cross River in western Cameroon. Gresens’ killifish (F. gresensi) is confined to the Bagwor and Mack rivers, tributaries of the upper Cross River in western Cameroon. Scheel’s killifish (F. scheeli) is known only from a small area of south-eastern Nigeria within the Cross River drainage. The Mamfe blue killifish (F. gardneri mamfensis) is confined to the rainforest lowlands within the upper Cross River drainage of western Cameroon. Teugels’ bichir (Polypterus teugelsi) is known only from the upper Cross River drainage. Ahl’s catfish (Parauchenoglanis ahli) is known only from the upper Cross River drainage. The Mamfue River The Mamfue River is a tributary of the upper Cross River located in western Cameroon. The Mamfue killifish (Fundulopanchax intermittens) is known only from two localities within the Mamfue River. The Nguti River is located is western Cameroon. The Nguti cichlid (Etia nguti) is confined to the Nguti River. The Ngongolon River The Ngongolon River is located in the upper Cross River drainage. The Ngongolon killifish (Fundulopanchax traudeae) is confined to the Ngongolon River. The Calabar River The Calabar River is located in south-eastern Nigeria (Cross River state). The russet squeaker catfish (Synodontis robbianus) is known only from its original collection in 1875 from the Calabar River.

The Guineo-Congolian Region

The Nyong River The Nyong River is located in south-western Cameroon. The Nyong minnow (Prolabeops nyongensis) is known only from the Nyong River drainage. Heinemann’s killifish (Aphyosemion heinemanni) is confined to tributaries (forest brooks) of the Nyong River. The long-headed catfish (Parauchenoglanis longiceps) is known only from two localities within the Nyong River drainage. The Nyong catfish (Chrysichthys nyongensis) is known only from the Nyong River drainage. The Nyong butter catfish (Schilbe nyongensis) is known only from a single locality in the Nyong River.

The Sanaga River The Sanaga River is located in western and central Cameroon. It originates with the confluence of the Djérem and Lom rivers in the Adamawa Plateau, and flows some 600 km to the Gulf of Guinea. Jaeger’s yellowfish (Labeobarbus jaegeri) and the Mbam yellowfish (L. mbami) are both known only from the Sanaga River drainage. The Sanaga pygmy herring (Thrattidion noctivagus) is known only from a few specimens collected in 1972 from the lower Sanaga River. The Sanaga citharine (Nannocharax rubrolabiatus) is known only from three localities within the Sanaga River drainage. The Bamileke killifish (Aphyosemion bamilekorum) and the Mbam killifish (A. dargei) are both confined to the Sanaga River drainage, where they are threatened by habitat destruction and degradation as well as by overcollection for the international aquarium trade. The Sanaga mormyrid (Marcusenius sanagaensis) is confined to the Sanaga River drainage. The Sanaga catfish (Chrysichthys longidorsalis) is confined to the Sanaga River drainage. The Sanaga squeaker catfish (Synodontis rebeli) is confined to the Sanaga River drainage. The Mbam River The Mbam River is the main tributary of the Sanaga River. The Djim catfish (Notoglanidium depierrei) is only known from specimens collected in the Djim River, a small tributary of the Mbam River.

The Kouilou-Niari River The Kouilou-Niari River is located in south-western Gabon and south-western Congo. General threats include deforestation and mining activities. The Baudon barb (Enteromius kuiluensis), Roux’s barb (E. rouxi), and the chained barb (E. catenarius) are both known only from their original collection in the mid-twentieth century from the Kouilou-Niari drainage of southern Republic of Congo. The blackish-yellow barb (E. nigroluteus) is known only from three localities in south-western Democratic

Republic of Congo, where it is threatened by deforestation, mining pollution, and loss of habitat due to dam construction. The Kouilou citharine (Nannocharax altus) is known only from two localities within the Kouilou River drainage in southwestern Congo. Thys’ killifish (Aphyosemion thysi) is known only from the Louvisi and Niari rivers. Daget’s catfish (Chrysichthys dageti) is confined to a few localities within the Kouilou-Niari River drainage. The Louvisi River The Louvisi River (also known as the Louesse River) is located in south-western Congo. The Louvisi yellowfish (Labeobarbus semireticulatus) is known only from the Louvisi River. The Louvisi whiptailed catfish (Phractura gladysae) is known only from the Louvisi River. The Mpoulou River The Mpoulou River is located in south-western Congo. The Mpoulou catfish (Amphilius mamonekenensis) and the Dimonika catfish (A. dimonikensis) are both confined to the Mpoulou River, within the Dimonika Biosphere Reserve.

The Campo River The Campo River (also known as the Ntem River) originates in Gabon and flows through Equatorial Guinea and Cameroon before emptying into the Gulf of Guinea. The Campo catfish (Parauchenoglanis pantherinus) is known only from the Campo River drainage in southern Cameroon, where it is threatened by habitat destruction. The Bolo River The Bolo River (Rio Bolo in Portuguese) is located in Equatorial Guinea. The Bolo barb (Enteromius aloyi) is known only from the Bolo River. The Kyé River The Kyé River (Rio Kyé in Portuguese) is located in Equatorial Guinea. The Kyé upside-down catfish (Microsynodontis nannoculus) is known only from the Kyé River. The Nye River The Nye River is located in northern Gabon. The Nye upside-down catfish (Microsynodontis hirsuta) is confined to the Nye River.

The Cuanza River The Cuanza River (also known as the Kwanza River) is located in central Angola. Ansorge’s trout barb (Raiamas ansorgii) is known only from its original collection in the early twentieth century from the Cuanza River.

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Gulielm’s yellowfish (Labeobarbus gulielmi) is known only from its original collection in the early twentieth century from the Cuanza River. The Cuanza mormyrid (Petrocephalus cunganus) is known only from the lower Cuanza River. The Cuanza butter catfish (Schilbe angolensis) is known only from the upper Cuanza River. The Cuemba River The Cuemba River is located in central Angola. Greenwood’s barb (Enteromius greenwoodi) and the Cuemba barb (E. brevilateralis) are both known only from the Cuemba River. The Cokema River The Cokema River (also known as the Kokema River) is located in central Angola. The Cokema shellear (Parakneria marmorata) is confined to the Cokema River. The Cokema squeaker catfish (Synodontis laessoei) is confined to the Cokema River. The Cokema spiny eel (Mastacembelus ansorgii) is confined to the Cokema River. The Cunje River The Cunje River is located in central Angola. The Cunje yellowfish (Labeobarbus clarkeae) is known only from the Cunje River. The Lucala River The Lucala River is located in north-western Angola (Uíge, Malanje, and Cuanza Norte provinces). Girard’s yellowfish (Labeobarbus girardi) is confined to the Lucala River. Ansorge’s shellear (Kneria ansorgii) is confined to the Lucala River.

The Cuvo River The Cuvo River is located in west-central Angola (Cuanza Sul province). Wellman’s barbel (Enteromius wellmani) is confined to the upper reaches of the Cuvo River drainage. The Cuvo catfish (Amphilius lentiginosus) is known only from Mount Moko, within the Cuvo River drainage.

The Nyanga River The Nyanga River runs through southern Gabon and northern Republic of Congo. Stiassny’s whiptailed catfish (Phractura stiassny) is known only from a single collection in the Nyanga River at Tchibanga, Gabon. The Moukalaba River The Moukalaba River is located in southern Gabon.

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The Moukalaba cichlid (Chromidotilapia nana) is confined to the Moukalaba River.

The Chari River The Chari River (also known as the Shari River) is located in Central Africa. It runs for some 1400 km through from the Central African Republic and Cameroon to Lake Chad. The Chari mormyrid (Pollimyrus guttatus) is known only from a few specimens collected in the Chari River drainage. The Logone River The Logone River is located in northern Cameroon, eastern Nigeria, and south-western Chad. The Lancrenon Falls garra (Garra lancrenonensis) is known only from a single locality in the upper Logone River.

The Saint Paul River The Saint Paul River (also known as the Diani or Niandi River) is located in south-eastern Guinea and western Liberia. The Saint Paul tilapia (Coptodon coffea) is a type of cichlid confined to a single locality within the Saint Paul River of Liberia. The Monrovia panchax (Callopanchax monroviae) is known only from a single locality within the lower Saint Paul River of western Liberia. The Via River The Via River is located in north-western Liberia. It has been heavily impacted by deforestation, farming and mining activities. The black-banded barb (Enteromius melanotaenia) and the sharklike barb (E. carcharhinoides) are both known only from the Via River. Currie’s labeo (Labeo curriei) is known only from the Via River, where it was last collected in 1969.

The Mano River The Mano River is located in northern Liberia and southeastern Guinea. The Mano barb (Enteromius trispiloides) is known only from a single specimen collected in the 1980s from the Mano River. Jeanpol’s panchax (Nimbapanchax jeanpoli) is known only from the upper Mano River drainage.

Miscellaneous Lakes, Rivers, and Marshes Lake Edward is the smallest of the African Great Lakes. It is located on the border between the Democratic Republic of the Congo and Uganda, with its northern shore just a few kilometres south of the equator. It lies completely within the Virunga National Park (Democratic Republic of Congo) and the Queen Elizabeth National Park (Uganda), and does not have extensive human habitation on its shores. The Lake Edward cichlid (Haplochromis paradoxus) is confined to the lake.

The Guineo-Congolian Region

Lake Kivu lies on the border between the Democratic Republic of Congo and Rwanda. It empties into the Ruzizi River, which flows southwards into Lake Tanganyika. Along with Lake Nyos and Lake Monoun, one of only three lakes in the world that experiences limnic eruptions. Adolph Friedrich’s cichlid (Haplochromis adolphifrederici) is confined to Lake Kivu. Lake Kachira is a small lake located in western Uganda. The Kachira cichlid (Haplochromis commutabilis) and Schraml’s cichlid (H. ampullarostratus) are both known only from Lake Kachira. Lake Ruhondo (also know as Lake Luhondo) is a small lake located in north-western Rwanda. The Ruhondo yellowfish (Labeobarbus microbarbis) is known only from a single specimen collected from Lake Ruhondo in the 1930s. It is possible, although not certain, that the species was described based upon a hybrid and therefore not valid. Lake Barombi-ba-Kotto is a small crater lake located in western Cameroon. The Barombi-ba-Kotto tilapia (Coptodon kottae) is confined to Lake Barombi-ba-Kotto and the smaller nearby Lake Mboandong. The Corubal River is located in northern Guinea and Guinea-Bissau. The Koumba barb (Enteromius anniae) is known only from the Koumba River, a tributary of the Corubal River in north-western Guinea. The Senegal River is located in Senegal, Mauritania, Mali, and Guinea. The Senegal River barb (Enteromius ditinensis) is confined to the Bafing and Téné rivers in the upper Senegal River drainage of Guinea. The Lofa River (historically known as the Little Cape Mountain River) is located in Guinea and Liberia. Lauzanne’s barb (Enteromius lauzannei) is known only from the upper course of the Lofa River. The Dubreka River is located in Guinea. Gruvel’s yellowfish (Labeobarbus gruveli) is known only from a single specimen collected in the Dubreka River. The Farmington River is located in the Gibi Mountains of Liberia. The Farmington barb (Enteromius boboi) is known only from the Farmington River. The Kolenté River (also known as the Great Scarcies River) is located in Guinea and Sierra Leone. The Kindia killifish (Scriptaphyosemion cauveti) is confined to the Santa River, a tributary of the Kolenté River. The Bagbwe River is located in Sierra Leone. The Bagbwe barb (Enteromius bagbwensis) is confined to the Bagbwe River drainage. The Sewa River is located in Sierra Leone. The Bagbwe suckermouth catfish (Chiloglanis polyodon) is known only from a single specimen collected in the headwaters of the Bagbwe River, one of the sources of the Sewa River. The Cestos River (also known as the Nuon or Nipoué River) is located in central Liberia and western Ivory Coast. The Cestos tilapia (Tilapia cessiana) is a type of cichlid confined to the Cestos River. The Komoé River (also known as the Comoé River) is located in Burkina Faso and Ivory Coast. The Komoé tetra (Micralestes

comoensis) is confined to a few localities in the upper Komoé, where it is threatened by drought and deforestation. The Kogon River is located in western Guinea. Teugels’ electric catfish (Malapterurus teugelsi) is confined to the Kogon River. The Jong River (also known as the Taia River) is located in Sierra Leone. The Jong catfish (Notoglanidium maculatum) is confined to the Jong River. The Saint John River is located in Liberia. Ruhkopf’s panchax (Epiplatys ruhkopfi) is known only from the upper Saint John River in north-central Liberia. The Yewa River is located in Benin and Nigeria. Clausen’s barb (Enteromius clauseni) is known only from its type locality in the upper course of the Yewa River. The Ogun River is located in south-western Nigeria. The Ogun mormyrid (Marcusenius brucii) is known for certain only from the Ogun River, within Oyo Oyo National Park. The Kienké River is located in south-western Cameroon. The Kienké tetra (Phenacogrammus stigmatura) is confined to the Kienké River. The Mungo River is located in western Cameroon. Celia’s killifish (Aphyosemion celiae celiae) is known from a few localities within the Mungo River. The Lokoundje River is located in south-eastern Cameroon. The Lokoundje suckermouth catfish (Chiloglanis harbinger) is known only from a single locality within the Lokoundje River. The Little Scarcies River is located in central Guinea and northern Sierra Leone. The Little Scarcies barb (Enteromius foutensis) is known only from the Little Scarcies River. The Chiloango River (Rio Chiluango in Portuguese) is located in Angola, Democratic Republic of Congo, and Republic of Congo. The Chiloango spiny eel (Mastacembelus shiloangoensis) is confined to the Chiloango River drainage. The Bengo River is located in north-western Angola. The Bengo mormyrid (Heteromormyrus pauciradiatus) is confined to the Bengo River. The Cubal River is located in central coastal Angola. The Cubal shellear (Kneria sjolandersi) is known only from the Cubal River. The Ruzizi River is located in the Democratic Republic of Congo, Burundi, and Rwanda. The Ruzizi suckermouth catfish (Chiloglanis ruziziensis) is confined to rocky, fastflowing stretches of the Ruzizi River. The Komo River is located in north-western Gabon and southern Equatorial Guinea. The Komo killifish (Aphyosemion kouamense) is confined to the Komo River drainage. The Ecucu River is located in western Equatorial Guinea. The Ecucu killifish (Fundulopanchax avichang) is known only from small temporary pools within the Ecucu River drainage.

Coasts and Satellite Islands This section includes the West African coast from Gambia to the mouth of the Congo including the islands of the Gulf of Guinea.

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The Afrotropical Realm

The African manatee (Trichechus senegalensis) inhabits coastal marine, brackish, and freshwater areas of West Africa from Senegal to Angola, including estuaries, lagoons, lakes, mangrove creeks, and adjacent rivers. In some of the latter, for instance the Niger River system, it can be found in upper watercourses a considerable distance from the coast (such individuals often die of starvation due to the periodic drying up of the river, leaving them stranded). Individuals have also been found as far as 75 km offshore. Although widespread and abundant they have long been heavily exploited everywhere for their meat and ‘traditional medicines’, in spite of the fact that hunting them is illegal in every country where they occur. Individuals are also frequently caught by accident in fishing nets.

Bioko Bioko (formerly known as Fernando Po) is located 32 km off the coast of Cameroon. Part of the Cameroon Volcanic Line, the island used to be the end of a peninsula attached to the mainland but was cut off when sea levels rose 10,000 years ago at the end of the last ice age. This isolation has resulted in a number of endemic species and subspecies, most of which are now highly threatened. The Bioko drill (Mandrillus leucophaeus poensis) is a large terrestrial monkey confined to the south-western part of the island, where the population in 2006 was estimated to be less than 5000. The Bioko russet-eared guenon (Cercopithecus erythrotis erythrotis) is confined to Bioko.

Figure 3.4 African manatee (Trichechus senegalensis). (Credit: Joseph Wolf.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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The Guineo-Congolian Region

The Bioko red colobus (Piliocolobus pennantii) appears to be largely confined to south-western Bioko, where the total population is thought to be less than 5000. The Bioko black colobus (Colobus satanas satanas) is confined to Bioko. The Bioko needle-clawed galago (Euoticus pallidus pallidus) is confined to Bioko. The Bioko squirrel galago (Sciurocheirus alleni alleni) is confined to Bioko. Father Basilio’s striped mouse (Hybomys basilii) is known only from specimens collected in the early 1960s from highland areas. The Bioko forest shrew (Sylvisorex isabellae) is known with certainty only from a few specimens. The Bioko screeching frog (Arthroleptis bioko) was first described in 2010, and is known only from a handful of individuals. Oeser’s killifish (Fundulopanchax oeseri) is a freshwater fish known only from single rainforest locality in northern Bioko. Mountains Bioko is very mountainous and is made up of three overlapping shield volcanoes. The Bioko Preuss’ monkey (Allochrocebus preussi insularis) is known from Pico Basilé National Park and the Gran Caldera Scientific Reserve. The Bioko montane chameleon (Trioceros feae) is confined to montane forest, where it is rare and decreasing. Pico Basilé, located in north-central Bioko, is the highest peak on the island. It is protected within Pico Basilé National Park. Eisentraut’s mouse shrew (Myosorex eisentrauti) is known only from Pico Basilé above 2000 m. It has not been recorded since 1968. The Bioko white-eye (Zosterops brunneus) is a warbler-like bird known only from Pico Basilé, although it is possible that it could also occur on the nearby Caldera de Luba.

São Tomé São Tomé lies just north of the equator about 240 km off the north-western coast of Gabon. The entire island is a massive shield volcano that rises from the floor of the Atlantic Ocean, over 3000 m below sea level. The higher slopes of the island are forested and form part of Obo National Park. The São Tomé shrew (Crocidura thomensis) is confined to São Tomé. The São Tomé collared fruit bat (Myonycteris brachycephala) is known only from a few localities. The São Tomé free-tailed bat (Chaerephon tomensis) is known only from three specimens. The São Tomé scops owl (Otus hartlaubi) is confined to São Tomé. The São Tomé dwarf ibis (Bostrychia bocagei) is confined to a few areas of São Tomé, where it is extremely rare and heavily hunted.

Figure 3.5 São Tomé fiscal shrike (Lanius newtoni pictured below). (Credit: Henrik Grönvold.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The São Tomé olive pigeon (Columba thomensis) is confined to São Tomé. The São Tomé green pigeon (Treron sanctithomae) is confined to São Tomé, where it is threatened by hunting and habitat destruction. The São Tomé grosbeak (Crithagra concolor) was long known only from three specimens collected during the nineteenth century. Rediscovered in 1991, it remains extremely rare and is known only from a few localities. The São Tomé fiscal shrike (Lanius newtoni) was long known only from records in 1888 and 1928. Rediscovered in 1990, it remains very rare and localized. The São Tomé short-tail (Motacilla bocagii) is a type of passerine bird confined to central and southern São Tomé. The São Tomé giant sunbird (Dreptes thomensis) is confined to São Tomé. The São Tomé oriole (Oriolus crassirostris) is confined to São Tomé. The São Tomé reed frog (Hyperolius thomensis) is confined to primary forest remnants on São Tomé.

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Newton’s ridged frog (Ptychadena newtoni) is confined to wetland areas on São Tomé.

Príncipe Príncipe consists of a heavily eroded volcano surrounded by several smaller islands. Uninhabited when discovered by the Portuguese, the north and centre were thereafter converted into cocoa and sugar plantations which have since largely reverted to forest. The Príncipe olive ibis (Bostrychia olivacea rothschildi) was long thought to be extinct. However, it was reportedly sighted in 1991 and a population of perhaps less than 10 may still survive. The Príncipe thrush (Turdus xanthorhynchus) was long thought extinct until its rediscovery in small numbers in 1997. The Príncipe white-eye (Zosterops ficedulinus) is a passerine bird confined to the forests of Príncipe. The palm forest tree frog (Leptopelis palmatus) is known from a few localities on Príncipe.

Annobón Annobón is another extinct volcano lying about 180 km southwest of São Tomé. Long uninhabited, it has evolved considerable biological diversity despite its small size. The Annobón scops owl (Otus feae) is confined to higherelevation moist forests on Annobón, where the total population is thought to be between 50 and 250. The Annobón white-eye (Zosterops griseovirescens) is a type of passerine bird confined to forest areas on Annobón. The Annobón red-bellied paradise flycatcher (Terpsiphone rufiventer smithii) is confined to Annobón. The Annobón lidless skink (Afroablepharus annobonensis) is confined to dry forests on Annobón.

Miscellaneous Islands McCarthy Island (also known as Lemain Island or Janjanbureh Island) is located approximately 270 km up the Gambia River, in eastern Gambia. The Gambian blind snake (Myriopholis natatrix) is known only from McCarthy Island, where it has not definitely been recorded since before 1937. Tumbo Island is located off the coast of Guinea. The Tumbo screeching frog (Arthroleptis bivittatus) is known only from a single specimen collected from Tumbo Island.

Balance for the Guineo-Congolian Region Africa is considered by most paleoanthropologists to be the world’s oldest inhabited region, with humans having actually originated on the continent. The fossilized remains of several species of early ape-like humans have been discovered there dating back at least seven million years, which are believed to have evolved into modern humans (Homo sapiens sapiens) between 300,000 and 200,000 years ago. Since that time Africa has been continuously populated by groups of huntergatherers, some of whom began to migrate into first Europe

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and Asia and from there the rest of the globe around 70,000 years ago. Around 4000 B . C ., the Saharan climate started to become drier at an exceedingly fast pace. This caused lakes and rivers to shrink significantly and increased desertification, which in turn decreased the amount of land conducive to settlements. As a result, humans began to migrate ino the more tropical climate of West Africa. The pygmies of the Congo Basin are thought to be the direct descendants of Middle Stone Age hunter-gatherers. While livestock, fire, cultivation, and hunting have all had an impact on the environment, these factors have had a much shorter time to affect the GuineoCongolian Region than elsewhere in Africa, and indeed longterm climatic changes during thousands of years have had a significantly greater impact on its fauna and climate. European contact with Africa has been relatively brief. The Portugese first began to explore the West African coast in 1444, reaching what is now Sierra Leone by 1460. The latter were also the first to visit the islands of the Gulf of Guinea between 1472 and 1475, discovering the Congo River in 1482. However, unlike in Asia or the Americas, in Africa Europeans would be content to avoid the interior, instead setting up a system of coastal trading posts. True exploration would not resume until the late eighteenth century, when the Scottish explorer Mungo Park reached the upper Niger River, in what is now Mali, in 1796. A few years later, in 1805–06, he would descend that same river as far as the Bussa Rapids, where he was drowned. In 1798 the Brazilian-born Portuguese explorer Francisco de Lacerda travelled from Mozambique north-west to Lake Mweru in the Congo. In 1823 members of an Anglo-Scottish expedition became the first Europeans to sight Lake Chad, and in 1830 the English explorer Richard Lander and his brother John descended the Niger River for more than 640 km, from what is now northern Nigeria to the river’s mouth. In the 1860s the French explorer Paul Du Chaillu became the first European to confirm the existence of gorillas and, later, of pygmy tribes within the Congo Basin. During 1875–77 the Welsh-American journalist Henry Morton Stanley descended the Lualaba and Congo rivers to the sea. Years later, in 1887–89, Stanley traversed the Ituri Rainforest, explored the Ruwenzori and followed the Semliki River to its source, which he named Lake Edward. In 1884–85 almost all of Africa was formally divided up between the European colonial powers, who defined boundaries that are still largely intact today among post-colonial states. Apart from hunting little damage was done to the environment and, indeed, it was during this period that the first national parks and environmental protections were established. During the mid-twentieth century African colonies began to achieve independence. Their populations also begin to explode, creating in the process extreme poverty, disease, corruption, and almost continuous warfare, all of which have put intense pressure on wildlife and habitats. In recent historical time (i.e. since A . D . 1500), the GuineoCongolian Region has lost at least one subspecies of vertebrates (a mammal). Another three species are possibly extinct, and one species is currently extinct in the wild.

The African Region

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

~ species

1 species

~ species

143 species

1 subspecies

~ subspecies

~ subspecies

24 subspecies

1 taxon

1 taxon

~ taxa

167 taxa

~ species

1 species

~ species

62 species

~ subspecies

~ subspecies

~ subspecies

6 subspecies

~ taxa

1 taxon

~ taxa

68 taxa

~ species

1 species

~ species

50 species

~ subspecies

~ subspecies

~ subspecies

3 subspecies

~ taxa

1 taxon

~ taxa

53 taxa

~ species

~ species

~ species

142 species

~ subspecies

~ subspecies

~ subspecies

1 subspecies

~ taxa

~ taxa

~ taxa

143 taxa

~ species

~ species

1 species

558 species

~ subspecies

~ subspecies

~ subspecies

13 subspecies

~ taxa

~ taxa

1 taxon

571 taxa

~ species

3 species

1 species

955 species

1 subspecies

~ subspecies

~ subspecies

47 subspecies

1 taxon

3 taxa

1 taxon

1002 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

In addition, there are 955 species/47 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 143 species/24 subspecies are mammals, 62 species/6 subspecies are birds, 50 species/3 subspecies are reptiles, 142 species/1 subspecies are amphibians, and 558 species/13 subspecies are freshwater fishes.

The African Region The African Region includes all of sub-Saharan Africa with the exception of the Guineo-Congolian Region. It includes a few coastal islands along with the remote islands of the South Atlantic, although not Madagascar or the island groups of the western Indian Ocean (which are treated as a distinct realm). In general the climate is drier than that of the Guineo-Congolian Region. The African bush elephant (Loxodonta africana) is the largest living terrestrial animal. The species is still found widely across sub-Saharan Africa outside the rainforest belt, most typically in savanna and miombo areas, but this distribution is becoming highly fragmented. Whole populations have been devastated by the illegal ivory trade in recent years, as well as by trophy and bushmeat hunters. They are known to have

become extirpated from The Gambia in 1913, Burundi in the 1970s, Mauritania in the 1980s and Swaziland in 1920 (where they were reintroduced in the 1980s and 1990s). Loss of habitat and human–elephant conflict is also a concern. The black or hook-lipped rhinoceros (Diceros bicornis) was historically found throughout most of African south of the Sahara. While primarily a lowland savanna animal, it exhibits some ecological flexibility, including desert areas and montane savannas. For most of the twentieth century it was the most numerous of all rhinos but has long been mercilessly hunted for meat and trophies as well as for its horns, which are erroneously believed to have magical or medicinal properties and are used as well in the making of ceremonial Arab daggers. Around 1900 there were still probably several hundred thousand (and perhaps as many as 850,000), but by the late 1960s their numbers were reduced to an estimated 70,000. From there on the decline became much more dramatic. By 1981 only 10,000–15,000 remained, and in the early 1990s the total dipped below 2500. Since then, thanks mainly to increased protection of wild stocks, the number had recovered somewhat to 4880 by 2010. Nevertheless, at least three of the eight described subspecies are already extinct. The western black rhinoceros (D. b. longipes) once lived in South Sudan, northern Central African Republic, southern Chad, northern Cameroon, north-eastern Nigeria, and south-eastern Niger. The range possibly stretched west to the Niger River in western

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The Afrotropical Realm Figure 3.6 Western black rhinoceros (Diceros bicornis longipes). (Credit: M. Brunel.)

Niger, although this is unconfirmed. The last known wild specimens lived in northern Cameroon. In 2006 an intensive survey across its putative range in Cameroon failed to locate any, and in 2011 it was officially declared extinct. The quagga (Equus quagga quagga), a distinctive horse-like mammal with striped foreparts, was long considered to have been a distinct species but is now generally thought to have been a subspecies of the plains zebra (E. quagga). Historically it lived on the grasslands (now Karoo subdeserts) of the former Cape Province, south of the Orange and Vaal rivers, and west of the Drakensberg. In 1775 the quagga was still found near the coast of Cape Agulhas, and in 1812 herds of 30–50 roamed the plains between Fraserburg and Hanover, often in the company of white-tailed gnus (Connochaetes gnou) and South African ostriches (Struthio camelus australus). European farmers hunted the quagga heavily, using the meat to feed their African workers. In about 1865 they began to kill the animals for their hides as well, which were in great demand in the coast district and among the Boers for use as grain sacks. The last wild quagga was probably killed in 1878. Prior to that, several had been brought to Europe and kept in zoological gardens, but no serious attempts to breed them seem to have been made. The last one died in the Amsterdam Zoo on 12 August 1883. Only one was ever photographed alive, and only about two dozen skins are preserved today. In 1984 it became the first extinct animal to have its DNA analysed. The Quagga Project is currently trying to recreate the phonotype of hair pattern and related characteristics by the selective breeding of Burchell’s zebras (E. q. burchellii). The black wildebeest or white-tailed gnu (Connochaetes gnou) formerly ranged across the grassy veld and karoo steppes of Cape Province and on the high veld of the Orange Free State

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and the Transvaal, where at one time it was probably the most common of the larger ungulates to be found there. Even these vast numbers, however, could not withstand the terrible slaughter that began at the end of the eighteenth century, and probably not a single black wildebeest was left alive in the whole of the Transvaal by the end of 1885. It would certainly have been completely exterminated if not for some farmers in the Orange Free State, who carefully protected what was left of the few remaining herds. With the establishment of a system of protected areas in South Africa numbers began to rise, and by the end of the 1960s there were some 1800 living wild in South Africa. Today there are thought to be more than 16,000. Populations have been reintroduced to Swaziland and Lesotho where they had long been extirpated, as well as introduced outside their historic range into Namibia, where they have now become well established. Grévy’s zebra (Equus grevyi) is a large, distinct species with a long, narrow head, making it look more like a mule than a horse. An inhabitant of the semi-arid grasslands of the Horn of Africa, it suffered a substantial reduction in numbers owing to habitat degradation and hunting. It is now confined to parts of Ethiopia and northern Kenya, having been extirpated from south-western Somalia in the mid-1970s. The total population in 2014 was estimated at about 2680, down from an around 15,600 in the early 1980s, and 5800 a decade later. Two subspecies of mountain zebra (E. zebra) are largely endemic to the Great Escarpment of the Central Plateau (see below). The Cape mountain zebra (E. z. zebra) is the smallest and most geographically isolated of all existing zebras. Historically it was widely distributed along the mountain ranges that formed the southern and western edge of the Central Plateau, from Namaqualand to the Drakensberg, where it climbed on steep

The African Region Figure 3.7 Quagga mare in the London Zoo, 1870. The only one of her kind to be photographed alive. (Credit: Frederick York.)

slopes up to about 2100 m. It must have been the first zebra ever seen by Europeans, although apparently it was not particularly numerous when they began to settle in the Cape area because, as early as 1656, it was already being protected by Jan van Riebeek. Nevertheless, hunting continued. By 1937 the Mountain Zebra National Park was established on a farm in the eastern Cape Province, where there were just six zebras. In 1965 there were 57 animals in the national park and 75 in the whole of South Africa. Since then their numbers have been steadily increasing, although today surviving natural populations occur only in Mountain Zebra National Park, Gamka Mountain Reserve, and in the Kamanassie Mountains. Others have been reintroduced to various parts of their former range, mainly in national parks and nature reserves. Hartmann’s mountain zebra (E. z. hartmannae) occurs in the dry mountains between the Namib Desert and the central plateau in western Namibia, and possibly in Iona National Park in far south-western Angola. It remains relatively common despite the threat of hunting for its skins and farming activities such as fencing, which reduces its access to water. Populations have been introduced outside their natural range in the Western Cape and Eastern Cape, where they potentially pose a hybridization threat to the indigenous Cape Mountain zebra (E. z. zebra). The largest antelopes in the world are the elands. Two species are found in Africa, although only the giant eland (Taurotragus derbianus) is threatened at present. The eastern giant eland (T. d. gigas) is known for certain only from national parks in Cameroon, Chad, and the Central African Republic, but may still exist as well in parts of Sudan and northern Uganda. The western giant eland (T. d. derbianus) formerly occurred across West Africa, but has been

exterminated from most of its former range by overhunting. It is now reduced to a few hundred individuals living in protected areas in Senegal and Mali. The East African oryx (Oryx beisa) is divided into two subspecies, both of which are threatened by hunting and habitat destruction. The beisa oryx (O. b. beisa) once inhabited a large area of north-eastern Africa from north-eastern Sudan to the Tana River in Kenya, but is today largely confined to parts of Ethiopia and northern Kenya, with a small population in South Sudan. The fringe-eared oryx (O. b. callotis) is found south of the Tana River in south-eastern Kenya and northeastern Tanzania. The dibatag (Ammodorcas clarkei) is a rare type of gazelle confined to the Ogaden region of south-eastern Ethiopia and northern and central Somalia. It has disappeared from substantial parts of its former range, as for instance parts of northern Ogaden and northern Somalia, and is under heavy pressure elsewhere from armed pastoralists, overgrazing of livestock, and drought. Soemmerring’s gazelle (Nanger soemmerringii) is divided into at least three rather ill-defined subspecies found patchily within the arid steppes and grasslands of north-eastern Africa. The Sudan Soemmerring’s gazelle (N. s. soemmeringii) is confined to a small area of eastern coastal Sudan. The Borani Soemmerring’s gazelle (N. s. butteri) occurs in south-eastern Ethiopia. The Berber Soemmerring’s gazelle (N. s. berberana) is found disjunctly in northern Ethiopia, Eritrea, and coastal Somalia. The small population of ‘dwarf’ Soemmerring’s gazelles inhabiting Dahlak Kebir Island off the coast of Eritrea may also qualify as distinct. All are threatened by uncontrolled hunting and habitat destruction.

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Speke’s gazelle (Gazella spekei), the smallest of all the gazelles, historically inhabited much of the Horn of Africa in eastern Ethiopia and Somalia, but had already been much reduced in number by the late 1980s due to uncontrolled hunting, drought, and overgrazing of its habitat by domestic livestock. The species is now likely extirpated in Ethiopia, and continues to be seriously threatened in Somalia. Two subspecies of cheetah (Acinonyx jubatus), previously discussed, occur in sub-Saharan Africa within a wide range of habitats. Both are threatened by loss of habitat, hunting for their skins and persecution by farmers. The south-east African cheetah (A. j. jubatus) is found patchily in Angola, Botswana, Mozambique, South Africa, and Zambia. The north-east African cheetah (A. j. soemmeringii) is confined to a few small population pockets in northern Central African Republic, Chad, Ethiopia, and South Sudan. The so-called king cheetah is a rare genetic mutation for cream-coloured fur with large, blotchy spots, and dark stripes. The black-footed cat (Felis nigripes), the smallest wild cat in Africa, remains widespread within the arid areas of southern Africa (South Africa, Botswana, Namibia, and possibly southwestern Zimbabwe). It is everywhere rare, however, and threatened by loss of habitat and human persecution. The African wild dog (Lycaon pictus), previously discussed, is divided into a number of subspecies that are threatened by loss of habitat, human persecution, and infectious disease. The West African wild dog (L. p. manguensis) was historically widespread from Senegal to Nigeria, but now survives only in two disjunct populations totalling less than 100. The East African wild dog (L. p. lupinus) and the the Somali wild dog (L. p. somalicus) are both found patchily in eastern Africa. The Rondo dwarf galago (Paragalago rondoensis), one of the world’s most threatened primates, was historically found in both montane and lowland forest in eastern Tanzania. Extirpated from many areas due to loss of habitat, it is now known only from eight isolated forest patches. Temminck’s ground pangolin (Smutsia temminckii) is the most widespread of African pangolins, being found in a variety of habitats throughout much of eastern and southern Africa. It has suffered massive declines, however, due to ongoing exploitation for use as bushmeat and in ‘traditional medicine’, with increasing intercontinental trade to markets in Asia. The Berbera gerbil (Gerbillus acticola) is known only from a few individuals collected at various elevations in northwestern Somalia. The white-tailed rat (Mystromys albicaudatus) is a rare species found sporadically in grassland and scrub areas of South Africa and Lesotho. Duthie’s golden mole (Chlorotalpa duthieae) is known only from a few localities within a narrow band along the southern coast of South Africa (Western Cape and Eastern Cape). Heller’s pipistrelle bat (Pipistrellus aero) is known only from a few disjunct localities in central Kenya and possibly north-western Ethiopia.

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Parisi’s slit-faced bat (Nycteris parisii) is known only from three specimens collected from southern Somalia, southern Ethiopia, and northern Cameroon. Morris’ mouse-eared bat (Myotis morrisi) is known only from two widely separated localities (north-eastern Nigeria and western Ethiopia). Hamilton’s tomb bat (Taphozous hamiltoni) is known only from a very few records, mainly in east-central Africa (Sudan, Somalia, Kenya, and Tanzania), with an additional report from southern Chad. The Somali ostrich (Struthio molybdophanes) is still found across the Horn of Africa in Ethiopia, Somalia, Djibouti, and Kenya, although populations have been declining since the late 1980s due to hunting and overcollection of eggs. Ludwig’s bustard (Neotis ludwigii) is a large, terrestrial bird found over a wide area of south-western Africa (southwestern Angola, Namibia, south-western Botswana, and western and central South Africa). It is threatened mainly by collisions with power lines. The southern black bustard (Afrotis afra) is a type of large terrestrial bird that was historically common throughout southwestern South Africa (Northern Cape, Western Cape, and Eastern Cape). It has been much reduced by loss of habitat. The southern ground hornbill (Bucorvus leadbeateri) is a large, carnivorous species still widespread in eastern and southern Africa, but is everywhere declining due to habitat destruction and human persecution. The blue crane (Anthropoides paradiseus) is found throughout much of South Africa, with an additional disjunct breeding population in northern Namibia. The species has undergone a drastic decline since the 1970s due to habitat destruction, poisoning (both accidental and intentional), collisions with power lines and other factors. The African crowned cranes (Balearica) are everywhere threatened by wetland drainage, overgrazing of livestock, and pesticide pollution. The grey crowned crane (B. regulorum) is still found over a large area of eastern and southern Africa, where it is divided into two subspecies. The South African grey crowned crane (B. r. regulorum) is found from Mozambique south through Zimbabwe to South Africa and west to Namibia and Angola. The East African grey crowned crane (B. r. gibbericeps) occurs from eastern Democratic Republic of Congo, Rwanda, Uganda, and Kenya south through Tanzania to Mozambique. The black crowned crane (B. pavonina) is usually found within the shallow wetlands of sub-Saharan Africa during the breeding season, at other times ranging into dry savanna and near croplands. The nominate form (B. p. pavonina) occurs patchily in Senegal, Gambia, southern Mauritania, southern Mali, Burkina Faso, and northern Togo, while Cecilia’s black crowned crane (B. p. ceciliae) is found in southern Niger, southern Chad, northern Central African Republic, northern Nigeria, Sudan, South Sudan, and western Ethiopia. The martial eagle (Polemaetus bellicosus) is a very large species found throughout sub-Saharan Africa outside the rainforest belt. It is generally rare, however, and threatened by

The African Region

habitat destruction, persecution (shooting and trapping) by farmers, and other factors. The black harrier (Circus maurus) is a type of hawk found throughout South Africa and Namibia. The species is everywhere rare and largely dependent upon protected areas. The Cape vulture (Gyps coprotheres) is found in the open country of southern Africa, where it nests on cliffs. It has been declining dramatically everywhere since at least the 1980s due mainly to the decrease in the available amount of large carrion, poisoning (often targeted), electrocution or collision with power lines, and loss of habitat. The fynbos buttonquail (Turnix hottentottus) is confined to a small area of lowland and montane scrub in southern South Africa (Western Cape and Eastern Cape), where it is threatened by habitat destruction. Fischer’s lovebird (Agapornis fischeri) is a type of small parrot native to northern and north-western Tanzania, although during times of drought flocks will temporarily migrate west into Rwanda, Burundi, and Uganda. Very common up until the 1970s, it has since undergone a considerable population decline due to trapping for the international cage-bird trade. The species has been introduced into central Kenya, and wild birds have also been observed in south-eastern France, Florida, Puerto Rico, and the U.S. Virgin Islands, although these appear to be escaped pets and not viable breeding populations. The Somali pigeon (Columba oliviae) is a rare species confined to coastal north-eastern Somalia. The Amani sunbird (Hedydipna pallidigaster) is found patchily in south-eastern Kenya and in central and northeastern Tanzania, in both lowland and montane forest pockets. It is threatened by loss of habitat. The Sokoke pipit (Anthus sokokensis) is a type of passerine bird found patchily in both dry and moist lowland forest in south-eastern Kenya and north-eastern Tanzania. Benson’s papyrus warbler (Calamonastides bensoni) is confined to Lake Mweru, at the mouth of the Luapula River in Zambia and to a few adjacent swampy areas in Democratic Republic of Congo. The thin-billed papyrus warbler (C. gracilirostris) has a severely fragmented range within the Great Lakes region of Central Africa, occurring in western Kenya, western Uganda, eastern Democratic Republic of Congo, Rwanda, Burundi, and north-western Tanzania. Both are threatened by loss of their wetland habitat. The white-winged apalis (Apalis chariessa) is a type of warbler with a highly disjunct range in East Africa, where it is divided into two subspecies (one of which is now extinct). Macpherson’s white-winged apalis (A. c. macphersoni) is known from the Uluguru and Udzungwa mountains of Tanzania, Mount Chiperone, Mount Mulanje, and the Namuli Massif in Mozambique, and a few localities in southeastern Malawi. The Karamoja apalis (A. karamojae) is known disjunctly from north-eastern Uganda and northern Tanzania. The African spurred tortoise (Centrochelys sulcata), one of the largest tortoises in the world, is confined to the southern edge of the Sahara Desert as well as the Sahel.

The pancake tortoise (Malacochersus tornieri) lives in semi-arid and savanna woodland areas of southern Kenya and eastern Tanzania, with an additional, introduced population possibly living in Zimbabwe. Thanks to its flat shell it is able to take refuge in narrow rocky crevices and under stones where, when disturbed, it can inflate the body to such a size that it is almost impossible to drag it out from its shelter. The species is threatened by habitat destruction and overexploitation by the international pet trade. The Dutumi house gecko (Hemidactylus tanganicus) is known only from two localities in Tanzania (Dutumi and Mkomazi national parks). Scheffler’s dwarf gecko (Lygodactylus scheffleri) is known only from two localities in south-eastern Kenya in the vicinity of the Chyulu Hills and Kibwezi, where it was last recorded in the 1930s. There is an additional disjunct record from an undefined locality in north-central Tanzania. Pickersgill’s reed frog (Hyperolius pickersgilli) is confined to coastal eastern South Africa (KwaZulu-Natal). The spotted shovel-nosed frog (Hemisus guttatus) is found patchily in eastern coastal South Africa (KwaZulu-Natal and Mpumalanga) and possibly Swaziland. Bilbo Baggins’ rain frog (Breviceps bagginsi) is confined to three disjunct areas in eastern coastal South Africa (Eastern Cape and KwaZulu-Natal). The Changamwe caecilian (Boulengerula changamwensis) is known only from a small number of specimens collected in southern Kenya and Malawi.

Mountains and Highlands The African Region includes a number of mountainous areas, the most important of which include the Ethiopian Highlands, the Eastern Rift Mountains and Eastern Arc Mountains, the Southern and Eastern Highlands, the Central Plateau, and the Cape Fold Mountains. All provide a wide range of habitats and very often a last refuge for many species. The highland crested mangabey or kipunji (Rungwecebus kipunji) is a type of monkey known only from two populations separated by some 350 km of non-forested land. The first occurs in the high-elevation Rungwe-Kitulo Forest in the Southern Highlands of south-western Tanzania. The latter includes the Mount Rungwe Forest Reserve and the Livingstone Forest, both of which lie within Kitulo National Park. The other population lives in the Vikongwa Valley, Ndundulu Forest, within the Kilombero Nature Reserve of the Udzungwa Mountains in south-central Tanzania. Simultaneously discovered by two separate research teams in 2003–2004, the species is highly threatened by loss of habitat and numbers considerably less than 2000 individuals in total. The mountain dwarf galago (Paragalago orinus) is found sporadically throughout the Eastern Arc Mountains and Southern Highlands of central and north-eastern Tanzania, where it is threatened by loss of habitat.

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Two subspecies of mountain reedbuck (Redunca fulvorfula), previously discussed, are threatened by loss of habitat and hunting. Chanler’s mountain reedbuck (R. f. chanleri) is found in the mountains of south-eastern Sudan, Ethiopia, Uganda, Kenya, and northern Tanzania. The southern mountain reedbuck (R. f. fulvorufula) occurs in South Africa, Lesotho, Swaziland, Botswana, and Mozambique. Abbot’s duiker (Cephalophus spadix) is a rare and elusive type of small antelope known only from a few scattered populations across Tanzania, specifically in the Eastern Arc Mountains, the Southern Highlands, and on Mount Kilimanjaro. It is threatened by snaring and habitat destruction. The Tanzanian groove-toothed rat (Otomys lacustris) is confined to isolated populations in extreme northern Malawi, south-central Tanzania, and south-western Kenya, with an additional record from northern Zambia. De Winton’s African dormouse (Graphiurus angolensis) is known only from a few localities in Angolia and western Zambia. Vernay’s climbing mouse (Dendromus vernayi) is known only from its original collection in the 1930s from east-central Angola. The giant golden mole (Chrysospalax trevelyani) is the largest and rarest golden mole species, being confined to a small area of montane forest in south-eastern South Africa (Eastern Cape). Kihaule’s mouse shrew (Myosorex kihaulei) is confined to the Udzungwa Mountains and Mount Rungwe in southcentral Tanzania. The desperate shrew (Crocidura desperata) is confined to relict forest patches within the Udzungwa Mountains and Mount Rungwe in southern Tanzania. The Taita falcon (Falco fasciinucha) is a small species found patchily in east-central Africa, chiefly in gorges and escarpments up to 3800 m where it uses cliffs for nesting. As a result of this specialization it is everywhere rare, although it appears to have formerly been associated with human settlements at lower elevations where it was often hunted. The Djibouti francolin (Pternistis ochropectus) is a type of terrestrial gamebird confined to montane cedar forests in the Goda and Mabla mountains of Djibouti. It is threatened by loss of habitat. Belcher’s spotted ground thrush (Geokichla guttata belcheri) is confined to the mountains of southern Malawi. The Thyolo alethe (Chamaetylas choloensis) is a type of passerine bird confined to a few scattered montane localities in southern Malawi and north-western Mozambique. The Namuli apalis (Apalis lynesi) is a type of passerine bird known only from Mount Namuli and Mount Mabu in northern Mozambique. The yellow-throated apalis (A. flavigularis) is confined to Mount Mulanje, Mount Zomba, and Mount Malosa in southern Malawi (Southern region). Both are threatened by loss of habitat. The blue swallow (Hirundo atrocaerulea) is a migratory species found throughout the mountains of Eastern Africa

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from Uganda to South Africa, but confined for breeding purposes to a few widely separated areas on montane grassland. Archer’s lark (Heteromirafra archeri) is known only from two small localities within the highlands of eastern Ethiopia and (formerly at least) north-western Somalia, where it is seriously threatened by loss of habitat. The dapple-throat (Arcanator orostruthus) is a type of passerine bird divided into three subspecies. The nominate form (A. o. orostruthus) is confined to a few scattered populations in north-eastern Mozambique. Swynnerton’s robin (Swynnertonia swynnertoni) is divided into three subspecies spread over a few small and scattered localities. The Mozambique Swynnerton’s robin (S. s. umbratica) is known only from Mount Gorongosa and Mount Mabu in central and north-eastern Mozambique. The sombre rockchat (Oenanthe dubia) is a passerine bird known only from the highlands of east-central Ethiopia and north-western Somalia. The Iringa akalat (Sheppardia lowei) is a type of flycatcher found patchily in montane forest fragments of the Eastern Arc Mountains and Southern Highlands of south-central Tanzania. The Namuli spiny pygmy chameleon (Rhampholeon tilburyi) is confined to a few isolated mountains in northern Mozambique (Zambézia and Nampula provinces). The Lolui mabuya skink (Trachylepis loluiensis) is confined to high-elevation areas on Lolui Island, in Lake Victoria. The Mpwapwa worm lizard (Chirindia mpwapwaensis) is known only from two specimens collected in east-central Tanzania. The Zambézia bush viper (Atheris mabuensis) is confined to Mount Mabu and Mount Namuli in northern Mozambique. The Udzungwa toad (Mertensophryne uzunguensis) is confined to the Udzungwa Mountains and the Southern Highlands of south-central Tanzania. The Mashonaland toad (M. anotis) occurs in the Chirinda Forest of eastern Zimbabwe and in the Dombé Forest of adjacent western Mozambique. An additional record from Quirimbas National Park in north-eastern Mozambique may suggest either a much wider distribution, or perhaps represents an as-yet undescribed species. The peacock tree frog (Leptopelis vermiculatus) and the large-headed forest tree frog (L. grandiceps) are both found patchily in the mountains of eastern and central Tanzania. The Rungwe big-fingered frog (Probreviceps rungwensis) occurs patchily in the Udzungwa and Mahenge mountains and the Southern Highlands of south-central Tanzania. The tiny reed frog (Hyperolius minutissimus) is confined to the Udzungwa Mountains and the Southern Highlands of south-central Tanzania. The Eastern Highlands reed frog (H. inyangae) is known only from Nyanga National Park in eastern Zimbabwe and from the Kaningina Forest Reserve in northern Malawi. The spiny-throated reed frog (H. spinigularis) is known from the Mulanje Massif in southern Malawi and from the Namuli Massif in adjacent northern Mozambique.

The African Region

Broadley’s ridged frog (Ptychadena broadleyi) is known only from the foothills of Mount Mulanje and from Mount Zomba, in southern Malawi. France’s screeching frog (Arthroleptis francei) is known only from the Mulanje Massif in southern Malawi and from Mount Mabu and Mount Namuli in northern Mozambique. Denhardt’s caecilian (Boulengerula denhardti) is known only from a small area of mid-altitude forest in central Kenya, where it is threatened by agricultural expansion and pesticides.

The Ethiopian Highlands The Ethiopian Highlands consist of two main mountainous regions divided by the Great Rift Valley, which taken together comprise the largest continuous high-altitude area on the African continent. Rugged and with little of the surface falling below 1500 m, they are sometimes called the Roof of Africa. The Ethiopian wolf (Canis simensis) is endemic to the Ethiopian Highlands. Up until the 1960s this species was common in the Simien Mountains and elsewhere, after which it was accused without justification of being a sheep-killer and a relentless persecution was initiated. In a few years it had become one of the rarest canids in the world, and is now confined to the higher elevations of seven isolated mountain ranges. Two subspecies are recognized. The northern Ethiopian wolf (C. s. simensis) occurs in the north-western Rift Valley (Simien Mountains, Mount Guna, and the Wollo Highlands). The southern Ethiopian wolf (C. s. citernii) is found in the southeastern Rift Valley (Arsi and Bale Mountains). The black-clawed brush-furred rat (Lophuromys melanonyx) is known from a few specimens collected from central Ethiopia. The species has been extirpated from west of the Rift Valley, and now appears to be confined to a few areas of highaltitude grassland further south. The Guramba shrew (Crocidura phaeura) is known only from a few localities within the Ethiopian Highlands. Scott’s mouse-eared bat (Myotis scotti) is confined to a few localities on either side of the Rift Valley in Ethiopia, where it is likely threatened by loss of its forest habitat. The Ethiopian long-eared bat (Plecotus balensis) is known only from a few localities within the Ethiopian Highlands. The blue-winged goose (Cyanochen cyanoptera) is confined to rivers and lakes within the Ethiopian Highlands, where it is threatened by habitat destruction and hunting. The Shoa forest tree frog (Leptopelis ragazzii) is confined to a few scattered localities on both sides of the Rift Valley, where it is threatened by the loss of its montane forest habitat. The Ethiopian leaf-folding frog (Afrixalus enseticola) is confined to a few scattered localities within the Ethiopian Highlands. Largen’s puddle frog (Phrynobatrachus inexpectatus) is known only from three scattered localities within the Ethiopian Highlands. Largen’s clawed frog (Xenopus largeni) is an aquatic species found patchily in the Ethiopian Highlands.

The Western Ethiopian Highlands The Western Ethiopian Highlands are located in southwestern, west-central, and north-western Ethiopia (Tigre and Amhara regions), extending into Eritrea. The Ethiopian groove-tailed rat (Mylomys rex) is known only from a single skin collected from the Charada Forest in central Ethiopia. Yalden’s rat (Desmomys yaldeni) is confined to a small area in south-western Ethiopia encompassing the Sheko Forest. The Ethiopian woolly bat (Kerivoula eriophora) is known only from its original collection in the late nineteenth century from northern Ethiopia. The Ankober serin (Crithagra ankoberensis) is a type of finch known only from a few localities in the highlands of central and northern Ethiopia. The Eritrean blind snake (Letheobia erythraea) is known only from two localities in the arid central highlands of Eritrea. Susana’s forest tree frog (Leptopelis susanae) is known only from the Gughe Mountains and from Saja Forest in south-western Ethiopia. Yalden’s forest tree frog (L. yaldeni) is confined to a few localities in north-western Ethiopia. Clarke’s leaf-folding frog (Afrixalus clarkei) is known only from two small areas in south-western Ethiopia. Wade’s ridged frog (Ptychadena wadei) is known only from a small area of the North-eastern Highlands. The Simien Mountains are located in north-western Ethiopia (Amhara region). Its summit, Ras Dashen, is the highest peak in Ethiopia. The important Simien Mountains National Park was established in 1970. The northern gelada (Theropithecus gelada gelada) is a large, terrestrial monkey confined to high-altitude Afroalpine grasslands adjacent to deep gorges and cliffs around the Simien Massif. It is threatened by loss of habitat and hunting. The walia ibex (Capra walie) is the southernmost of all ibexes and seems to have been long found only in the Simien Mountains, where it was formerly widespread. It was heavily hunted, however, and since at least the mid-twentieth century has been mostly restricted to a 4250-m escarpment on the northern side. In 1970 the Simien Mountains National Park was created here specifically to protect the last remaining walia ibex, then estimated to number around 150. Today the total population is estimated at around 500, both within the park and at four small localities outside its borders. Habitat destruction and degradation is now the main threat. The Eastern Ethiopian Highlands The Eastern Ethiopian Highlands are located in southcentral Ethiopia (Oromia region). They are further subdivided by two mountainous blocks separated by the Shabelle River Valley. The Djaffa Mountains guereza (Colobus guereza gallarum) is restricted to the Ethiopian Highlands east of the Rift Valley. The mountain nyala (Tragelaphus buxtoni) is a type of antelope found only in the highlands of southern Ethiopia south and east of the Rift Valley. It has been eliminated from

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much of its former range by hunting and human disturbance. Currently, the majority live in Bale Mountains National Park and on the eastern escarpments of the Bale Massif. Smaller relict populations occur in the Amhar Mountains, the Arsi Mountains, and on the Somkaro-Kurduro ridge. The total number was estimated at between 7000 and 12,500 in the 1960s, falling to 2000–4000 by the 1980s. Numbers are believed to have declined since then. Nikolaus’ mouse (Megadendromus nikolausi) is known only from a few specimens collected in the Bale and Chilalo Mountains. Lucina’s shrew (Crocidura lucina) is known only from a few localities in the Bale and Arsi mountains. The Bale Mountains shrew (C. bottegoides) occurs in the Harenna Escarpment and on Mount Albasso. Both are threatened by loss of habitat due to wood collection and fires. Prince Ruspoli’s turaco (Tauraco ruspolii) inhabits midelevation dry forest and savanna areas in south-central Ethiopia. It has suffered population declines due to habitat degradation and hybridization with the related white-cheeked turaco (T. leucotis). The Ethiopian bushcrow (Zavattariornis stresemanni) is a micro-habitat specialist confined to a small area in southern Ethiopia (Oromia region). The white-tailed swallow (Hirundo megaensis) is confined to scrubby areas in south-central Ethiopia. Salvadori’s seedeater (Crithagra xantholaema) is a type of finch confined to scrubby areas and high-altitude dry forests of the south-eastern Ethiopian Highlands. Osgood’s Ethiopia toad (Altiphrynoides osgoodi) is known only from the south-central Ethiopia. The species has not been recorded since 2003 despite intensive surveys, and may be extinct. The Ethiopian ridged frog (Ptychadena nana) is known only from two localities in the Bale and Arsi mountains. The Kouni Valley striped frog (Paracassina kounhiensis) is confined to a few scattered localities in the Mendebo and Ahmar mountains. The Mendebo Mountains are located south of the Shabelle River Valley in south-central Ethiopia. The important Bale Mountains range, located within the north-western part, are nearly as high as the Simien Mountains and include several peaks over 4000 m. They are protected within Bale Mountains National Park. The Harenna Forest, located in the foothills of the Bale Mountains, is one of the few remaining forest areas in Ethiopia. The Bale Mountains monkey (Chlorocebus djamdjamensis) is a terrestrial species largely confined to high-elevation bamboo forests within the Bale Mountains and the Sidamo Highlands. The large-headed mole-rat (Tachyoryctes macrocephalus) is confined to high-elevation grasslands in the Bale Mountains. The Harenna shrew (Crocidura harenna) is known only from the Harenna Forest. Malcolm’s Ethiopia toad (Altiphrynoides malcolmi) is confined to the Bale Mountains.

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The Harenna tree frog (Balebreviceps hillmani) is confined to the Harenna Forest. The Harenna ridged frog (Ptychadena harenna) is confined to the Harenna Forest. The Bale Mountains frog (Ericabatrachus baleensis) is confined to a single tiny area of montane forest in the Bale Mountains. It is threatened by loss of habitat and perhaps chytrid fungus. The Ethiopian Rift Valley The Ethiopian Rift Valley (also known as the Great Rift Valley) is a branch of the East African Rift that runs through Ethiopia in a south-west direction. It is notably arid and contains a number of salt lakes. The Ethiopian thicket rat (Grammomys minnae) is known only from two localities within the Rift Valley of Ethiopia. The Nechisar nightjar (Caprimulgus solala) is known for certain only from a single specimen collected in 1990 from the Nechisar Plains in south-western Ethiopia. There have been a few unconfirmed sightings since then. The yellow-throated seedeater (Crithagra flavigula) is a type of finch long known only from three specimens collected during the nineteenth century from central Ethiopia. It was rediscovered in very small numbers in 1989 at the same area, and has since been recorded from two other localities. The Langano toad (Sclerophrys langanoensis) is known only from the northern Rift Valley of Ethiopia, where it has been recorded from Lake Langano and from Awash National Park. The hot springs ridged frog (Ptychadena filwoha) is known only from two closely adjacent springs and pools within the northern Rift Valley of Ethiopia. The Gughe Mountains are located in south-western Ethiopia, within the Great Rift Valley. Rupp’s mouse (Stenocephalemys ruppi) is confined to the Gughe Mountains.

The Northern Somalia Highlands The northern Somalia Highlands are located in northern coastal Somalia, in the Horn of Africa. The Somali thrush (Turdus ludoviciae) is confined to mountaintop woodlands in the Northern Somalia Highlands. It remains relatively common in the Daalo Forest and on Mount Wagar. Sharpe’s lark (Mirafra sharpii) is known only from northwestern Somalia, though it may possibly extend into adjacent areas of eastern Ethiopia. The Warsangli linnet (Linaria johannis) is a type of finch known only from two small areas of the Northern Somalia Highlands.

The Eastern Rift Mountains The Eastern Rift Mountains are associated with the East African Rift Valley in Kenya, Tanzania, Sudan, South Sudan, and Uganda. Volcanic in origin, they include some of the largest peaks in Africa.

The African Region

The eastern or mountain bongo (Tragelaphus eurycerus isaaci) is a large and spectactular forest antelope confined to a few isolated populations in the forested mountains of central Kenya, including Mount Kenya, Mount Elgon, the Aberdare Range, Mau Forest, and the Cherengani and Chepalungu hills. The survival of the remaining wild population of around 150 is dependent upon more effective protection of remaining forest patches. Fortunately, the subspecies is well established in captivity. The East African highland shrew (Crocidura allex) is found patchily in highland areas of south-central Kenya and northern Tanzania, where it is threatened by habitat destruction. Sharpe’s longclaw (Macronyx sharpei) is a type of passerine bird confined to a few localities in the highlands of western and central Kenya. Abbott’s starling (Poeoptera femoralis) is confined to a few montane forest localities in south-central Kenya and northern Tanzania. Hinde’s babbler (Turdoides hindei) is confined to a few localities in south-central Kenya. The Eastern Rift sentinel chameleon (Kinyongia excubitor) is found in the Aberdare Range as well as on Mount Kenya and the Nyambeni Hills. It is threatened by loss of habitat and perhaps collection for ‘traditional medicine’. The Mount Elgon day gecko (Cnemaspis elgonensis) is confined to eastern Uganda and adjacent Kenya. Schmidt’s bold-eyed tree snake (Thrasops schmidti) is confined to highland forest fragments in central Kenya. The Eastern Rift worm snake (Leptotyphlops keniensis) is known only from Mount Kenya and environs, and from Mount Kilimanjaro in Tanzania. Lönnberg’s toad (Mertensophryne lonnbergi) is found in scattered localities in the highlands on either side of the Great Rift Valley in south-central Kenya, including Mount Kenya. While relatively common and adaptable, it is threatened by loss of habitat. Mocquard’s toad (M. mocquardi) is confined to Mount Kenya, the Kanangop Plateau, and the highlands surrounding Nairobi. The Kinangop puddle frog (Phrynobatrachus kinangopensis) and the Irangi puddle frog (P. irangi) are both known only from a few localities in south-central Kenya, where they are threatened by loss of habitat. The Nuba Mountains The Nuba Mountains are located in southern Sudan (South Kordofan). Wilson’s snake-eyed skink (Panaspis wilsoni), one of the smallest of all the skinks, is known only from a single specimen collected in the southern Nuba Mountains during the early twentieth century. The Imatong Mountains The Imatong Mountains are located in southern South Sudan and northern Uganda.

Rupp’s climbing mouse (Dendromus ruppi) is known only from the Imatong Mountains. Mount Kineti is the highest peak in the Imatong Mountains. The Kineti chameleon (Trioceros kinetensis) is known only from Mount Kineti. The Didinga Hills The Didinga Hills are located in south-eastern South Sudan. The Didinga Hills guereza (Colobus guereza dodingae) is a type of leaf-eating monkey confined to the Didinga Hills. Mount Nyiro Mount Nyiro is located within the Ndoto Range, south of Lake Turkana in north-western Kenya (former Rift Valley province). Although surrounded by desert, its upper slopes are forested. Macow’s shrew (Crocidura macowi) is a little-known species known only from Mount Nyiro. The Nyiru chameleon (Trioceros ntunte) is confined to the montane moorlands of Mount Nyiru. The Aberdare Range The Aberdare Range (formerly known as the Sattima Range) is a 160-km stretch of mountains located in west-central Kenya (former Central and Rift Valley). Aberdare National Park was established in 1950. The ultimate shrew (Crocidura ultima) is known only from its original collection in the early twentieth century from the Jombeni Mountains. The Aberdare cisticola (Cisticola aberdare) is a warblerlike bird confined to high-elevation grassland areas of the Aberdare Range. The Matthews Range The Matthews Range (known locally as the Lenkiyio Hills) is located in central Kenya (former Rift Valley province). It stretches for about 150 km in a north–south orientation. Percival’s guereza (Colobus guereza percivali) is confined to the forested slopes of Mount Gargues, where it continues to be hunted for its beautiful fur. Rainey’s shrew (Crocidura raineyi) is known only from lower elevations on Mount Gargues, where it has not been recorded since 1912. Mount Kenya Mount Kenya is located in central Kenya (former Central province). The second highest mountain in Africa after Kilimanjaro, it along with the surrounding forest was designated as a national park in 1949. The Kenyan oribi (Ourebia ourebi kenyae), a type of small antelope, formerly occurred on the lower slopes of Mount Kenya but is now extinct. The Mount Kenya potto (Perodicticus ibeanus stockleyi) is a type of nocturnal prosimian known only from a single

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specimen collected in 1938 from montane forest on Mount Kenya. It is most likely extinct. The giant thicket rat (Grammomys gigas) is confined to montane forest and shrubland on Mount Kenya. The Mount Kenya mole shrew (Surdisorex polulus) is known only from two or three specimens collected on Mount Kenya. Drews’ worm snake (Epacrophis drewsi) is known only from a single specimen collected on Mount Kenya in 1957. Mount Elgon Mount Elgon is an extinct shield volcano located on the border between Uganda and Kenya. Barbour’s groove-toothed rat (Otomys barbouri) is confined to high-elevation grasslands on Mount Elgon. The Elgon mole shrew (Surdisorex schlitteri) is known only from a single specimen collected on Mount Elgon in 1984. Du Toit’s rocky river frog (Arthroleptides dutoiti) is confined to montane forest streams on Mount Elgon, where it was last recorded in 1962. Mount Kilimanjaro Mount Kilimanjaro is located in northern Tanzania near the border with Kenya, within Kilimanjaro National Park. A dormant stratovolcano with a distinctive, snow-covered peak, it is the highest mountain in Africa. The Kilimanjaro stream frog (Strongylopus kilimanjaro) is known only from three specimens collected in 1936 from the middle slopes of Mount Kilimanjaro. Mount Kitumbeine Mount Kitumbeine is an extinct volcano located in northern Tanzania (Arusha region). The Kitumbeine stream frog (Strongylopus kitumbeine) is confined to Mount Kitumbeine.

The Eastern Arc Mountains The Eastern Arc Mountains are the oldest in East Africa, having been created at least 100 million years ago. They form a roughly crescent-shaped arc of isolated mountains running from south-eastern Kenya through much of Tanzania, which, although physically separated from each other, share a similar geomorphology and ecology. Some 30 million years ago the entire area was covered by extensive rainforest, but when the climate cooled and dried some 10 million years ago the lowland forests were converted to savanna, leaving the mountain ranges as ‘sky islands’ where the tropical forests continued to survive. This isolation has led to a great deal of endemism and a very diverse flora and fauna. Lowe’s servaline genet (Genetta servalina lowei) was long known only from a single skin collected in the Udzungwa Mountains in 1932. Rediscovered in 2000, it has since been caught on camera traps in other areas of the Eastern Arc Mountains, where it is threatened by hunting and habitat destruction.

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Two species of shrew (Crocidura) endemic to the Eastern Arc Mountains are threatened by loss of their montane forest habitat. The Usambara shrew (C. usambarae) is confined to the Usambara and South Pare mountains in north-eastern Tanzania. Telford’s shrew (C. telfordi) is confined to parts of the Uluguru and Udzungwa mountains of east-central Tanzania. Vosseler’s eagle-owl (Bubo poensis vosseleri) is a rare subspecies known from the Usambara and Uluguru mountains of north-eastern and east-central Tanzania, with a possible sighting in the Nguru Mountains as well. The banded sunbird (Anthreptes rubritorques) is confined to montane forest patches in eastern Tanzania. The Eastern Arc Swynnerton’s robin (Swynnertonia swynnertoni rodgersi) is confined to the Udzungwa Mountains and, marginally, in lowland forest patches within the East Usambara Mountains. Nicoll’s weaver (Ploceus nicolli) is divided into two subspecies. The southern Nicoll’s weaver (P. n. anderseni) is known from the Udzungwa and Uluguru mountains of east-central and south-central Tanzania and possibly also the Rubeho Mountains. The Rubeho akalat (Sheppardia aurantiithorax) is a type of flycatcher confined to the mountains of central Tanzania. Winifred’s warbler (Scepomycter winifredae) is confined to montane forest areas within the Uluguru, Ukaguru, Udzungwa, and Rubeho mountains of east-central Tanzania. The Ukinga girdled lizard (Cordylus ukingensis) is known only from two montane grassland localities in southcentral Tanzania. The montane rock agama (Agama montana) is known only from remnant forest patches within the Usumbara and Uluguru mountains of eastern Tanzania. Usambara flap-nosed chameleon (Kinyongia tenuis) is known from the East Usumbara Mountains and Magrotto Hill in north-eastern Tanzania. A population in the Shimba Hills of south-eastern Kenya may represent an as-yet undescribed species. The Usambara dwarf gecko (Lygodactylus gravis) is confined to the Usambara and South Pare mountains of northeastern Tanzania. Two species of bush viper (Atheris) endemic to the Eastern Arc Mountains are threatened by loss of habitat. Barbour’s bush viper (A. barbouri) is confined to the Udzungwa and Ukinga mountains of south-central Tanzania. The eyelash bush viper (A. ceratophora) is found pachily in the mountains of eastern Tanzania. The Eastern Arc forest snake (Buhoma procterae) is confined to forest fragments in the Udzungwa and Uluguru mountains. The black garter snake (Elapsoidea nigra) confined to montane forest patches in north-eastern Tanzania and possibly south-eastern Kenya. The Eastern Arc vine snake (Thelotornis usambaricus) is confined to isolated mountain ranges in Tanzania and on the Kenyan coast.

The African Region

The Rondo worm snake (Afrotyphlops rondoensis) is confined to the Rondo and Makonde plateaus of extreme southeastern Tanzania. The Usambara spotted worm snake (A. gierrai) is confined to the Usambara and Nguru mountains of north-eastern Tanzania. Parker’s forest tree frog (Leptopelis parkeri) is found patchily in the mountains of eastern and central Tanzania. Burgess’ reed frog (Hyperolius burgessi) is confined to a few isolated massifs in the East Usambara, Uluguru, and South Nguru mountains of eastern Tanzania. Yakusin’s rocky river frog (Arthroleptides yakusini) is confined to the mountains of central-eastern Tanzania. The Eastern Arc leaf-folding frog (Afrixalus uluguruensis) is found patchily throughout the Eastern Arc Mountains. Two species of blue-bellied frog (Hoplophryne) endemic to the Eastern Arc Mountains are threatened by loss of habitat. The Eastern Arc blue-bellied frog (H. uluguruensis) is confined to montane forest fragments in the Uluguru, Nguru, Udzungwa, and Mahenge mountains of eastern Tanzania. Rogers’ blue-bellied frog (H. rogersi) is known only from the Usambara and Nguu mountains of north-eastern Tanzania. Nieden’s big-fingered frog (Probreviceps macrodactylus) is confined to the East Usambara and Nguru mountains of north-eastern Tanzania. Loveridge’s big-fingered frog (P. loveridgei) and the hard-nosed big-fingered frog (P. durirostris) are both found patchily in the mountains of eastcentral Tanzania. The Taita Hills The Taita hills are located in south-eastern Kenya (former Coast province). They consist of three massifs (Dabida, Sagalla, and Kasigau) historically covered by moist forest, of which only patches remain today. The Taita pygmy white-toothed shrew (Suncus aequatorius) is known only from Mount Sagalla and the Chawia Forest in the Taita Hills. The Taita thrush (Turdus helleri) is confined to forest patches in the Taita Hills and nearby Mount Mbololo. The Taita white-eye (Zosterops silvanus) is confined to the Taita Hills and nearby Mount Kasigau. The Taita apalis (Apalis fuscigularis) is a warbler-like bird confined to forest fragments in the Dabida and Mbololo massifs. The Taita purple-glossed snake (Amblyodipsas teitana) is known only from a single specimen collected in the Taita Hills in 1936. The Taita warty frog (Callulina dawida) is confined to forest fragments in the Dabida and Mbololo massifs. Loveridge’s caecilian (Boulengerula taitana) is confined to three of the four Taita Hills (Dawida, Mbololo, and Kasigau). The Sagalla Hill caecilian (B. niedeni) is confined to Sagalla Hill. The Pare Mountains Located in north-eastern Tanzania (Tanga and Kilimanjaro regions), the Pare Mountains are comprised of two subranges (the North and South Pare mountains, respectively).

The Pare pygmy chameleon (Rhampholeon viridis) is confined to the North and South Pare Mountains. The South Pare Mountains are located in Kilimanjaro and Tanga regions. The South Pare white-eye (Zosterops winifredae) is a type of bird confined to the South Pare Mountains. The South Pare screeching frog (Arthroleptis anotis) is confined to the Chome Forest Reserve in the South Pare Mountains. The Shengena warty frog (Callulina shengena) and Stanley’s warty frog (C. stanleyi) are both confined to the Chome Forest Reserve in the South Pare Mountains. The North Pare Mountains are located in Kilimanjaro region. Lapham’s warty frog (Callulina laphami) is confined to the Kindoroko and Minja forest reserves in the North Pare Mountains. The Usambara Mountains Located in north-eastern Tanzania (Tanga region), the Usambaras comprise the easternmost ranges of the Eastern Arc Mountains. Approximately 90 km long and about half that wide, they were historically clad in tropical forest whose long isolation has resulted in a great deal of endemism. They have been heavily deforested but patches of habitat remain. The Usambara shrew (Crocidura tansaniana) is confined to the Usambara Mountains. The northern Nicoll’s weaver (Ploceus nicolli nicolli) is confined to the East and West Usambaras. The Usumbara spiny pygmy chameleon (Rhampholeon spinosus) is confined to the Usambara Mountains. The Usambara leaf-folding frog (Afrixalus dorsimaculatus) is confined to the Usambara Mountains. Krefft’s puddle frog (Phrynobatrachus krefftii) is confined to the Usambara Mountains. The West Usambara Mountains comprise the main body of the Usambara Mountains. The Usambara akalat (Sheppardia montana) is a type of flycatcher confined to the West Usambaras. The West Usambara blade-horned chameleon (Kinyongia multituberculata) is confined to remnant forest patches within the West Usambaras. Vestergaard’s tree toad (Nectophrynoides vestergaardi) is known only from the Shume-Magamba and Mazumbai forest reserves and the Ambangulu Tea Estate in the West Usambaras. Tanner’s reed frog (Hyperolius tanneri) is known only from a locality near the Mazumbai Forest Reserve and from the Shume-Magambo Forest Reserve in the West Usambaras. The hidden screeching frog (Arthroleptis fichika) and Tanner’s screeching frog (A. tanneri) are both confined to the West Usambaras. The Mazumbai warty frog (Callulina kisiwamsitu) is confined to remnant forest patches within the West Usambaras.

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The East Usambara Mountains, being closer to the coast, receive more rainfall than the West Usambaras, and are also significantly smaller. The Usambara hyliota (Hyliota usambara) is a type of passerine bird confined to the East Usambaras, where it is found mainly in the foothills. The long-billed forest warbler (Artisornis moreaui) is confined to the East Usambaras. The Amani dapple-throat (Arcanator orostruthus amani) is a type of passerine bird confined to the East Usambaras, where in 1990 the population was estimated somewhere in the low thousands. It thereafter underwent a rapid decline and by 2006 had been reduced to only a few pairs. Vosseler’s blade-horned chameleon (Kinyongia vosseleri) and the giant East Usambara blade-horned chameleon (K. matschiei) are both confined to the East Usambaras. The East Usambara pygmy chameleon (Rhampholeon temporalis) is confined to the East Usambaras and Magrotto Hill. The Amani tree toad (Nectophrynoides frontierei) is known only from the Amani Nature Reserve in the East Usambaras. The East Usambara toad (Mertensophryne usambarae) is confined to the foothills of the East Usambaras. The Amani forest frog (Parhoplophryne usambarica) is known only from the heavily deforested hills west of Amani in the East Usambaras, where it was last seen in the 1920s. The Amani screeching frog (Arthroleptis xenodactylus) is confined to the East Usambaras. The East Usambara rocky river frog (Arthroleptides martiensseni) is confined to the East Usambaras. The Uluguru Mountains The Uluguru Mountains are located in east-central Tanzania (northern Morogoro region). The main portion consists of a ridge running roughly north–south and rising to 2630 m at its highest point. The Geata mouse shrew (Myosorex geata) is known only from its original collection during the early twentieth century within the Uluguru Mountains. The Uluguru bush-shrike (Malaconotus alius) is confined to a small area encompassing part of the Uluguru Nature Reserve. Loveridge’s sunbird (Cinnyris loveridgei) is confined to the Uluguru Nature Reserve, on the eastern slopes of the northern part of the Uluguru Mountains. The turquoise dwarf gecko (Lygodactylus williamsi) is confined to the Kimbozo and Ruvu forest reserves in the eastern foothills of the Uluguru Mountains, where it is threatened by habitat destruction and illegal collection for the international pet trade. The Uluguru limbless skink (Melanoseps emmrichi) is known only from a few specimens collected in the Uluguru Mountains. The ornate shovel-snout (Prosymna ornatissima) is a type of snake known only from a small area of forest in the Uluguru Mountains.

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The Uluguru gracile blind snake (Letheobia uluguruensis) is known only from the northern Uluguru Mountains. Four species of tree toad (Nectophrynoides) endemic to the Uluguru Mountains are threatened by loss of habitat. The minute tree toad (N. minutus) is confined to a small area of montane rainforest. The pseudo tree toad (N. pseudotornieri) is known only from the Uluguru Nature Reserve. The secret tree toad (N. cryptus) is known only from its original collections during the 1920s in the northern Uluguru Mountains. The smooth forest toad (N. laevis) is known only from a single specimen collected from the Uluguru South Forest Reserve. The Uluguru big-fingered frog (Probreviceps uluguruensis) is confined to the Uluguru Mountains. The Nyingwa caecilian (Scolecomorphus uluguruensis) is confined to the Uluguru Mountains. The Uluguru shellear (Kneria uluguru) is a small freshwater fish confined to streams and rivers of the upper Ruvu drainage descending from the Uluguru Mountains. The Nguru Mountains The Nguru Mountains are located in east-central Tanzania (northern Morogoro region). The Nguru spiny pygmy chameleon (Rhampholeon acuminatus) is confined to the Nguru Mountains. Three species of warty frog (Callulina) endemic to the Nguru Mountains are seriously threatened by loss of habitat. The Kanga warty frog (C. kanga) is known only from two localities within the Kanga Forest Reserve. The Nguru South warty frog (C. meteora) is known only from the Nguru South Forest Reserve. Hansen’s warty frog (C. hanseni) is known only from the western side of the Nguru Mountains. The Nguru screeching frog (Arthroleptis nguruensis) and the tiny screeching frog (A. kidogo) are both known only from the Nguru South Forest Reserve. The Ukaguru Mountains The Ukaguru Mountains are located in east-central Tanzania (north-western Morogoro region). The wide-headed tree toad (Nectophrynoides laticeps) and Paula’s tree toad (N. paulae) are both known only from the Mamiwa-Kisara Forest Reserve, where they are seriously threatened by loss of habitat. Maridad’s tree toad (Churamiti maridadi) is known only from four specimens collected from a single forest fragment in the Ukaguru Mountains. The Rubeho Mountains The Rubeho Mountains are located north of the Udzungwa Mountains in central Tanzania (Dodoma and Morogoro regions). The Rubeho forest partridge (Xenoperdix obscuratus) is confined to the Rubeho Mountains. The Rubeho spiny-throated reed frog (Hyperolius ukwiva) is known only from a single high-elevation locality within the Rubeho Mountains.

The African Region

Nike’s screeching frog (Arthroleptis nikeae) appears to be entirely confined to the Mafwemiro Catchment Forest Reserve, where it is seriously threatened by loss of habitat. The Mahenge Massif The Mahenge Massif is located in central Tanzania (Morogoro region). The Mahenge pygmy chameleon (Rhampholeon beraduccii) is confined to the Mahenge Massif. The Lupiro reed frog (Hyperolius lupiroensis) is known only from a single locality on the northern tip of the Mahenge Massif. The Udzungwa Mountains The Udzungwa Mountains are a biologically important range in south-central Tanzania (Iringa region). The Sanje River mangabey (Cercocebus sanjei) is confined to the vicinity of the Mwanihana Forest and Udzungwa Scarp Forest Reserve, on the eastern slopes of the Udzungwa Mountains. The Udzungwa red colobus (Piliocolobus gordonorum) is confined to forest fragments within the Udzungwa Mountains. While nominally safe within a few protected areas, illegal hunting continues. The grey-faced elephant shrew (Rhynchocyon udzungwensis) is found only in the Ndundulu-Luhomero and Mwanihana forests, Udzungwa Mountains. Phillips’ Congo shrew (Congosorex phillipsorum) is confined to the Udzungwa Mountains. The Udzungwa forest partridge (Xenoperdix udzungwensis) is confined to the Udzungwa Mountains. The rufous-winged sunbird (Cinnyris rufipennis) is confined to the Udzungwa Mountains. The Sanje dapple-throat (Arcanator orostruthus sanjei) is a type of passerine bird confined to a few forested areas on the Udzungwa Escarpment. The spiny-sided chameleon (Trioceros laterispinis) is confined to the Udzungwa Mountains. The Magombera single-horned chameleon (Kinyongia magomberae) is confined to the Mwanihana and Magombera Forests, Udzungwa Mountains. The Udzungwa long-tailed seps (Tetradactylus udzungwensis) is a type of lizard known only from the Gendawaki Valley in the Udzungwa Mountains. The Udzungwa skink (Leptosiaphos rhomboidalis) is known only from a single specimen collected in 1989 in the Mwanihana Forest, Udzungwa Mountains National Park. Three species of tree toad (Nectophrynoides) are endemic to the Udzungwa Mountains, where they are seriously threatened by loss of habitat. Poynton’s tree toad (N. poyntoni) is known only from a single specimen collected from the Udzungwa Scarp Forest Reserve in 2003. It has not been recorded since despite targeted searches, and may be extinct. The Kihansi tree toad (N. asperginis) was historically known only from a small area around Kihansi Falls, in the Kihansi Gorge, Udzungwa

Mountains. Extinct in the wild after 2009 owing to the construction of a dam, it fortunately survived in captivity and attempts have since been made to try and reintroduce it with the help of artificial sprinkler systems. The Udzungwa Scarp tree toad (N. wendyae) is known only from a single valley within the Udzungwa Scarp Forest Reserve. The Udzungwa leaf-folding frog (Afrixalus morerei) is confined to the Udzungwa Mountains. Keith’s striped frog (Phlyctimantis keithae) is confined to the area of the Kigogo Forest Reserve, in the southern Udzungwa Mountains. The Udzungwa reed frog (Hyperolius kihangensis) is confined to the Udzungwa Mountains. The Mafinga puddle frog (Phrynobatrachus breviceps) is known only from a single locality on the Udzungwa Plateau.

The Southern Highlands The Southern Highlands are a group of plateaus and volcanic mountains located in south-western Tanzania, on the northern end of Lake Malawi. Matilda’s bush viper (Atheris matildae) is known only from a single montane forest fragment within the Southern Highlands, the precise location of which is being withheld by conservationists for its security. The Livingstone Mountains The Livingstone Mountains (known locally as the Kipengere Range) are a plateau-like ridge located in south-western Tanzania (Njombe and Mbeya regions). The northern end of the range is also known as the Poroto Mountains, while more southerly areas are referred to as the Kinga Mountains. The overlooked screeching frog (Arthroleptis kutogundua) is known only from its original collection in 1930 from the Ngozi Crater. The species is most likely extinct due to loss of habitat. Davenport’s reed frog (Hyperolius davenporti) is known from a single locality in the Sakara Nyumo Forest Reserve of the Livingstone Mountains, where it is threatened by ongoing habitat destruction.

The Mulanje Massif The Mulanje Massif is located in south-eastern Malawi (Southern region). It rises sharply from the surrounding plains and features many individual peaks reaching over 2500 m. Much of the massif consists of rolling grassland intersected by deep forested ravines. It is protected by the Mulanje Mountain Forest Reserve. The Mulanje chameleon (Nadzikambia mlanjensis) is known only from the Mulanje Massif. The Mulanje pygmy chameleon (Rhampholeon platyceps) is found only in mid- to high-elevation forest fragments of Mount Mulanje and adjacent Mount Mchese. Broadley’s mongrel frog (Nothophryne broadleyi) is confined to the Mulanje Massif. The Mulanje river frog (Amietia johnstoni) is confined to the high plateau of the Mulanje Massif.

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The Eastern Highlands The Eastern Highlands (also known as the Manica Highlands) are a mountainous area located in eastern Zimbabwe (Manicaland province) and west-central Mozambique (Manica province). They are comprised of three main mountain groups: the Nyanga Mountains, the Bvumba Mountains, and the Chimanimani Mountains, all of which are sparsely populated and covered by grasslands and forest patches. They include the important Chirinda Forest of eastern Zimbabwe, the southernmost tropical rainforest in Africa. The Chirinda veld rat (Aethomys silindensis) is known only from three localities within the Eastern Highlands. The Eastern Highlands Swynnerton’s robin (Swynnertonia swynnertoni swynnertoni) is confined to the Chirinda Forest and a few other tiny forest patches on the Zimbabwe/ Mozambique border. Marshall’s pygmy chameleon (Rhampholeon marshalli) is confined to forest fragments within the Eastern Highlands. The Rhodesian big-fingered frog (Probreviceps rhodesianus) is known only from three isolated localities in eastern Zimbabwe. The Eastern Highlands stream frog (Strongylopus rhodesianus) is confined to a few scattered localities. The Nyanga Mountains The Nyanga Mountains are located in eastern Zimbabwe and south-western Mozambique. Arend’s golden mole (Carpitalpa arendsi) is confined to the Nyanga Mountains. The Nyanga river frog (Amietia inyangae) is known only from two localities in the Nyanga Mountains. Mount Nyangani (formerly Mount Inyangani) is the highest mountain in Zimbabwe. It is located within Nyanga National Park. The Nyangani toad (Vandijkophrynus inyangae) is known only from the plateau on Mount Nyangani. The Chimanimani Mountains The Chimanimani Mountains are located in eastern Zimbabwe and south-western Mozambique. The cave screeching frog (Arthroleptis troglodytes) is confined to the Chimanimani Mountains. Last recorded in 1962 and thought to be extinct, it was rediscovered in small numbers in 2016.

The Central Plateau The Central Plateau is a highland area dominating much of southern Africa in South Africa, Lesotho, Swaziland, Zambia, Angola, and Namibia. It is characterized mainly by highveld (high-elevation grassland). The southern bald ibis (Geronticus calvus) is confined to the Central Plateau in north-eastern South Africa, Lesotho, and western Swaziland. It is threatened by habitat destruction and human interference with its breeding colonies.

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The giant spiny-tailed lizard (Smaug giganteus) is confined to highveld grasslands of north-eastern South Africa (northern Free State and south-western Mpumalanga). The Snow Mountains adder (Bitis inornata) is known only from a small area of montane grassland in south-central South Africa (Eastern Cape). Breyer’s long-tailed seps (Tetradactylus breyeri) is a snakelike lizard that was historically widespread in the montane and highveld grasslands of north-eastern South Africa, but is now confined to a few disjunct populations in Mpumalanga, Free State, and KwaZulu-Natal. The Great Escarpment The Great Escarpment consists of the steep slopes from the high Central Plateau downwards in the direction of the oceans that surround southern Africa on three sides. While it lies predominantly within the borders of South Africa, in the east it extends northwards to eastern Zambia and in the west into southern Namibia. Gunning’s golden mole (Neamblysomus gunningi) and Juliana’s golden mole (N. julianae) are both known only from a few localities in north-eastern South Africa, where they are threatened by loss of habitat. The long-toed forest tree frog (Leptopelis xenodactylus) is confined to a few upland bogs, grassy wetlands, and marshes in south-eastern South Africa (southern KwaZulu-Natal). Two species of chirping frog (Anhydrophryne) endemic to the Great Escarpment are threatened by loss of habitat. The Hogsback chirping frog (A. rattrayi) is known only from the Amathole, Katberg, and Keiskammahoek mountains of southeastern South Africa (Eastern Cape). The mistbelt chirping frog (A. ngongoniensis) is known only from a few localities in eastern South Africa (KwaZulu-Natal). The Drakensberg (Drakensberge in Afrikaans; uKhahlamba in Zulu) is the name given to the eastern portion of the Great Escarpment in eastern South Africa (Mpumalanga, KwaZuluNatal, Limpopo, and Free State) and Lesotho. The robust golden mole (Amblysomus robustus) is known only from a few localities within the Drakensberg. Rudd’s lark (Heteromirafra ruddi) is confined to two main areas of high-altitude grassland, where it is declining due to loss of habitat. Botha’s lark (Spizocorys fringillaris) is confined to a few areas of southern Mpumalanga and eastern Free State, where the total population in 2019 was estimated at 1500–5000. The yellow-breasted pipit (Hemimacronyx chloris) is a rare bird species confined to eastern South Africa and Lesotho. Methuen’s dwarf gecko (Lygodactylus methueni) is known only from a few areas of montane grassland in north-eastern South Africa (southern Limpopo). Eastwood’s longtailed seps (Tetradactylus eastwoodae) was a type of lizard known only from two specimens collected in the early twentieth century. Its montane grassland habitat has long since been eradicated, and it is now thought to be extinct.

The African Region

The woodbush legless skink (Acontias rieppeli) is confined to a small area of north-eastern South Africa (Limpopo). The Amatola or Amathole Mountains are a range of densely forested mountains located in south-eastern coastal South Africa (Eastern Cape). They form a part of the southern portion of the Great Escarpment. The Amatola toad (Vandijkophrynus amatolicus) is known only from the Winterberg and Amatola Mountains, although it has not been reported from the former since the 1970s.

The Cape Fold Mountains The Cape Fold Mountains form a series of parallel ranges that run along the south-western and southern coastlines of South Africa for some 850 km. The long-tailed forest shrew (Myosorex longicaudatus) is divided into two subspecies. The nominate form (M. l. longicaudatus) is confined to a small area of the Cape Fold Mountains in southern South Africa (Western Cape and Eastern Cape), where it is threatened by loss of habitat. Smith’s dwarf chameleon (Bradypodion taeniabronchum) is confined to a few disjunct localities within the Elandsberg and Kareedouw mountains of southern South Africa (Eastern Cape), as well as an additional wetland area near Cape St. Francis. The species may have been historically widespread in the lowlands, from where it has been extirpated. Rose’s mountain toadlet (Capensibufo rosei) is confined to a few, mainly higher-elevation fynbos heathland localities in south-western South Africa (Western Cape). The northern moss frog (Arthroleptella subvoce) is known only from three localities within the Groot Winterhoek Wilderness Area in south-western South Africa (Western Cape). The rough moss frog (A. rugosa) is confined to a single inselberg in south-western South Africa (Western Cape). Hewitt’s ghost frog (Heleophryne hewitti) is known only from a few localities within the Elandsberg and Cockscomb mountains of south-eastern South Africa (Eastern Cape). The Hottentots Holland Mountains The Hottentots Holland Mountains are located in southwestern South Africa (Western Cape). The dwarf crag lizard (Hemicordylus nebulosus) is confined to higher elevations of the Hottentots Holland Mountains. The Langeberg Mountains The Langeberg Mountains are located in south-western South Africa (Western Cape). The Langeberg long-tailed forest shrew (Myosorex longicaudatus boosmai) is confined to the Langeberg Mountains. Table Mountain Table Mountain is located near Cape Town, south-western South Africa (Western Cape). The Table Mountain ghost frog (Heleophryne rosei) is confined to the southern, eastern, and marginally western slopes of Table Mountain.

The Angolan Highlands The Angolan Highlands are located just inwards from the coast in eastern Angola. They encompass some of the highest mountains in the country along with a few outlying inselbergs, remnants of the Great Escarpment from a time when it was more extensive. The heather shrew (Crocidura erica) is known only from western Angola. Swierstra’s francolin (Pternistis swierstrai) is a partridgelike bird confined to a few montane forest patches. Two species of bush shrike (Laniarius) endemic to the Angolan Highlands are threatened by loss of habitat. Braun’s bush shrike (L. brauni) is known only from a very small area of north-western Angola (Cuanza Norte province). The Gabela bush shrike (L. amboimensis) is a rare species from central-western Angola. Pulitzer’s longbill (Macrosphenus pulitzeri) is a warblerlike bird confined to the Angolan Highlands. The Gabela akalat (Sheppardia gabela) is a type of flycatcher known only from a few forest patches on the escarpement of central-western Angola. The Congulu forest tree frog (Leptopelis jordani) is known only from a single specimen collected in 1934 from centralwestern Angola (Cuanza Sul province). The Quissange forest tree frog (Leptopelis marginatus) is known only from a single (now lost) specimen collected in 1895 from Benguela province. The taxonomic validity of both species, however, has been questioned. Parker’s white-lipped frog (Amnirana parkeriana) is known only from a small area of the northern Angolan Highlands.

Miscellaneous Mountains and Highlands The Nyika Plateau is located mainly in northern Malawi, with a small portion in north-eastern Zambia. The Nyika burrowing shrew (Myosorex gnoskei) is endemic to the Nyika Plateau within Nyika National Park, Malawi. The Njesi Plateau is located in north-western Mozambique (Niassa province). The Njesi forest warbler (Artisornis sousae) is confined to the Njesi Plateau, where the total population is very small. The Namuli Massif is located in northern Mozambique (Zambézia province). It consists of a level plateau with the granite dome of Mount Namuli rising above. Historically clad in tropical rainforest and still an important biodiversity hotspot, the lower slopes have now been mostly converted to agriculture. Vincent’s bush squirrel (Paraxerus vincenti) is known only from Mount Namuli. Mount Chiperone is located in northern Mozambique (Zambézia province). The Chiperone pygmy chameleon (Rhampholeon nebulauctor) is confined to montane forest on Mount Chiperone. Mount Mabu is located in northern Mozambique (Zambézia province). It is well-known for the Mabu Forest, which is perhaps the largest remaining mid-altitude rainforest

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in Africa. The Mabu pygmy chameleon (Nadzikambia baylissi) is confined to Mount Mabu. Mount Inago is located in northern Mozambique (Nampula province). The Inago pygmy chameleon (Rhampholeon bruessoworum) is known only from Mount Inago. Mount Ribáuè is located in northern Mozambique (Nampula province). It consists of two narrow inselbergs (the other being Mount M’pàluwé) separated by a narrow valley. The Ribáuè mongrel frog (Nothophryne ribauensis) is confined to Mount Ribáuè. Mount Gorongosa is an inselberg located in central Mozambique (Sofala province). While nominally protected within Gorongosa National Park most of its montane forest has been destroyed, with only fragments now remaining. The Gorongosa pygmy chameleon (Rhampholeon gorongosae) is confined to Mount Gorongosa. The Natundu Hills are located in southernmost Malawi. Chapman’s pygmy chameleon (Rhampholeon chapmanorum) is confined to forest fragments in the Natundu Hills. The Lebombo Mountains (sometimes called the Lubombo Mountains) are a narrow, 800-km-long range in South Africa, Mozambique, and Swaziland. Marley’s golden mole (Amblysomus marleyi) is known only from two localities (Ubombo and Ingwavuma) on the eastern slopes of the Lebombo Mountains in KwaZulu-Natal.

Gallery Forests Gallery forests are tropical forests that grow along rivers or wetlands, sometimes extending far from the banks and projecting into landscapes that are otherwise only sparsely treed such as savannas, grasslands or deserts. Ansell’s shrew (Crocidura ansellorum) is known only from a few specimens collected from north-western Zambia and southern Democratic Republic of Congo. The Vilhena’s reed frog (Hyperolius vilhenai) is known only its original collection in the 1960s from gallery forests along the Luita River in north-eastern Angola. Five additional specimens have since been obtained from Cangandala National Park some 350 km to the south-west, which have been provisionally assigned to this species.

The Tana River Forest The Tana River in south-eastern Kenya still has remnant patches of gallery forest along its lower course. Long isolated by heavy deforestation in the surrounding areas, they have effectively ‘trapped’ a number of unique species (in particular primates) and now provide a last refuge for them. In 1976, in deference to their extreme importance, these forests were set aside as the Tana River Primate National Reserve. Unfortunately, a legal challenge resulted in the reserve being degazetted in 2007. The Tana River mangabey (Cercocebus galeritus) is endemic to gallery forests along the lower Tana River.

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Cultivation in the region has long since eliminated most of the forests, forcing the species to withdraw to forest patches, where they continue to be subject to intensive trapping. In 1994 the total population was estimated at between 1000 and 1200. The Tana River red colobus (Piliocolobus rufomitratus) has a similar history and range, with a total population in 2003 of 1100–1300 (down from 1200 to 1800 in 1975). The Tana River cisticola (Cisticola restrictus) is a passerine bird known only from a small number of specimens, all collected within the lower Tana River drainage. It was last seen in 1972. The Tana River white-winged apalis (Apalis chariessa chariessa) was a type of passerine bird known only from the lower Tana River, where it was last observed in 1961.

Coastal Moist Forests A strip of moist tropical as well as subtropical lowland forests extends along Africa’s Indian Ocean coast from southern Somalia to southern South Africa. Taken together they are among the most biologically important and threatened in the African Region.

East African Coastal Forests The East African coastal forests are a tropical moist forest region located in a narrow band along the Indian Ocean coast from southern Somalia in the north, through coastal Kenya and Tanzania to just north of the mouth of the Limpopo River in southern Mozambique. It also includes the Zanzibar Archipelago, here treated in a separate section. The region has been heavily impacted by deforestation. The Arabuko Sokoke Forest, located on the central coast of Kenya, is the only remaining, larger coastal forest in East Africa. It is nominally protected within the Arabuko-Sokoke National Forest Reserve, a small portion of which is further set aside as a national park. Ader’s duiker (Cephalophus adersi) was long thought to be endemic to the island of Zanzibar, until found to occur as well in the Arabuko-Sokoke Forest on the coast of Kenya. In recent years populations have been discovered in other coastal forest pockets in Kenya and Tanzania and the total population is believed to be fairly large. The species remains threatened, however, by loss of habitat, predation by feral dogs and hunting. The Sokoke dog mongoose (Bdeogale omnivora) appears to be mostly or entirely confined to the coastal forests of southeastern Kenya and north-eastern Tanzania. The golden-rumped elephant shrew (Rhynchocyon chrysopygus) occurs in the Arabuko Sokoke Forest and a few other small coastal forests fragments in south-eastern Kenya. The Tanzanian woolly bat (Kerivoula africana) is known only from a few localities in coastal Tanzania, where it was long thought to be extinct until its rediscovery in 2000. It is threatened by loss of habitat and capture for use in ‘traditional medicine’.

The African Region

The Dar-es-salaam pipistrelle bat (Pipistrellus permixtus) is known only from a single specimen collected from coastal forest in Tanzania in the 1950s. The Kenyan wattled bat (Glauconycteris kenyacola) is known only from a single locality in coastal south-eastern Kenya (former Coast province). Hildegarde’s tomb bat (Taphozous hildegardeae) is a cavedependant species known only from a few localities along the coast of south-eastern Kenya and north-eastern Tanzania, including the islands of Zanzibar and Pemba. The Sokoke scops owl (Otus ireneae) is known only from Arabuko-Sokoke Forest of south-eastern Kenya and from the lowlands of the East Usambara Mountains in northeastern Tanzania. Fischer’s spotted ground thrush (Geokichla guttata fischeri) is confined to the coastal forests of Kenya and Tanzania, where it is rare and decreasing. Clarke’s weaver (Ploceus golandi) is a type of passerine bird known only from a few localities in south-eastern Kenya. Günther’s house gecko (Hemidactylus modestus) is known only from a few coastal forest fragments in south-eastern Kenya, where it has not been recorded in several decades. The Dar-es-Salaam dwarf gecko (Lygodactylus inexpectatus) is known only from the Dar-es-Salaam area of Tanzania, where it has not been recorded since its original description in the mid-twentieth century. The sandy limbless skink (Scolecoseps acontias) is known for certain only from a single, now lost specimen collected in 1913 from central coastal Tanzania. The Kilwa sharp-snouted worm lizard (Ancylocranium ionidesi) is known only from a few localities in south-eastern Tanzania, including the Litipo Forest Reserve. The Tanga blind snake (Afrotyphlops platyrhynchus) is known only from its original collection from a single locality in extreme north-eastern Tanzania in the early twentieth century. The Mbanja worm snake (Leptotyphlops mbanjensis) is known only from five specimens collected from extreme south-eastern Tanzania. Schiotz’s leaf-folding frog (Afrixalus sylvaticus) is found patchily in coastal south-eastern Kenya and northeastern Tanzania. The Ruvu reed frog (Hyperolius ruvuensis) is known only from the Ruvu South Forest Reserve in eastern Tanzania. The Shimba Hills reed frog (H. rubrovermiculatus) is confined to the Shimba Hills of coastal south-eastern Kenya. Lonnberg’s screeching frog (Arthroleptis lonnbergi) is known for certain only from lowland forests in eastern and north-eastern Tanzania. The Zanzibar puddle frog (Phrynobatrachus ungujae) is known only from the Jozani Forest on Zanzibar and from the Arabuko-Sokoke Forest in south-eastern Kenya. The Litopo Forest The Litipo Forest is located in coastal Tanzania.

The Litipo sand skink (Scolecoseps litipoensis) is known only from a single specimen collected in the Litipo Forest in 1993.

South-East African Coastal Forests The South-East African coastal forests are a tropical moist forest region running along a narrow band from central coastal Mozambique to south-eastern South Africa. The Knysna warbler (Bradypterus sylvaticus) is confined to remnant coastal forest patches of the Eastern and Western Cape of South Africa. The total population is thought to be around 2500. The South African spotted ground thrush (Geokichla guttata guttata) is confined to Eastern Cape province (possibly extending into a small area of KwaZulu-Natal), where in 2005 the total population was estimated at less than 300 pairs. The Transkei dwarf chameleon (Bradypodion caffer) is confined to a small area of coastal forest in Eastern Cape province. Günther’s dwarf burrowing skink (Scelotes guentheri) is known only from a single specimen collected in 1886 from KwaZulu-Natal. It is considered to be extinct, a victim of habitat destruction. The legless dwarf burrowing skink (S. inornatus) is known only from a few localities along a small stretch of coast near Durban in KwaZulu-Natal, where it is seriously threatened. The Knysna leaf-folding frog (Afrixalus knysnae) is confined to a few areas of Eastern Cape and Western Cape provinces. The Kloof frog (Natalobatrachus bonebergi) is confined to a few lowland forest patches in Eastern Cape and KwaZuluNatal provinces.

Lowland Mediterranean Forest (Fynbos) Fynbos, which means ‘fine-leaved plants’ in Afrikaans, is a small belt of maquis-like shrubland and heathland located in the Western Cape and Eastern Cape provinces of South Africa (roughly from north of Cape Town to Port Elizabeth). Two types are recognized: lowland fynbos (i.e. areas below 300 m) and montane fynbos, which is discussed separately in the mountains section. Known for its exceptional degree of biodiversity and endemism, lowland fynbos in particular has long been under threat by introduced plant species and coastal development. The geometric tortoise (Psammobates geometricus), one of Africa’s rarest reptiles, was thought to have become extinct about 1910, but in 1946–47 and 1958 specimens were obtained north and east of Cape Town. Today it is entirely confined to two small protected areas, although drought, grass fires, and illegal capture for the pet trade remain serious threats. It has never successfully reproduced in captivity. The western leopard toad (Sclerophrys pantherina) is confined to a few lowland areas of the Cape Peninsula, the Cape Flats, and the Agulhas Flats in south-western South Africa

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(Western Cape). It is threatened mainly by loss of habitat and road kill. The Cape Flats frog (Microbatrachella capensis) occurs only in the coastal lowlands of south-western Western Cape province.

Lowland Tropical and Subtropical Savannas Areas of lowland tropical and subtropical savannas, often interspersed with woodlands of varying type, occur throughout the African Region. During the pluvial periods of the Ice Age great forests spread over wide areas of Africa now covered by savannas, but evidently there were sufficiently large open areas during these forested periods to allow savanna species to survive in unparalleled variety. After the last ice sheet had melted from the Northern Hemisphere, a new dry period began in Africa, and about 20,000 years ago the savannas started increasing in area. Only a century or two ago, the mighty savannas of Africa provided scenes that we today find difficult to visualize. Numerous species of large and small antelopes, mixed with giraffes, buffaloes, rhinoceroses, and elephants, lived on the various kinds of savannas. These plains were regions of biological equilibrium, highly productive climax areas created by Nature herself through countless ages. In this highly specialized ecological system, each organism was a necessary element in its perpetuation. Different animal species made efficient use of the whole array of available niches within the habitat. The various species of hoofed animals grazed and browsed together, choosing different kinds of bulbs, grass, twigs, leaves, and fruits as food. Selection had eliminated competition, and in this way the effect of grazing was evenly distributed. With exceptions for species such as common eland (Tragelaphus oryx), Beisa oryx (Oryx beisa), impala (Aepyceros melampus), gerenuk (Litocranius walleri), Grant’s gazelle (Nanger granti), and Thomson’s gazelle (Eudorcas thomsonii), which need little or no water, herds congregated on the savanna regions around lakes, rivers, springs, and small water holes, for not many savanna species can live for long periods without water. During dry periods most of these hoofed animals moved to other regions that had recently had rain and where the grass was plentiful and green. And so it went, all year round. The hoofed animals followed the rain, changing pastures regularly, and as a consequence the grazing never destroyed the vegetation, which after the next rainy period was as fresh and green as ever. The giraffe (Giraffa camelopardalis) is the world’s tallest living land mammal, with fully grown adults standing between 4.3 and 5.7 m in height (although the tallest male ever recorded was just under 5.9 m). Despite many independent studies indicating multiple forms there is as yet no general agreement on the taxonomy, and so for the purposes of this book we shall consider it to be a single species with nine subspecies. These gentle creatures remain widespread in savanna woodlands across eastern and southern Africa, with smaller, isolated populations in western and central Africa, but are now

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increasingly threatened everywhere by hunting (both legal and illegal) for meat and trophies. The Nubian giraffe (G. c. camelopardalis) is found in eastern South Sudan and south-western Ethiopia, Kenya, and Uganda, where around 2150 are thought to remain in the wild. A distinct ecotype known as ‘Rothschild’s giraffe’ may be included with this form. The latter lives in parts of Kenya and Uganda, where the wild population is around 1500, although there are several hundred in captivity. The Kordofan giraffe (G. c. antiquorum) is found in southern Chad, the Central African Republic, northern Cameroon, and north-eastern Democratic Republic of Congo. Some 2000 remain in the wild. The reticulated giraffe (G. c. reticulata) occurs in north-eastern Kenya, south-eastern Ethiopia, and south-western Somalia. An estimated 8660 remain in the wild, with an additional 450 in captivity. The Namibian giraffe (G. c. angolensis) is found in northern Namibia, south-western Zambia, Botswana, and western Zimbabwe, where the total population is estimated at around 13,000. The Masai giraffe (G. c. tippelskirchi) ranges in central and southern Kenya and Tanzania, where a total of 32,550 are thought to remain. The Luangwa giraffe (G. c. thornicrofti) is confined to the Luangwa Valley in eastern Zambia, where there are no more than 550 remaining. Finally, the Niger giraffe (G. c. peralta) is found only in south-western Niger, where just 400 survive. The white or square-lipped rhinoceros (Ceratotherium simum), one of the largest and heaviest of terrestrial land mammals, is divided into two subspecies. The northern white rhinoceros (Ceratotherium simum cottoni) is, at time of writing, functionally extinct, with only two females remaining. At the beginning of the nineteenth century the form was widely but discontinuously distributed in parts of what is now north-western Uganda, southern Chad, south-western Sudan, eastern Central African Republic, and north-eastern Democratic Republic of the Congo. Long persecuted by trophy hunters, by the early twentieth century it had already all but disappeared everywhere except in Sudan, where it enjoyed strict protection, and in Garamba National Park, Democratic Republic of Congo, which had been specially created for it in 1938. The rhinos held out in both areas for the next few decades, but endless civil war combined with poaching for its horns ultimately wiped it out in both countries. The last wild individual was reported in Garamba in 2006. The last four survivors, all zoo captives, were ultimately brought together and returned to Africa in an expensive and ultimately misguided attempt at semi-reintroduction on a private reserve in Kenya, where the only male, ‘Sudan’, died in 2017. The southern white rhinoceros (C. s. simum) was and remains the most widespread, living as it does in the grasslands of central and southern Africa. Long a favourite target for trophy hunters, by the early twentieth century it was nearly wiped out, with less than 20 remaining and all living on a single reserve in South Africa. From this tiny remnant it has since made a remarkable comeback, reaching 1000 by the 1980s. As of 2015 the population stands at around 20,000,

The African Region Figure 3.8 ‘Angalifu’, a male northern white rhinoceros at the San Diego Wild Animal Park (deceased 2014). (Credit: San Diego Wild Animal Park.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

making it by far the most abundant rhino in the world, although the animals remain vulnerable to poaching and loss of habitat. There is a large captive population, and many reintroductions within the historical range have taken place or are planned. The black or hook-lipped rhinoceros (Diceros bicornis), previously discussed, was historically divided into several subspecies within the African Region. The now-extinct northeastern black rhinoceros (D. b. brucii) lived in central Sudan, Eritrea, northern and south-eastern Ethiopia, Djibouti, and northern and south-eastern Somalia. The last relict populations in northern Somalia vanished sometime during the early twentieth century. Of the surviving forms the south-central black rhinoceros (D. b. minor) is the most widely distributed, historically being found from north-eastern South Africa (KwaZulu-Natal) to north-eastern Tanzania and south-eastern Kenya. It survives in protected areas throughout most of its former range, but has probably been extirpated in eastern Angola, southern Democratic Republic of Congo, and possibly Mozambique. It was formerly extirpated in Malawi, Botswana, and Zambia as well, but has since been reintroduced. The eastern black rhinoceros (D. b. michaeli) was historically distributed from South Sudan, Uganda, and Ethiopia down through Kenya into north-central Tanzania. Today, its range is limited primarily to Kenya and Tanzania. The Ugandan black rhinoceros (D. b. ladoensis) was formerly found from South Sudan, across Uganda into western Kenya and southwesternmost Ethiopia. Black rhinos are considered to be extirpated across most of this area and their conservation status is unclear, although they probably survive in small numbers in

Kenyan reserves. The south-western black rhinoceros (D. b. occidentalis) is a notably small subspecies adapted to survival in desert and semi-desert conditions. Originally distributed in north-western Namibia and south-western Angola, today it is restricted to wildlife reserves in Namibia, with sporadic sightings in Angola. The Chobe black rhinoceros (D. b. chobiensis) is restricted to the Chobe Valley in south-eastern Angola, the Zambezi region of Namibia, and (at least formerly) northern Botswana. The secretary bird (Sagittarius serpentarius) is a large, mainly terrestrial bird of prey found over most of subSaharan Africa outside the rainforest belt, but has declined in many areas due to loss of habitat and hunting.

Lowland Tropical Savannas Areas of lowland tropical savanna, ranging from sparsely wooded grassland areas to open savanna dotted with acacias, are to be found across north-central, south-central, and eastern Africa. Hunter’s antelope or hirola (Beatragus hunteri), one of the world’s rarest antelopes, is confined to the border area of south-eastern Kenya and south-western Somalia. The wild population is estimated at 300–500 and declining, and there are none in captivity. The hartebeest (Alcelaphus buselaphus) is a large antelope that has been extensively hunted for both food and sport. Swayne’s hartebeest (A. b. swaynei) formerly occurred across the southern Rift Valley. It disappeared from Somaliland (north-western Somalia) in the early twentieth century due to rinderpest, and has since been effectively restricted to two sites

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in Ethiopia (Senkelle Swayne’s Hartebeest Sanctuary and Maze National Park). Reintroductions have been attempted in Awash and Nech Sar national parks, but without success. In 2008 the total population was estimated at less than 800. The tora hartebeest (A. b. tora) formerly occurred in western and southwestern Eritrea, north-western Ethiopia, and the adjacent border regions of Sudan (where it is now definitely extinct). Small numbers may survive in Eritrea or Ethiopia, but there have been no records since the 1990s and the subspecies is possibly extinct. The lelwel hartebeest (A. b. lelwel) is a large antelope that was historically found primarily in the Central African Republic, south-western Ethiopia, in northern and north-eastern Democratic Republic of the Congo, southern Sudan, and the north-western extreme of Tanzania. A massive population decline since the 1980s has resulted in most individuals now being confined to protected areas both inside and outside this range (southern Chad and northern Uganda). The tsessebe (Damaliscus lunatus) is a type of large, social antelope found in pockets across sub-Saharan Africa outside the rainforest belt. A few subspecies are considered threatened due to displacement by cattle and uncontrolled hunting for meat. The korrigum (D. l. korrigum) historically occurred from southern Mauritania and Senegal to western Chad, but underwent a dramatic decline during the twentieth century. It is now extirpated in Mauritania, Mali, Senegal, and The Gambia, and most likely from Nigeria and Chad as well except as vagrants. The coastal topi (D. l. topi) is known from southern Somalia within the riverine grasslands of the lower Shebelle and Jubba rivers and in parts of northern Kenya. Haggard’s oribi (Ourebia ourebi haggardi) is a type of small antelope confined to coastal savannas of southern Somalia and northern Kenya. It has long been threatened by loss of habitat and hunting. The Somali gerbil (Ammodillus imbellis) is known only from a few specimens collected from Somalia and southeastern Ethiopia. The Somali golden mole (Calcochloris tytonis) is known only from a single specimen (partial owl pellet remains) collected near Mogadishu, Somalia, in 1964. Johnston’s African dormouse (Graphiurus johnstoni) is confined to the savanna woodlands of southern Malawi and possibly adjacent areas of Zambia and Zimbabwe. The dusky elephant shrew (Elephantulus fuscus) is known only from a handful of specimens collected from southern Zambia, Malawi, and south-western Mozambique. All but one were obtained prior to 1968. The dusky-footed elephant shrew (E. fuscipes) is known only from a handful of specimens, all obtained prior to 1968, from southern South Sudan, northwestern Uganda and north-eastern Democratic Republic of Congo. Fischer’s shrew (Crocidura fischeri) is known only from two localities in southern Kenya and north-eastern Tanzania. Vinson’s slit-faced bat (Nycteris vinsoni) is known only from two specimens smoked out of a baobab tree in the mid1960s in southern Mozambique.

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The Zambian barbet (Lybius chaplini) is a type of bird confined to a small area of south-central Zambia, where it is threatened by loss of habitat. Friedmann’s lark (Mirafra pulpa) is a rare species known only from a small number of specimens collected in three disjunct areas of central and southern Kenya, south-western Ethiopia, and north-eastern Tanzania. Parker’s house gecko (Hemidactylus megalops) is known only from its original collection in Somalia in the early 1930s. Grandison’s dwarf gecko (Lygodactylus grandisonae) is known only from a few specimens collected from a single locality on the Kenya/Ethiopia border. The angulated dwarf gecko (L. grzimeki) is known only from a few specimens collected from Lake Manyara National Park in northcentral Tanzania. The imperial flat lizard (Platysaurus imperator) is confined to a small area of savanna in north-western Mozambique and north-eastern Zimbabwe. The Caprivi rough-scaled lizard (Ichnotropis grandiceps) is a rare species known only from the Caprivi Strip of northeastern Namibia and adjacent north-western Botswana. Chelazzi’s garter snake (Elapsoidea chelazzii) is known only from a small area of dry open woodland in southern Somalia. Parker’s worm snake (Leptotyphlops parkeri) is known only from two specimens collected many decades ago from two widely separated localities (the northern Ogaden region of Ethiopia and Tsavo National Park in Kenya). The Nairobi toad (Mertensophryne nairobiensis) is confined to a small area of savanna woodland in southcentral Kenya. The Bata marsh toad (Sclerophrys chudeaui) is known only from its original collection in the early twentieth century from an undefined locality in Mali. The Merera toad (S. reesi) is known only from a few specimens collected from south-central Tanzania. The Mossamedes toad (Poyntonophrynus grandisonae) is known only from a few specimens collected in south-western Angola (Namibe province). The Mapacha ridged frog (Ptychadena mapacha) is known only from the eastern Caprivi Strip in north-eastern Namibia, but likely also occurs in south-western Zambia, south-eastern Angola, and northern Botswana. The Masiliwa shovel-nosed frog (Hemisus brachydactylus) is known only from its original collection in the 1960s from an undefined locality in central Tanzania. De Witte’s shovelnosed frog (H. wittei) is known only from south-eastern Democratic Republic of Congo and north-eastern Zambia. The Mongu shovel-nosed frog (H. barotseensis) is known only from the Barotse Floodplain along the Zambezi River in central-western Zambia. Miombo Woodlands ‘Miombo’ is the vernacular word for Brachystegia, a genus of tree comprising a large number of species notable for shedding

The African Region

their leaves during the dry season. Miombo woodlands are characterized by the predominance of these species. They form a broad belt across south-central Africa, running from Angola and the Republic of Congo through the Democratic Republic of Congo, Burundi, Zambia, and Zimbabwe to Malawi, Mozambique, and Tanzania. The giant sable antelope (Hippotragus niger variani) is confined to the region between the Luando and Cuanza rivers (and to an area immediately north of the Luando) in central Angola. In the late 1960s there were an estimated 2000–3000, but there was a great deal of uncertainty regarding its survival of the Angolan Civil War (1975–2002). However, in 2004 photographic evidence was obtained of a surviving herd in Cangandala National Park. Others were found, and in 2007 the total population was estimated at 200–400. Since then the subspecies has continued to be heavily hunted. Today there are perhaps 40 remaining in Cangandala, with a similar number in the Luando Reserve. D’Anchieta’s fruit bat (Plerotes anchietae) is known only from a few scattered localities in Angola, the Republic of Congo, the Democratic Republic of Congo, Malawi, and Zambia. Ansell’s epauletted fruit bat (Epomophorus anselli) is known only from Malawi, where it was collected in Kasungu National Park in the 1980s and from the Karonga area in 1961. The Sakeji horseshoe bat (Rhinolophus sakejiensis) is known only from its original collection in north-western Zambia. The Gabela helmet-shrike (Prionops gabela) was long known only from a small area near Gabela, in central coastal Angola. The species was unrecorded after the mid-1970s until 2003, when a few small flocks began to be discovered sporadically over a relatively wide but narrow belt of lowland dry forest. The black-tailed cisticola (Cisticola melanurus) is a rare type of warbler found in north-eastern Angola and southeastern Democratic Republic of Congo. The Tanzanian rough-scaled lizard (Ichnotropis tanganicana) is known only from a single specimen collected in 1917 from an undefined locality east of Lake Tanganyika in western Tanzania. The Katavi burrowing skink (Typhlacontias kataviensis) is known only from sandy ridges within Katavi National Park in south-eastern Tanzania. Mopane Scrub and Woodlands Mopane (Colophospermum mopane), also known as balsam tree, butterfly tree, or turpentine tree, is a type of legume that grows in hot, dry, usually low-lying areas in the far northern parts of southern Africa. Varying in height between 4 and 18 m, its distinctive butterfly-shaped leaf and thin seed pod make it easy to identify. There are two main regions where mopane is the predominant vegetation, one in south-western Angola and northern Namibia and the other extending over the lowlands of the Zambezi River and its tributaries in Botswana,

Namibia, Zambia, Zimbabwe, Mozambique, Malawi, northern South Africa, and Swaziland. The plant is extensively used for firewood. The black-cheeked lovebird (Agapornis nigrigenis) is confined to a small area of south-western Zambia. Sudanian Savanna Sudanian savanna is a broad belt of tropical savanna running east to west across Africa. It consists of two main regions separated by the Cameroon Highlands. West Sudanian Savanna runs from the Atlantic Ocean in southern Senegal and Gambia through south-western Mali, north-eastern Guinea, southern Burkina Faso, north-eastern Ivory Coast, northern Ghana, northern Togo, northern Benin, south-western Niger, and parts of Nigeria. The Takwashara worm lizard (Cynisca nigeriensis) is known only from a single locality in central Nigeria. Joger’s saw-scaled viper (Echis jogeri) is confined to southeastern Senegal and south-western Mali. The Gabou file snake (Mehelya gabouensis) is known only from two specimens collected in 1996 from south-western Senegal. The Senegal worm lizard (Cynisca senegalensis) is known only from Niokolo-Koba National Park in south-eastern Senegal. East Sudanian Savanna consists of two main blocks in eastcentral Africa. The western block covers portions of northern Cameroon, southernmost Chad, northern Central African Republic, and South Sudan. The eastern block lies in a belt stretching from northern Uganda along the Sudan/Ethiopia border region. Vegetation typically consists of deciduous Terminalia trees with an undergrowth of shrubs and a wide array of grasses. Hoogstraal’s striped grass mouse (Lemniscomys hoogstraali) is known only from a single specimen collected in 1961 from what is now north-eastern South Sudan. Flooded Tropical Grasslands Important areas of flooded tropical grassland include the Okavango Delta in north-central Botswana, the Kafue Flats in central Zambia, and the Bangweulu Flats of northeastern Zambia. The Somali white-collared guenon (Cercopithecus albogularis zammaranoi) is confined to a small area of flooded grassland in southern Somalia along the Jubba and Shabelle rivers. The southern lechwe (Kobus leche) is a small, semi-aquatic antelope admirably adapted to the inundated flood plains and wetlands of south-central Africa. There are a number of isolated subspecies, two of which are now extinct and three others that are considered threatened. Roberts’ southern lechwe (K. l. robertsi) is believed to have been restricted to the Luongo and Kalungwishi drainage systems of the lower Luapula River of north-eastern Zambia, where it has been extinct for decades. The Cape southern lechwe (K. l. venterae) is a poorly known

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and somewhat controversial form, variously considered either as a species in its own right or not recognized as distinct at all. However it is to be classified, the population occurred widely within suitable habitat in the North West, Gauteng, Free State, Northern Cape, and Eastern Cape provinces of South Africa. Long gone from these areas, it was first described by the paleontologist Robert Broom from a frontlet and horn core discovered from Haggenstad (now known as Florisbad). Another subspecies, the red southern lechwe (K. l. leche) is currently widespread and relatively common in the wetlands of Zimbabwe, Botswana, Namibia, and Zambia. Fox’s weaver (Ploceus spekeoides) is a colonial-breeding passerine bird known only from a small area of seasonally flooded grasslands in east-central Uganda. The Kilombero weaver (P. burnieri) is confined to a few areas within the Kilombero River floodplain in south-central Tanzania. Both are threatened by loss of habitat. The Waza running frog (Kassina wazae) is known only from a single ephemeral marsh within seasonally flooded grassland near Waza, in northern Cameroon. The Kafue Flats are a vast, seasonally inundated floodplain on the middle Kafue River located in central Zambia (Southern, Central, Western, and North-Western provinces). Some 240 km long and about 50 km wide, it floods to a depth of less than a metre during the rainy season (deeper in some lagoons and permanent swampy areas), drying out to a clayey black soil in the dry season. The Kafue lechwe (Kobus leche kafuensis) is confined to the Kafue Flats, where its population has been monitored for almost a century. From an estimated 250,000 in 1934 it was reduced to a low of around 25,000 by the 1960s, but increased once more to between 90,000 and 110,000 during the early 1970s. By the early 1980s the numbers had been reduced again to about 40,000–45,000, then increased gradually to 78,000 by the end of the twentieth century. At this point the population suddenly crashed, with just under 29,000 estimated in 2015. While poaching has been a factor in the decline, maintenance of a seasonal flooding regime is critically important to the survival of this subspecies. Water flow has been regulated almost completely in this region according to human needs since construction of hydroelectric dams in the 1970s, and significant alteration to the current hydrological status could prove catastrophic. The Kafue mole-rat (Fukomys kafuensis) is known only from a small area of the Kafue Flats in Southern province. The Kafue worm lizard (Zygaspis kafuensis) is known only from a few specimens collected in the vicinity of the Kafue Flats. The Bangweulu Flats are an extensive wetland area located adjacent to Lake Bangweulu in north-eastern Zambia. The Bangweulu tsessebe (Damaliscus lunatus superstes) is a type of antelope confined to the Bangweulu Flats. The black lechwe (Kobus leche smithemani) is today confined to the southern half of the Bangweulu Flats, although at one time it was also found on the Chambeshi floodplains along

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the upper Luapula between Zambia and south-eastern Democratic Republic of Congo, where it is unlikely to survive. In terms of population it has declined from about 150,000 in 1932 to no more than 4000 in 1966. By the early 1980s it had increased to around 40,000 but subsequently decreased to 30,000 by the end of that decade. Today the population is more or less stable at between 35,000 and 50,000. However, it remains vulnerable to poaching and is totally dependent on the maintenance of a favourable hydrological regime.

Lowland Subtropical Grasslands and Savannas Lowland subtropical grasslands and savannas are found patchily in Angola, northern and central Namibia, northern and eastern Botswana, southern Zambia, Zimbabwe, south-western Mozambique, and central and north-eastern South Africa. Historically they were much more extensive. When Europeans first began settling in southern Africa three and a half centuries ago, they gradually spread inland from the coast. There they found rich grasslands and a wealth of wild animals. Intensified farming and the raising of cattle and sheep led to the occupation of more land and the driving out of the wild mammals. The opening up of the country to settlers increased the demand for meat. Many men became professional hunters; others killed for fun. Hecatombs of wild mammals fell before their guns. The intensive exploitation of the land produced a serious decline in grass cover and soils along with the wildlife. The southern or Cape black rhinoceros (Diceros bicornis bicornis) was the largest black rhinoceros subspecies. Once abundant from the Cape of Good Hope to Transvaal, South Africa and probably into southern Namibia, it became extinct due to excessive hunting and habitat destruction around 1850. The South African or Cape giraffe (Giraffa camelopardalis giraffa) is found in northern South Africa, southern Botswana, southern Zimbabwe, and south-western Mozambique. A maximum of 31,500 are estimated in the wild. The Cape buffalo (Syncerus caffer caffer) is found patchily in eastern and southern Africa, where it is threatened by habitat destruction, hunting, drought, and disease. The eastern and southern African populations of the African lion (Panthera leo) are nowadays included together as a distinct subspecies (P. l. melanochaita), where they live in a variety of habitats. The southernmost population, known as the Cape lion, inhabited South Africa’s Natal and Cape provinces. In the early nineteenth century lions still occurred in the karoo plains and in the northern Cape. The last individuals reported south of the Orange River were sighted between 1850 and 1858, but they may have survived in the northern Orange Free State into the 1860s. In any case they were soon wiped out by hunting. As with the Barbary lion population discussed earlier in this book, it is possible that some zoo specimens with Cape lion ancestry may yet survive. The bluebuck (Hippotragus leucophaeus) was the first African antelope to be hunted to extinction by European settlers. Historically limited to a small area of what was then grassland on the south-western Cape coast, it first became

The African Region

South Africa as well as in Namibia, Botswana, and Zimbabwe. The bontebok and the blesbok are no longer considered threatened. The Cape wild dog (Lycaon pictus pictus) is found patchily in south-western Mozambique, south-eastern Zimbabwe, and north-eastern South Africa (Limpopo, Mpumalanga, and KwaZulu-Natal). The rough-haired golden mole (Chrysospalax villosus) is known only from a few scattered localities in eastern and north-eastern South Africa (Eastern Cape, KwaZulu-Natal, Gauteng, and Mpumalanga). The white-chested tinkerbird (Pogoniulus makawai) is known only from a single specimen collected in 1964 from north-western Zambia.

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Figure 3.9 Drawing of a bluebuck museum specimen by François Levaillant, 1781. (Credit: François Levaillant.)

known in 1719 at which time it was already rare. The last survivors were shot around 1800. Only four stuffed specimens are preserved in European museums. The bontebok (Damaliscus pygargus pygargus) is a beautiful antelope that was historically confined to the coastal plain of the Western Cape in South Africa, where overhunting reduced it from locally abundant to the verge of extinction. It was saved from extermination during the mid-nineteenth century by a few Cape farming families, who protected the small, remnant population living on the poor grazing lands where it had been driven. In 1931 Bontebok National Park was established near Bredasdorp, where the original stock of just 17 animals grew to 84 by 1960, at which point the animals were translocated to the more suitable site of the current Bontebok National Park near Swellendam. From there the population grew to 320 as of 1981. Numbers have since been maintained at around 250, with surplus individuals forming the nucleus of reintroduced populations in other protected areas and private farms. The total number of bontebok now stands at around 3500, though only about 1500 actually occur within the historical range. The largest subpopulation is in the De Hoop Nature Reserve and environs. Another subspecies, the blesbok (D. p. phillipsi), formerly lived in an area some 300 km north and east of the bontebok’s range in the highveld of Free State and Gauteng provinces, parts of KwaZulu-Natal, and the northern karoo in the Eastern and Northern Cape, South Africa. It was also present in western Lesotho and possibly Swaziland. Originally quite abundant, it too was decimated by overhunting and reduced to around 2000 by the late nineteenth century. It has since made a spectacular recovery, and has been translocated to private game farms throughout

Areas of lowland dry grassland within the African Region are to be found in a narrow band across the whole of northern Africa in northern Senegal, southern Mauritania, southcentral Mali, southern Niger, central Chad, and Sudan, with further areas within the Horn of Africa extending to Tanzania. Other pockets occur in Namibia, Botswana, and South Africa. The black-faced impala (Aepyceros melampus petersi) is confined to north-western Namibia and south-western Angola. Brockman’s gerbil (Gerbillus brockmani) is known only from its original collection in the early twentieth century from north-western Somalia. The Somali gerbil (G. somalicus) is known only from two historical records (north-western Somalia and Djibouti). The Somali elephant shrew (Galegeeska revoilii) was historically known only from northern Somalia, where it was last recorded after 1968 and thought to be extinct. A small population was rediscovered in Djibouti in 2020. The southern shield-backed lizard (Philochortus rudolfensis) is known only from five specimens collected in the early twentienth century from arid bushland areas of northern Kenya. Campbell’s girdled lizard (Namazonurus campbelli) is confined to a small area of south-central Namibia. Hughes’ saw-scaled viper (Echis hughesi) is known only from a single locality near the tip of the Horn of Africa in Somalia. The Somali snake (Aeluroglena cucullata) is known only from five specimens collected from northern Somalia. The Kenyan dwarf blind snake (Afrotyphlops nanus) is known only from its original collection during the nineteenth century from arid bushland in south-eastern Kenya. The Turkana toad (Sclerophrys turkanae) is known only from the south-eastern shores of Lake Turkana and from near the Ewaso Ng’iro River in northern and central Kenya.

The Sahel The Sahel is essentially a semi-arid transition zone between the Sahara Desert to the north and the Sudanian savanna to the

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south. It stretches across northern Africa between the Atlantic Ocean and the Red Sea, roughly in northern Senegal, southern Mauritania, central Mali, northern Burkina Faso, southern Niger, extreme northern Nigeria, extreme northern Cameroon, central Chad, Sudan, and Eritrea. The red-fronted gazelle (Eudorcas rufifrons) was historically common across the Sahel, but has now been reduced to scattered pockets due to hunting, competition with domestic livestock and habitat loss due to drought and clearance for agriculture. A number of ill-defined subspecies have been described, including the Senegal red-fronted gazelle (E. r. rufifrons), the eastern Chad red-fronted gazelle (E. r. centralis), the north Nigeria red-fronted gazelle (E. r. hasleri), the Kanuri red-fronted gazelle (E. r. kanuri), and the Nubian red-fronted gazelle (E. r. laevipes). The so-called Atlas red-fronted gazelle (E. r. rufina) is a rather mysterious form known only from three skins bought in Algeria toward the end of the nineteenth century. Never observed in the wild, it may have lived in mountainous forests of the Algerian– Moroccan border region and was said to be well-known to furriers in Oran. Variously treated either as a colour variation or subspecies of red-fronted gazelle (as here) or as a distinct species in its own right, the most plausible explanation is that the purchased skins had been transported to market from somewhere further south. Etheridge’s blind snake (Xerotyphlops etheridgei) is known only from a single locality in west-central Mauritania.

Deserts and Semi-Deserts Deserts, subdeserts, and arid scrubs cover about one-third of the continent of Africa. In the north, of course, there is the immense Sahara Desert, which is dealt with elsewhere in this book. Apart from that there are the Namib and Kalahari deserts of the south-west, the karoo subdesert of the south, and the Danakil and other large deserts of the Horn of Africa. The man-made subdeserts of Kenya, Tanzania, Botswana, Lesotho, and elsewhere in the African Region are also included here.

Karoo The karoo is an ill-defined semi-desert area of south-western South Africa and southern Namibia. It is usually subdivided into the Great Karoo to the north of the Swartberg Mountains and Little Karoo to the south. The latter is separated from the sea by the east–west running Outeniqua–Langeberg Mountains. Characterized mainly by its low rainfall, the karoo region was once covered by grasslands but has expanded to its present size as a result of long overuse by sheep and goats. The riverine rabbit (Bunolagus monticularis) is a highly threatened species that inhabits the growth along seasonal river courses in the central karoo. The total population is thought to be less than 1500. Visagie’s golden mole (Chrysochloris visagiei) is known only from a single specimen collected from an area of

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Northern Cape province that has since been dramatically disrupted by agriculture. The ferruginous sand lark (Calendulauda burra) is a rare species confined to the karoo of western South Africa (Northern Cape). It is threatened by loss of habitat. The karoo dwarf chameleon (Bradypodion karrooicum) is confined to a small area of central South Africa (eastern Northern Cape and western Free State). The dwarf karoo girdled lizard (Cordylus aridus) is confined to a small area of the southern karoo in south-central South Africa (Western Cape). Fisk’s house snake (Lamprophis fiskii) is a little-known species from the south-western karoo, where it may be affected by habitat destruction.

The Danakil Desert The Danakil Desert lies in the Afar Triangle and covers northeastern Ethiopia, southern Eritrea, southern Djibouti ,and the north-westernmost part of Somalia. It is known for its active volcanoes and extreme heat. The Somali wild ass (Equus africanus somaliensis) was historically found in Somalia, Ethiopia, Eritrea, and Djibouti, where it was considered common up to the mid-twentieth century. By 1980 it had been reduced to perhaps 6000 or 7000 living mainly in northern Somalia and the Danakil subdeserts and Awash basin of Ethiopia. As with the Nubian wild ass this sharp decline was due mainly to excessive hunting for its meat and hides (exacerbated by the endless warfare that has plagued this part of Africa since 1935), along with drought and the resulting competition for limited food and water from increasing numbers of livestock. Today there are thought to be only a few hundred wild asses living in the Ethiopia/Eritrea border region; its status (or even survival) in Somalia is unknown. Another 200 or so live in captivity. The beira antelope (Dorcatragus megalotis) is a small species found only in the arid coastal areas of southern Djibouti, northern Somalia and, marginally, eastern Ethiopia (where it was last observed in 1972). Considered rare even at the time of its discovery in 1885, it has since suffered further declines due mainly to habitat destruction and competition with livestock.

The Namib Desert The Namib is a coastal desert that, according to its broadest definition, extends for some 2000 km along the Atlantic coasts of southern Angola and of Namibia, along with a small area of adjacent western South Africa. Its northernmost portion, located along the Angola–Namibia border, is known as the Moçâmedes Desert, while its southern portern approaches the neighbouring Kalahari. Having endured arid or semi-arid conditions for roughly 55–80 million years it may be the oldest desert in the world, and contains some of its driest regions. It remains almost completely untouched by humans. The Nama dwarf tortoise (Chersobius solus) has a very restricted distribution in the extreme deserts of southern

The African Region

Namibia, but is not currently thought to be threatened beyond occasional collection for the international pet trade. The Namaqua dwarf adder (Bitis schneideri), perhaps the world’s smallest viper, is confined to the coastal dunes of the south-western Namibia/South Africa border region. It is potentially threatened by future mining activity.

The Kaokoveld Desert The Kaokoveld Desert is located in north-western Namibia and southern Angola, east of the Namib Desert. The Damaraland pygmy toad (Poyntonophrynus damaranus) is confined to the Kaokoveld Desert. The Damaraland sand frog (Tomopterna damarensis) is known only from a small area of north-western Namibia.

Lakes, Rivers, and Marshes The major lakes and rivers within the African Region include Lake Victoria and Lake Malawi, along with the upper Nile and Zambezi river drainages, in addition to numerous smaller, more ephemeral ones. The Cape clawless otter (Aonyx capensis) is the largest Old World otter species and one of the most widely distributed, being found over much of sub-Saharan Africa outside the rainforest belt. It has suffered significant declines however due to hunting, habitat destruction, and pollution. Hopkins’ groove-toothed swamp rat (Pelomys hopkinsi) is confined to swampy areas between Lake Victoria and the Albertine Rift Valley. It has been recorded from western Kenya, Rwanda, and Uganda, and possibly also occurs in Burundi and Tanzania. The wattled crane (Bugeranus carunculatus) is found patchily in wetland areas across much of sub-Saharan Africa outside of the rainforest belt, but is everywhere declining due to loss of habitat. The shoebill (Balaeniceps rex) is a large, stork-like bird so named for its enormous shoe-shaped bill. Known to both the ancient Egyptians and medieval Arabs, the species was, however, only classified in the mid-nineteenth century after skins and, eventually, live specimens were brought to Europe. Widely but very locally distributed in large swamps from South Sudan to Zambia, the total population is thought to be between 5000 and 8000. Threats include habitat destruction, disturbance, and hunting. The slaty egret (Egretta vinaceigula) is a rare species found primarily Zambia and northern Botswana, with occasional vagrants reported from north-eastern South Africa, southwestern and central Mozambique, Zimbabwe, southern Angola, and northern Namibia. The Maccoa duck (Oxyura maccoa) is found widely but patchily across the African Region, where it has been declining steadily for decades due to loss of habitat and pollution. The white-winged flufftail (Sarothrura ayresi) is an extremely rare rail-like bird with a highly disjunct range, being known only from seasonal marshlands in central Ethiopia and

eastern South Africa, with a few additional historical reports from Zimbabwe. The Zambezi flapshell turtle (Cycloderma frenatum) is confined to the rivers and lakes of east-central Africa, in particular the Zambezi River drainage. The Cape clawed frog (Xenopus gilli) is confined to a few localities in extreme south-western South Africa (Western Cape). Tweddle’s cichlid (Haplochromis tweddlei) is known only from lakes Chiuta and Chilwa and associated rivers in southeastern Malawi and northern Mozambique, where it is threatened by overfishing and drought. The lowveld largemouth cichlid (Serranochromis meridianus) is confined to north-eastern South Africa and southern Mozambique. It is threatened by loss of habitat. Several species of commercially important tilapia cichlids (Oreochromis) have been severely depleted by overfishing and other factors. The greenhead tilapia (O. macrochir) and the threespot tilapia (O. andersonii) are both native to the Zambezi River drainage and associated lakes in south-central Africa, from where they have been translocated to many other parts of southern Africa and even Madagascar. They are threatened by competition with introduced Nile tilapia (O. niloticus). Trewavas’ tilapia (O. lidole), Günther’s tilapia (O. squamipinnis), and the Karonga tilapia (O. karongae) were each historically found in Lake Malawi, Lake Malombe, and the upper and middle Shire River, with the latter also present in two crater lakes in Tanzania (Ikapu and Itamba). Both are now close to extinction. Hilgendorf’s tilapia (O. amphimelas) is confined to Lake Manyara and a few other small Rift Valley Lakes in north-central Tanzania. Graham’s tilapia (O. esculentus) was historically found within the Lake Victoria drainage where it has been virtually extirpated apart from a few smaller satellite lakes due to overfishing and introduced species. The species has, however, been introduced into the Pangani River drainage of south-eastern Kenya and north-eastern Tanzania. The soda tilapia (Alcolapia alcalicus) is confined to the hypersaline Lake Natron drainage, Lake Magardi, and the Shombole swamps of southern Kenya and northern Tanzania. The Guineas tilapia (Tilapia guinasana) was historically confined to Lake Guineas, a small sinkhole lake located in north-central Namibia. The species has since been introduced into Lake Otjikoto and other farm dams. It is threatened mainly by groundwater extraction. The rhinoceros yellowfish (Labeobarbus rhinoceros) is confined to the Athi and Tana River drainage in Kenya, where it is heavily fished. Four barbs of the genus Enteromius are threatened by habitat destruction and degradation as well as introduced species. The Marico barb (E. motebensis) is confined to small streams within the Marico, Elands, and Crocodile (West) rivers of northern South Africa (North West and Gauteng provinces). The Zalbi barb (E. zalbiensis) is confined to the Benue River in northern Cameroon, the Mayo-Kebbi River of Chad, and the Chari River of Chad and Central African

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Republic. The Amatola barb (E. amatolicus) is known only from the Kei and Mbashe rivers of south-eastern South Africa (Eastern Cape). The silver barb (E. choloensis) is confined to a few rivers and streams in north-eastern Mozambique and southern Malawi. The Somalian blind barb (Barbopsis devecchi) is known from a few streams, springs, and wells in the Nogal Valley of north-central Somalia. Several species of redfin (Pseudobarbus) are endemic to southern Africa, where they are threatened by loss of habitat and invasive species. The Eastern Cape redfin (P. afer) is confined to the Baakens, Swartkops, and Sundays river drainages of southern coastal South Africa (Eastern Cape). The smallscale redfin (P. asper) is confined to the Gamtoos and Gouritz river drainages of southern South Africa. The Maloti redfin (P. quathlambae) is confined to a few river drainages in Lesotho and eastern South Africa (KwaZulu-Natal). The border redfin (P. trevelyani) is confined to the Keiskamma and Buffalo river drainages of south-eastern South Africa (Eastern Cape). The Cape redfin (P. capensis) is confined to the Berg and Breede river drainages of south-western South Africa (Western Cape). Bottego’s sardine (Neobola bottegoi) is known only from Lake Turkana and the Omo River of Ethiopia and the Jubba River of Somalia. The Eastern Cape rocky (Sandelia bainsii) is confined to a few river drainages in south-eastern South Africa (Eastern Cape). Andruzzi’s cavefish (Phreatichthys andruzzii) is confined to a few isolated wells in the vicinity of Bud-bud Uaesele in south-central Somalia. The Tigray garra (Garra duobarbis) is confined to the Dirma River and one reservoir in north-western Ethiopia. Leleup’s robber tetra (Rhabdalestes leleupi) is confined to Lake Nyumba ya Mungu and its outlet to the Pangani River in north-eastern Tanzania, where it is seriously threatened by overfishing and habitat deterioration. The Sibayi goby (Silhouettea sibayi) is confined to a few coastal freshwater lakes in north-eastern South Africa (KwaZulu-Natal) and southern Mozambique. Several species of killifish (Nothobranchius) are threatened by loss of their seasonal freshwater habitats. Lueke’s killifish (N. luekei) and the red-finned killifish (N. rubripinnis) are both known only from seasonal pools and swamps within the Mbezi and Luhule/Luhute river drainages of eastern coastal Tanzania. The elongated killifish (N. elongatus) and the Kikambala killifish (N. interruptus) are both confined to seasonal pools and marshes in south-eastern coastal Kenya. The Somali killifish (N. fasciatus) is confined to temporary pools, marshes, and reservoirs in south-eastern Somalia. The arrow killifish (N. sagittae) is confined to temporary pools and swamps within the Lake Victoria drainage of northern Tanzania. The Pwani killifish (N. albimarginatus) and the Rufiji killifish (N. annectens) are both confined to a few disjunct localities in central coastal Tanzania.

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The East Coast lampeye (Pantanodon stuhlmanni) is confined to the lower reaches of river and brooks in the coastal areas of south-eastern Kenya and north-eastern Tanzania, including salt pans. The Kibiti lampeye (Aplocheilichthys lacustris) is confined to Lake Kibiti in the Luhute River drainage and to the Mbezi River drainage in eastern coastal Tanzania. The giant catfish (Pardiglanis tarabinii) is known from the Juba and Shebelle rivers of Somalia and the lower Tana River of Kenya, with an additional possible record from the Ewaso Ng’iro River. The bifurcated suckermouth catfish (Chiloglanis bifurcus) historically occurred throughout the Crocodile, Elands, and Inkomati river drainages of north-eastern South Africa (Mpumalanga), Swaziland, and south-western Mozambique. The species has undergone a considerable decline due to dam construction and introduced fish species, and is now confined to a few higher-elevation localities. Engelsen’s airbreathing catfish (Clarias engelseni) is known only from its original collection in the 1920s from the Yei River of eastern Sudan. The Aigamas Cave airbreathing catfish (C. cavernicola) is known only from a small, deep pool within Aigamas Cave in northern Namibia, where it is threatened by water extraction. The total population is estimated at between 200 and 400. Zammarano’s cave catfish (Uegitglanis zammaranoi) is confined to a few caves near the Jubba and Shebelle rivers in central Somalia.

Lake Afrera Lake Afrera (formerly known as Lake Giulietti) is a hypersaline lake located in northern Ethiopia (Afar region). General threats include water extraction and salt mining. The Afrera cichlid (Danakilia franchettii) is confined to freshwater hot springs on the shores of Lake Afrera and nearby swamps. The Afrera toothcarp (Aphanius stiassnyae) is confined to the freshwater hot springs of Lake Afrera.

Lake Magadi Lake Magadi is a hot, saline, slightly alkaline lake located in south-central Kenya. Sir David Attenborough once noted that it was perhaps the most hostile environment on Earth still capable of supporting life. The Magadi tilapia (Alcolapia grahami) is a type of cichlid historically confined to the springs and lagoons around Lake Magadi. In the 1950s and 1960s it was also introduced into Lake Elmenteita and Lake Nakuru in Kenya and to Lake Natron in Tanzania (where it does not appear to have become established). It is everywhere declining.

Lake Natron Lake Natron is a salt and soda lake in the Arusha region of northern Tanzania. While most animals find the lake’s high temperature (up to 60°C) and salt content inhospitable, a few

The African Region

species of cichlid manage to survive in the slightly less salty water around its margins. The wide-lipped Natron tilapia (Alcolapia latilabris) and the narrow-mouthed Natron tilapia (A. ndalalani) are both confined to southern areas of the Lake Natron basin, where they inhabit a few springs and effluents.

Lake Victoria Lake Victoria is Africa’s largest lake and the world’s second largest by surface area. Located in a shallow depression, it has a maximum depth of just 80–85 m and is surrounded by three countries (Kenya, Uganda, and Tanzania). The lake historically had an incredible number of endemic fish species including over 350 types of mouthbrooding cichlid, whose extraordinary diversity and speed of evolution were of enormous interest to those concerned with the forces that create and maintain the richness of life everywhere, and which served a vital role in the lake’s bountiful ecosystem. Unfortunately, during the twentieth century the latter was dramatically altered by a combination of factors. In the 1950s a proposal to increase fish catches in the lake by introducing the Nile perch (Lates niloticus), known locally as mbutta or sangara, was adamantly opposed by scientists, who feared that the lack of a natural predator for this exotic species would devastate the native ones. Despite the controversy, a colonial fisheries officer was ordered to clandestinely put the perch into the Ugandan portion of the lake in 1952. Thereafter, it was introduced intentionally in both 1962 and 1963. By 1964 perch were recorded in Tanzania, by 1970 they were well established in Kenya, and by the early 1980s were abundant throughout Tanzania, Kenya, and Uganda. Today, virtually all of the natural, biological wealth unique to Lake Victoria has been destroyed. Only a handful of the original fish species still survive, and their loss has resulted in eutrophication of the water and an increasing frequency of algal blooms. The variable tilapia (Oreochromis variabilis) is still found in some localized portions of Lake Victoria and a few satellite lakes and ponds. As mentioned above, it is the endemic mouthbrooding cichlids of the genus Haplochromis that have been hardest hit by changes to the lake. The following forms are most likely extinct: the thick-lipped cichlid (H. crassilabris), not seen since 1980; Linnaeus’ cichlid (H. vonlinnei), not seen since 1980; Ushind’s cichlid (H. ushindi), not seen since 1980; Plutonian cichlid (H. plutonius), not seen since 1980; the hiatus cichlid (H. hiatus), not seen since 1980; the iris cichlid (H. iris), not seen since 1980; the crowned cichlid (H. cinctus), not seen since 1982; the coprologus cichlid (H. coprologus), not seen since 1982; Katunzi’s cichlid (H. katunzii), not seen since 1982; the silvery cichlid (H. argenteus), not seen since 1983; the mylergates cichlid (H. mylergates), not seen since 1983; the pyrrhopteryx cichlid (H. pyrrhopteryx), not seen since 1983; Cnester cichlid (H. cnester), not seen since 1983; the differentmouthed cichlid (H. xenostoma), not seen since 1983; the yellowfin cichlid (H. flavipinnis), not seen since 1984; the

ptistes cichlid (H. ptistes), not seen since 1984; Greenwood’s cichlid (H. aelocephalus), not seen since 1985; Heusinkveld’s cichlid (H. heusinkveldi), not seen since 1985; Michael Graham’s cichlid (H. michaeli), not seen since 1985; the obese cichlid (H. obesus), not seen since 1985; Barbara William’s cichlid (H. barbarae), not seen since 1985; Guiart’s cichlid (H. guiarti), not seen since 1985; Martin’s cichlid (H. martini), not seen since 1985; the yellow-draped cichlid (H. crocopeplus), not seen since 1985; the large-jawed cichlid (H. macrognathus), not seen since 1985; the small-toothed cichlid (H. microdon), not seen since 1985; the sulfurous cichlid (H. sulphureus), not seen since 1985; the Victorianus cichlid (H. victorianus), not seen since 1985; the long-snouted cichlid (H. longirostris), not seen since 1985; Cassius’ cichlid (H. cassius), not seen since 1986; Teunisras’ cichlid (H. teunisrasi), not seen since 1986; Teegelaar’s cichlid (H. teegelaari), not seen since 1986; Grant’s cichlid (H. granti), not seen since 1986; the dichrourus cichlid (H. dichrourus), not seen since 1986; the wasp cichlid (H. sphex), not seen since 1986; the dwarf sea bass cichlid (H. nanoserranus), not seen since 1986; Perrier’s cichlid (H. perrieri), not seen since 1986; the pancitrinus cichlid (H. pancitrinus), not seen since 1986; the perch-like cichlid (H. percoides), not seen since 1986; the dentex cichlid (H. dentex), not seen since 1987; Brown’s cichlid (H. brownae), not seen since the 1980s; George Ishmael’s cichlid (H. ishmaeli), not seen since 1991; the theliodon cichlid (H. theliodon), not seen since 1993; the small-toothed cichlid (H. parvidens), not seen since 1995. The surviving forms (if indeed they do survive) must all be considered seriously threatened: these include the red-headed cichlid (H. erythrocephalus), the few-scaled cichlid (H. oligolepis), Paraguiart’s cichlid (H. paraguiarti), the arcane cichlid (H. arcanus), Artaxerxes’ cichlid (H. artaxerxes), the azure cichlid (H. azureus), Barel’s cichlid (H. bareli), Barton’s cichlid (H. bartoni), Bayon’s cichlid (H. bayoni), the bicolour cichlid (H. bicolor), Bwathondi’s cichlid (H. bwathondii), the hollow-fronted cichlid (H. cavifrons), the grey cichlid (H. cinereus), the twobanded cichlid (H. diplotaenia), Fischer’s cichlid (H. fischeri), the giant cichlid (H. gigas), Gilbert’s cichlid (H. gilberti), Goldschmidt’s cichlid (H. goldschmidti), Gowers’ cichlid (H. gowersii), Howes’ cichlid (H. howesi), Kujunju’s cichlid (H. kujunjui), the large-headed cichlid (H. macrocephalus), Maisome’s cichlid (H. maisomei), the black-finned cichlid (H. melanopus), the small-headed cichlid (H. microcephalus), the blackish cichlid (H. nigrescens), the cloudy cichlid (H. nubilus), the Nyanza cichlid (H. nyanzae), the thick-headed cichlid (H. pachycephalus), the pallid cichlid (H. pallidus), Pellegrin’s cichlid (H. pellegrini), Pitman’s cichlid (H. pitmani), the prognathus cichlid (H. prognathus), Sauvage’s cichlid (H. sauvagei), Speke’s cichlid (H. spekii), the trident cichlid (H. tridens), Vanoijen’s cichlid (H. vanoijeni), Welcomme’s cichlid (H. welcommei), the acidens cichlid (H. acidens), the argens cichlid (H. argens), the altigenis cichlid (H. altigenis), the antleter cichlid (H. antleter), the apogonoides cichlid (H. apogonoides), the cow-eyed cichlid (H.

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boops), the greenish cichlid (H. chlorochrous), the chromogynos cichlid (H. chromogynos), the chrysogynaion cichlid (H. chrysogynaion), the Cronos cichlid (H. cronus), the hiddentoothed cichlid (H. cryptodon), the cryptogramma cichlid (H. cryptogramma), the decticostoma cichlid (H. decticostoma), the dolichorhynchus cichlid (H. dolichorhynchus), the empodisma cichlid (H. empodisma), the jealous cichlid (H. estor), the eutaenia cichlid (H. eutaenia), the spindle-shaped cichlid (H. fusiformi), Van Oijen’s cichlid (H. harpakteridion), the lowly cichlid (H. humilior), the igneopinnus cichlid (H. igneopinnis), the labriformis cichlid (H. labriformis), the weeping cichlid (H. lacrimosus), the leaden-blue cichlid (H. lividus), the yellow cichlid (H. luteus), the large-eyed cichlid (H. macrops), the spot-eared cichlid (H. maculipinna), the mandibular cichlid (H. mandibularis), the jaw cichlid (H. maxillaris), the honey-yellow cichlid (H. melichrous), the largeeyed cichlid (H. megalops), the black-finned cichlid (H. melanopterus), the chin cichlid (H. mento), the Nile cichlid (H. niloticus), Hilgendorf’s cichlid (H. nuchisquamulatus), the obliquidens cichlid (H. obliquidens), the blunt-toothed cichlid (H. obtusidens), the paraplagiostoma cichlid (H. paraplagiostoma), the parorthostoma cichlid (H. parorthostoma), the pharyngomylus cichlid (H. pharyngomylus), the plant-eating cichlid (H. phytophagus), the piceatus cichlid (H. piceatus), the oblique-toothed cichlid (H. plagiodon), the plagiostoma cichlid (H. plagiostoma), the precursor cichlid (H. prodromus), the pseudo-Pellegrin’s cichlid (H. pseudopellegrini), the retrodens cichlid (H. retrodens), the saxicola cichlid (H. saxicola), the serranus cichlid (H. serranus), the simotes cichlid (H. simotes), the small-scaled cichlid (H. squamulatus), the thereuterion cichlid (H. thereuterion), the thuragnathus cichlid (H. thuragnathus), the purple-velvet cichlid (H. tyrianthinus), the Victoria cichlid (H. victoriae), and the xanthopteryx cichlid (H. xanthopteryx). The Lake Victoria labeo (Labeo victorianus) is confined to the Lake Victoria basin, where it is seriously threatened by overfishing and loss of habitat. Loveridge’s barb (Xenobarbus loveridgei) is known only from three specimens collected from off the southern shore of Lake Victoria in the 1920s. The Lake Victoria deepwater catfish (Xenoclarias eupogon) has not been reported since 1997, and is possibly extinct. The Lake Victoria snake catfish (Clariallabes petricola) is known only from the northern part of Lake Victoria and from an area of the Victoria Nile near Owen Falls. Lake Nabugabo Lake Nabugabo is a small satellite lake of Lake Victoria located in central Uganda. It formed as a result of sand dunes resulting from strong winds which separated the two around 5000 years ago. Simpson’s cichlid (Haplochromis simpsoni), Beadle’s cichlid (H. beadlei), the sailfin cichlid (H. velifer), the hunter cichlid (H. venator) and the linking-toothed cichlid (H. annectidens) are all confined to Lake Nabugabo.

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The Katonga River The Katonga River is located in south-western Uganda. Its channel is continuous between Lake Victoria and Lake George, reflecting that it once drained away from the former into the latter. The Katonga cichlid (Haplochromis katonga) is known only from the Katonga River. Degen’s mormyrid (Petrocephalus degeni) is known only from a single specimen collected in 1906 in the mouth of the Katonga River, Lake Victoria basin.

Lake Malawi Lake Malawi (also known as Lake Nyasa in Tanzania) is the southernmost lake in the East African Rift system. Located between eastern Malawi, western Mozambique, and southern Tanzania, it is home to more types of fish than any other lake in the world, including at least 700 species of cichlids. Lake Malawi National Park is located at the southern end of the lake. Friedemann’s reed frog (Hyperolius friedemanni) is known only from two localities on the shores of Lake Malawi (Karonga and Monkey Bay). Four cichlids of the genus Melanochromis endemic to small areas of Lake Malawi are threatened by collection for the international aquarium trade. The Membe Point cichlid (M. baliodigma) is known only from a single locality along the eastern shore of the lake. The Chipoka cichlid (M. chipokae) is known only from two localities in the southern part of the lake. The Masinje cichlid (M. dialeptos) and the Makanjila Point cichlid (M. lepidiadaptes) are both known only from a small area along the south-eastern shore of the lake. Several cichlids of the genus Labidochromis endemic to Lake Malawi are threatened by sedimentation and collection for the international aquarium trade. The Masimbwe cichlid (L. zebroides) is confined to Masimbwe Islet near Likoma Island in the eastern part of the lake, where the total population is thought to be less than 250. Freiberg’s cichlid (L. freibergi) occurs naturally only around Likoma Island in the eastern part of the lake, but has been introduced near Thumbi West Island. The Chizumulu cichlid (L. chisumulae) and the strigated cichlid (L. strigatus) occur naturally only around Chizumulu Island in the eastern part of the lake, although the latter has been introduced near Thumbi West Island. The Boadzulu cichlid (L. heterodon) is confined to Boadzulu Island in the southern part of the lake. The violet cichlid (L. ianthinus) is confined to Mbenji Island and Nkhomo Reef in the southern part of the lake. The livid cichlid (L. lividus) is confined to the western coast of Likoma Island in the eastern part of the lake. The Mbenji cichlid (L. mbenjii) is confined to Mbenji Island in the south-western part of the lake. The Maleri cichlid (L. pallidus) is confined to the Maleri Islands in the southern part of the lake. The mylodon cichlid (L. mylodon) is confined to Mumbo Island and Thumbi West Island in the southern part of the lake. Several cichlids of the genus Copadichromis endemic to Lake Malawi are threatened by overfishing with seine nets

The African Region

and collection for the international aquarium trade. The Nkata Bay cichlid (C. nkatae) is known only from specimens collected in 1960 from the western shore of the lake. Geerts’ cichlid (C. geertsi) is known only from four localities along the eastern shore of the lake. Verduyn’s cichlid (C. verduyni) is confined to a small area of the south-eastern coast of the lake. Konings’ cichlid (C. mbenji) is confined to Mbenji Island in the south-western part of the lake. Trewavas’ cichlid (C. trewavasae) is known from Likoma Island, Chizumulu Island, and a couple of other localities in the eastern part of the lake. The Boadzulu cichlid (Nyassachromis boadzulu) is confined to the south-eastern arm of Lake Malawi, where it is has been recored from off of Boadzulue Island, Kanchedza Island, Domwe Island, Crocodile Rocks, and Chemwesi Rock. Several cichlids of the genus Chindongo endemic to Lake Malawi are threatened by sedimentation and collection for the international aquarium trade. Saulos’ cichlid (C. saulosi) is confined to Taiwanee Reef north of Chizumulu Island in the eastern part of the lake. Demason’s cichlid (C. demasoni) is confined to Pombo and Ndumbi reefs on the north-eastern shore of the lake. Staeck’s cichlid (C. heteropictus) is confined to Chizumulu Island in the eastern part of the lake. Stauffer’s cichlid (C. ater) is confined to Chinyamwezi Island and Chinyankwazi Island in the south-eastern part of the lake. Seegers’ cichlid (C. longior) is confined to a small area along the north-eastern shore of the lake. Boulenger’s cichlid (Mchenga inornata) is known only from two specimens collected during the early twentieth century from an unknown locality within Lake Malawi. The Nankumba cichlid (M. cyclicos) from the Nankumba Peninsula in the southern part of the lake and the conophorus cichlid (M. conophorus) with an undefined distribution are said to be seriously threatened. The cruel cichlid (Rhamphochromis ferox) is known only from a few specimens collected from Lake Malawi and the Shire River. However, it has been established in the international aquarium trade. Axelrod’s cichlid (Cynotilapia axelrodi) is known only from Nkhata Bay and off Chirombo Point off the west-central shore of Lake Malawi. Several species of peacock cichlid (Aulonocara) endemic to Lake Malawi are threatened by collection for the international aquarium trade. The Vua peacock cichlid (Aulonocara auditor) is known only from a single specimen collected in the 1930s from the northern part of the lake. It is most likely extinct. The Kande peacock cichlid (A. kandeense) is known only from Kande Island in the western part of the lake. The Chitande peacock cichlid (A. ethelwynnae) is known only from near Chitande Island in the north-western part of the lake. The Chizumulu peacock cichlid (A. korneliae) is known only from near Chizumulu Island in the eastern part of the lake. The sulfurhead peacock cichlid (A. maylandi) is known only from West Reef (Luwala Reef ) and Eccles Reef (Chimwalani Reef ) in the south-eastern part of the lake. The emperor peacock cichlid (A. nyassae) was long known only from a single specimen

collected during the early twentieth century, until rediscovered near Boadzulu Island and in Manzinzi Bay in the south-eastern part of the lake in the 1990s. Hueser’s peacock cichlid (A. hueseri) is known only from near Likoma Island in the eastern part of the lake. The Mdoka peacock cichlid (A. aquilonium) is known only from a single locality off the north-western coast of the lake. Boulenger’s peacock cichlid (A. trematocephala) is known only from a single specimen collected during the early twentieth century from an undefined locality in the northern part of the lake. Several species of sandsifter cichlid (Lethrinops) endemic to Lake Malawi are threatened by overfishing by means of seine nets and bottom trawls. The large-eyed sandsifter cichlid (L. macrophthalmus) is confined to a stretch of the south-western shore of the lake. The small-toothed sandsifter cichlid (L. microdon) is confined to the southern part of the lake. Stride’s cichlid (L. stridei) along with the large-spined sandsifter cichlid (L. macracanthus) and Regan’s sandsifter cichlid (L. micrentodon) are still to be found throughout the lake but have suffered massive declines going back many decades. Four cichlids of the genus Tropheops endemic to Lake Malawi are threatened by sedimentation and collection for the international aquarium trade. The smallmouth cichlid (T. microstoma), Romand’s cichlid (T. romandi), and the red-cheeked cichlid (T. tropheops) are all confined to the area around the Nankumba Peninsula in the southern part of the lake. The modest cichlid (T. modestus) is confined to the Maleri Islands and Chidunga Rocks in the south-western part of the lake. Several cichlids of the genus Pseudotropheus endemic to Lake Malawi are threatened by sedimentation, overfishing, and collection for the international aquarium trade. The Maingano cichlid (P. cyaneorhabdos) is confined to the north-eastern shore of Likoma Island in the eastern part of the lake. The Malawi yellow cichlid (P. flavus) is confined to the shallow shores of Chinyankwazi Island in the south-eastern part of the lake. Joan Johnson’s cichlid (P. joanjohnsonae) is confined to rocky areas around Likoma Island in the eastern part of the lake. The elongated cichlid (P. elongatus) is known only from Mkata Bay, Mbamba Bay, and Hongi Island in the eastern part of the lake. The interrupted cichlid (P. interruptus) is native to Chizumulu Island in eastern part of the lake, but has been introduced by fish exporters to nearby Likoma Island and to Nkhata Bay on the west-central shore. The cyan cichlid (P. cyaneus) is known only from Chinyamwezi Island in the south-eastern part of the lake. The blue-grey cichlid (P. johannii) is confined to a stretch of the eastern shore of the lake. The Chisumulu cichlid (P. tursiops) is known only from Chisumulu Island in the eastern part of the lake. The dark cichlid (P. fuscus) is confined to the north-western shore of the lake. The mottled cichlid (P. galanos) is known only from Mbenji Island in the south-western part of the lake, while the Likoma cichlid (P. perileucos) is native to Likoma Island in the eastern part of the lake but has been introduced to Thumbi West Island and to Otter Point in the south-west.

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The Afrotropical Realm

The lavender cichlid (Iodotropheus sprengerae) is confined to Boadzulu Island, Chinyamwezi Island, Chinyankwazi Island, and Makokola Reef in the south-eastern part of the lake. Stuart Grant’s cichlid (I. stuartgranti) is confined to a small stretch of the south-eastern coast of the lake. Both are threatened by sedimentation and collection for the international aquarium trade. The Nkhotakota cichlid (Protomelas macrodon) is known only from two specimens collected from a single locality on the south-western shore of the lake. The fire-blue cichlid (P. dejunctus) is known only from around Chinyankwazi Island and Chinyamwezi Island, in south-eastern Lake Malawi. The pointed-head cichlid (Trematocranus microstoma) has a relatively wide but highly disjunct distribution within Lake Malawi, where it is threatened by overfishing and collection for the international aquarium trade. The Jafua Bay cichlid (T. pachychilus) is known only from a single locality on the eastcentral shore of the lake. The rubberlip cichlid (Otopharynx pachycheilus) is known only from two disjunct localities in the northern and central parts of the lake. Oliver’s cichlid (O. lithobates) is confined to a few islands in the southern part of the lake. Both are threatened by overfishing, sedimentation, and collection for the international aquarium trade. Several cichlids of the genus Maylandia endemic to Lake Malawi are threatened by sedimentatation and collection for the international aquarium trade. Greshake’s cichlid (M. greshakei) is known only from Makokola Reef, west of Boadzulu Island in the south-eastern part of the lake. Hans Mayland’s cichlid (M. hajomaylandi) is confined to Chizumulu Island in the eastern part of the lake. Lombardo’s cichlid (M. lombardoi) is native to Mbenji Island and Nkhomo Reef in the south-western part of the lake, and has been introduced to Namalenje Island. The redtop cichlid (M. pyrsonotos) is known from a few disjunct localities in the southern part of the lake. The Mazinzi Reef cichlid (M. benetos) is known only from a few localities in the southern part of the lake. The Mpanga Rocks cichlid (M. emmiltos) is confined to a single locality in the north-western part of the lake. The red zebra cichlid (M. estherae) is confined to a few disjunct stretches along the eastern coast of the lake. The red-top cobalt cichlid (M. mbenjii) is confined to Mbenji Island in the south-western part of the lake. The Phaeos cichlid (M. phaeos) is found discontinuously along the central-eastern coast of the lake, while the xanstomachus cichlid (M. xanstomachus) and the golden fleece cichlid (M. chrysomallos) are known only from a few localities in the southern part of the lake. The cyneusmarginata cichlid (M. cyneusmarginata) is known from three localities in the south-western part of the lake and the pure cichlid (M. purum) is known only from an undefined locality (‘Nankhumba Peninsula’) somewhere within Lake Malawi. The Lake Malawi stone cichlid (Petrotilapia chrysos) is known only from the south-eastern shore of the lake, including the waters around the islands of Chinyamwezi and

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Chinyankwazi. Marsh’s stone cichlid (P. nigra) is known from two small, disjunct areas along the southern shore of the lake. The Lake Malawi labeo (Labeo mesops) is endemic to Lake Malawi and the Shire River system, where it is highly threatened by habitat destruction and overfishing. The Lake Malawi yellowfish (Labeobarbus latirostris) is an uncommon species endemic to Lake Malawi, from where it ascends the Bua and Shire rivers to spawn. The Lake Malawi salmon (Opsaridium microlepis) is endemic to Lake Malawi, from where it ascends affluent streams at certain times of the year to spawn. It is threatened by overfishing.

The Upper Nile River Drainage The upper Nile River drainage includes its two major tributaries (the Blue Nile and the White Nile, respectively), from their sources to their junction with the lower Nile at Khartoum in the Sudan. The Abyssinian stone loach (Afronemacheilus abyssinicus) is known from the Blue Nile at its outlet from Lake Tana and from the Baro River. The Blue Nile The Blue Nile (an-nil al-‘Azraqu in Arabic/T’ik’uri Abayi in Amharic) is one of the two major tributaries of the Nile River and the source of most of its water and silt. It originates at Lake Tana in Ethiopia and flows south-east into South Sudan before ultimately joins the White Nile near Khartoum, in Sudan. The Tana barb (Enteromius pleurogramma) is known only from a few specimens collected within the Blue Nile basin, including Lake Tana. Lake Tana (T’ana Hayk’ in Amharic) is located in the Ethiopian Highlands. It is the source of the Blue Nile. The sharp-beaked yellowfish (Labeobarbus acutirostris), Gorguar’s yellowfish (L. gorguari), Dainelli’s yellowfish (L. dainellii), the large-eyed yellowfish (L. macrophtalmus), the flat-backed yellowfish (L. platydorsus), Rüppell’s yellowfish (L. surkis), the long yellowfish (L. longissimus), the Osse yellowfish (L. osseensis), the Gorgor yellowfish (L. gorgorensis), and the thick-barbed yellowfish (L. crassibarbis) are all endemic to Lake Tana and its feeder rivers, where they are threatened by overfishing, habitat destruction and damming projects that obstruct their spawning grounds. The regressive garra (Garra regressus) and the Tana garra (G. tana) are endemic to Lake Tana, where they have been heavily impacted by overfishing. The Lesser Abay River is located in the Choke Mountains of north-western Ethiopia. The Ethiopian amphibious rat (Nilopegamys plumbeus) is a semi-aquatic species known only from a single specimen collected in 1928 from the Lesser Abay River. Most of its habitat has since been destroyed and it is likely extinct, although there have been a few unconfirmed sightings over the years.

The African Region

The White Nile The White Nile (an-nil al-‘abyad in Arabic) is the other of the two main tributaries of the Nile River. It is the longer of the two and arises in the Great Lakes region of central Africa, from where it flows north through Tanzania, Uganda, and South Sudan. The Tonga barb (Enteromius tongaensis) is known only from its original collection in 1935 from a single locality in northern South Sudan. The Tonga labeo (Labeo tongaensis) is known only from its original collection in northern South Sudan in 1935. The White Nile sardine (Neobola nilotica) is known only from its original collection in Sudan in 1919. Keating’s mormyrid (Petrocephalus keatingii) is known only from a few specimens collected from the White Nile between the Sobat confluence and Khartoum in the early twentieth century. The Nile elephant-snout fish (Mormyrus niloticus) is known only from outdated records taken along the entire length of the Nile from the delta to Lake Albert in Uganda, the most recent of which was in 1929. The construction of the Aswan High Dam is believed to have caused the extirpation of this species in North Africa. The white-finned squeaker catfish (Synodontis caudovittatus) is known only from a few specimens collected from Ethiopia, South Sudan, and Sudan during the early twentieth century. It was last recorded in 1907. The Sudd is a vast swamp formed by the White Nile in north-central South Sudan. One of the world’s largest wetlands, it serves as a vitally important refuge for migrating birds. The Nile lechwe (Kobus megaceros) is a type of antelope largely confined to the Sudd swamps along the White Nile and Sobat rivers of South Sudan, with smaller populations in the Machar-Gambella marshes extending (although in very low numbers) into Gambela National Park in south-western Ethiopia. Human displacement and resettlement due to ongoing civil war has resulted in much habitat destruction, hunting pressure, and even large-scale slaughter in most of these areas. The proposed construction of a canal through the species’ core habit would have a devastating effect. The Achwa River (also known as the Aswa River) is located in northern Uganda and (marginally) southern South Sudan. The Achwa squeaker catfish (Synodontis macrops) is confined to the Achwa River.

The Omo River The Omo River is located in south-western Ethiopia. Aramboug’s yellowfish (Labeobarbus arambourgi) is currently known only from a single locality in the Omo River. The Kaffa stone loach (Afronemacheilus kaffa) is confined to the Omo River.

The Shabelle–Jubba River Drainage The Shabelle, Jubba, and Lagh Dera rivers share a common mouth in south-eastern Somalia.

The Shabelle River The Shabelle River (Webi Shabeelle in Somali/Uebi Scebeli in Italian) originates in the highlands of Ethiopia and flows south-east into Somalia, where it becomes seasonal along the coast below Mogadishu. During most years it dries up near the mouth of the Jubba River. Boulenger’s labeo (Labeo boulengeri) is known only from a single locality within the Shabelle River drainage of eastern Ethiopia. The Shabelle suckermouth catfish (Chiloglanis modjensis) is known only from a single locality within the Shabelle River drainage of north-eastern Ethiopia. Lampe’s mountain catfish (Amphilius lampei) is known only from a single, undefined locality within the Shabelle River drainage of central Ethiopia. The Jubba River The Jubba or Juba River (Wabiga Jubba in Somali/Giuba in Italian) is located in southern Somalia. The Jubba yellowfish (Labeobarbus jubae) is confined to the Jubba River. The Jubba catfish (Bagrus urostigma) is confined to the Jubba River. The Lagh Dera River The Lagh Dera River is located in central and north-eastern Kenya and southern Somalia. The Ewaso Nyiro River is located in central Kenya. The Ewaso Nyiro labeo (Labeo percivali) is confined to the Ewaso Nyiro River. The Boji Plains killifish (Nothobranchius bojiensis) is confined to intermittent freshwater marshes within the northern Ewaso Nyiro drainage.

The Athi-Galana River Drainage The Athi-Galana River is the second longest in Kenya. It rises in the Gatamaiyo Forest as the Athi River and runs some 390 km in the south-eastern part of the country before entering the Indian Ocean as the Galana River (also known as the Sabaki River). The Athi labeo (Labeo trigliceps) is confined to the Athi River. The Athi elephant-snout fish (Mormyrus hildebrandti) is a little-known species confined to the Athi River system, including the Tsavo drainage and Mzima Springs. It is threatened by habitat destruction and overfishing.

The Tana River The Tana River is located in central and eastern Kenya. At around 1000 km in length it is the longest in the country, running from the Aberdare Mountains to the Indian Ocean. The Tana delta smooth snake (Meizodon krameri) is known only from two specimens collected in 1934 from the Tana River delta.

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The Afrotropical Realm

The Mnanzini killifish (Nothobranchius willerti) is confined to the lower Tana River drainage, where it inhabits ephemeral water bodies in the floodplain or connected streams located in open as well as forested savanna. The feather-barbelled squeaker catfish (Synodontis manni) is confined to the lower Tana River.

The Pangani River The Pangani River is located in north-eastern Tanzania and adjacent areas of southern Kenya. Bloyet’s cichlid (Astatotilapia bloyeti) is confined to the Pangani River drainage, including Lake Jipe. The Pangani cichlid (Ctenochromis pectoralis) is a poorly known species that appears to have been extirpated from the Pangani River proper and may be extinct. However, isolated populations of similar fish have been reported from springs flanking Mount Kilimanjaro, the Chemka springs upstream of the Pangani and the Mzima springs in the Tsavo River drainage of Kenya. The red Pangani barb (Enteromius venustus) and the fourspotted barb (E. quadripunctatus) are both endemic to the Pangani River drainage. Lake Jipe Lake Jipe is located on the Kenya/Tanzania border, within the Pangani River system. The Jipe tilapia (Oreochromis jipe) is a type of cichlid historically endemic to Lake Jipe, where it is highly threatened by overfishing. In recent areas it has managed to extend its range to adjacent tributaries.

The Malagarasi River The Malagarasi River is located in north-western Tanzania and south-eastern Burundi. Devos’ cichlid (Neolamprologus devosi) is known only from a few localities within the Malagarasi River. The Malagarasi largemouth cichlid (Serranochromis janus) is confined to the Malagarasi River. The Uvinza tilapia (Orthochromis uvinzae) is known only from a single locality within the middle Malagarasi River drainage in Tanzania. The Mazimero tilapia (O. mazimeroensis) is known only from the Mazimero and Nanganga rivers within the upper Malagarasi River drainage of Burundi. The Moso tilapia (O. mosoensis) is known only from the upper Malagarasi River drainage. The Malagarasi tilapia (O. malagaraziensis) is confined to the Malagarasi River, where it is threatened by habitat destruction. The Karomo tilapia (Oreochromis karomo) is confined to the lower Malagarasi River and its delta, where it is seriously threatened by overfishing and habitat destruction. The Malagarasi barb (Enteromius quadralineatus) is known only from the Malagarasi River. The Malagarasi salmon (Opsaridium splendens) is confined to the Malagarasi River drainage, where it is threatened by habitat destruction and sedimentation.

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The Mutsindozi snake catfish (Clariallabes mutsindoziensis) is confined to swampy areas within the Malagarasi River. The Ruchugi River The Ruchugi River is located in western Tanzania. The Ruchugi tilapia (Orthochromis kasuluensis) is confined to the upper Ruchugi drainage. The Ugalla River The Ugalla River is located in western Tanzania. The Majamazi tilapia (Orthochromis rubrolabialis) is known only from the Majamazi River, a tributary of the Ugalla River.

The Wami River The Wami River is located in eastern Tanzania. The Wami barb (Enteromius laticeps) is known only from two localities within the Wami River. Steinfort’s killifish (Nothobranchius steinforti) is known only from two intermittent pools within the upper Wami River drainage. Possibly extinct in the wild, it has fortunately been established in captivity by aquarium hobbyists. The Wami upside-down catfish (Atopochilus vogti) is known only from a single specimen collected from the Wami River in the early twentieth century. The Wami loach catfish (Zaireichthys wamiensis) is known only from a single locality within the Wami River.

The Ruvu River The Ruvu River is located in eastern Tanzania. It originates in the southern Uluguru Mountains and flows eastwards to the Indian Ocean. Lourens’ killifish (Nothobranchius lourensi), Foersch’s killifish (N. foerschi), and the Kisaki killifish (N. flammicomantis) are all confined to temporary pools and marshes within the Ruvu River floodplain.

The Rufiji River The Rufiji River is located in central, southern, and eastern Tanzania. Peters’ disichodus (Distichodus petersii) is confined to the Ruaha and Rufiji river drainages of central and eastern Tanzania, where it is threatened by overfishing and pollution. The Great Ruaha River The Great Ruaha River is located in south-central and southwestern Tanzania, where it flows through the Usangu wetlands and Ruaha National Park east into the Rufiji River. The Ruaha shellear (Kneria ruaha) is confined to a few tributaries of the Great Ruaha River on the north-western slopes of the Southern Highlands in south-western Tanzania. The Kimami shellear (Parakneria tanzaniae) is known only from two tributaries of the upper Great Ruaha River. The Little Ruaha River is located in south-western Tanzania.

The African Region

The Ruaha lampeye (Aplocheilichthys omoculatus) and the Usangu lampeye (A. usanguensis) are both confined to the Little Ruaha River. The Kilombero River The Kilombero River (also known as the Ulanga River) is located in western Tanzania. It originates on the eastern slopes of the East African Rift and flows north-east into the Rufiji River. The Kilombero killifish (Nothobranchius kilomberoensis) and the Gemini killifish (N. geminus) are both confined to temporary pools and marshes within the Kilombero River floodplain.

The Zambezi River Drainage At 2574 km the Zambezi is the fourth longest river in Africa as well as its longest east-flowing. It rises in the highlands of Zambia and flows first south through eastern Angola before winding its way along the border between Zambia and Zimbabwe to Mozambique, where it eventually empties into the Indian Ocean. Its most notable physical feature is the spectacular Victoria Falls on the Zambia/Zimbabwe border. The Kariba tilapia (Oreochromis mortimeri) is confined to Lake Kariba and the middle Zambezi River drainage, where it is seriously threatened by competition with introduced Nile tilapia (O. niloticus). The gorgeous barbel (Cyplobarbus bellcrossi) is known with certainly only from a single locality within the upper Zambezi drainage, all other records being highly questionable. The Longa River The Longa River is a tributary of the upper Zambezi located in east-central Angola. Rousselle’s barb (Enteromius roussellei) is known only from the Longa River. The ghost mormyrid (Paramormyrops jacksoni) is known only from a single specimen collected in the Longa River. The Kataba River The Kataba River is a tributary of the upper Zambezi located in south-western Zambia. The banded tetra (Neolebias lozii) is confined to the Kataba River and its tributary stream, the Siandra. The Kafue River The Kafue River is located in central Zambia (Southern, Central, and Lusaka provinces). The Kafue Flats are discussed under the flooded grasslands sections. The Lufupa yellowfish (Labeobarbus lufupensis) is confined to the Lufupa River, a tributary of the Kafue River.

The Okavango River The Okavango River (Rio Cubango in Portuguese) is located in Angola, Botswana, and Namibia. It is notable for not having an outlet to the sea: instead, it discharges into the Okavango delta within an endorheic basin of the Kalahari Desert.

The Cutato River The Cutato River is located in west-central Angola. The Cutato shellear (Parakneria fortuita) is confined to the Cutato River.

The Komati River The Komati River is located in coastal southern Mozambique and north-eastern South Africa (Limpopo and Mpumalanga) and Swaziland. The orange-fringed cichlid (Chetia brevis) was historically endemic to the Komati River drainage, where it is threatened by habitat destruction, water extraction, and introduced species. However, it was successfully translocated into a few impoundments within Kruger National Park in 1975.

The Limpopo River The Limpopo River is located in north-eastern South Africa (Mpumalanga and Limpopo), Swaziland, and southern Mozambique (Gaza province). The Blyde River The Blyde River is located in north-eastern South Africa (Mpumalanga province). The Treur barb (Enteromius treurensis) is confined to a 4.5-km stretch of the upper reaches of the Blyde River. It was historically also found in the Treur River, a tributary of the Blyde River, but was extirpated there by the introduction of exotic fish species in the 1960s. The remaining population in the Blyde River is protected by a downstream waterfall, but remains vulnerable.

The Olifants River The Olifants River (Olifantsrivier in Afrikaans) is located in south-western South Africa (Western Cape and Northern Cape). The Clanwilliam yellowfish (Labeobarbus capensis) is confined to the Olifants River drainage. The fiery redfin (Pseudobarbus phlegethon) is confined to a few tributaries within the Olifants River drainage. The Clanwilliam labeo (Labeo seeberi) was historically widespread within the Olifants River drainage, but has been much-reduced due to the introduction of predatory fish species. Barnard’s rock catfish (Austroglanis barnardi) is confined to three small tributaries of the Olifants River in Western Cape province, where it is threatened by water extraction and introduced fish species. The Twee River The Twee River is located in western Western Cape province. The Twee redfin (Pseudobarbus erubescens) is confined to the Twee River and its tributaries, where it is seriously threatened by habitat degradation and introduced fish species.

Miscellaneous Lakes, Rivers, and Marshes Lake Chad is located on the edge of the Sahara in Cameroon, Chad, Niger, and Nigeria. A historically large, shallow,

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The Afrotropical Realm

endorheic lake, it has varied greatly in size over the centuries. The Bahr Sara trout barb (Raiamas shariensis) is a littleknown and perhaps doubtful species confined to the Lake Chad basin. Lake Abaeded is a saline lake located in Eritrea. The Abaeded cichlid (Danakilia dinicolai) is confined to Lake Abaeded. Lake Ziway is a freshwater lake located in the Rift Valley of central Ethiopia. The Ziway yellowfish (Labeobarbus ethiopicus) is confined to the deeper waters of Lake Ziway. Lake Abaya is a large lake located in the Rift Valley of southern Ethiopia. Brunelli’s labeo (Labeo brunellii) is currently known only from Lake Abaya. Lake Turkana (formerly known as Lake Rudolf ) is located in the Rift Valley of Kenya and Ethiopia. It is the world’s largest permanent desert lake and largest alkaline lake, and supports a rich fauna. An active volcano forms an island in the lake’s centre. Lake Turkana formerly contained Africa’s largest population of Nile crocodiles (Crocodylus niloticus), which remain relatively common in some areas. The Turkana hinged terrapin (Pelusios broadleyi) is confined to Lake Turkana. Lake Chala is a small crater lake located on the eastern slopes of Mount Kilimanjaro, on the border between Kenya and Tanzania. The Chala tilapia (Oreochromis hunteri) is a type of cichlid confined to Lake Chala, where it is seriously threatened by siltation and introduced species. Lake Chungruru is a small, isolated, endorheic crater lake located in southern Tanzania (Mbeya Region). The Chungruru tilapia (Oreochromis chungruruensis) is a type of cichlid confined to Lake Chungruru, where it is seriously threatened by overfishing and introduced species. Lake Kyoga is a large, notably shallow lake located in western Uganda. The shoreline is quite swampy and much of its surface is covered by water lilies. Three cichlids of the genus Haplochromis are endemic to the Lake Kyoga basin. Worthington’s cichlid (H. worthingtoni) is confined to Lake Kyoga, while the broad-banded cichlid (H. latifasciatus) and the Kyoga cichlid (H. orthostoma) are found in Lake Kyoga and two smaller satellites (Lake Bisina and Lake Nawampasa). Lake Kijanebalola is located in southern Uganda. The Kijanebalola cichlid (Haplochromis exspectatus) is confined to Lake Kijanebalola. The Luiche River is located in western Tanzania. The Luiche tilapia (Orthochromis luichensis) is a type of cichlid confined to a few small effluents of the Luiche River. The Rufugu River is located in western Tanzania. The Rufugu tilapia (Orthochromis rugufuensis) is a type of cichlid known only from the upper Rufugu River drainage. The Kalambo River is located on the Tanzania–Zambia border. The Kalambo suckermouth catfish (Chiloglanis kalambo) is confined to the Kalambo River drainage. The Buzi River (Rio Búzi in Portuguese) is located in south-central Mozambique. The Buzi catfish (Chrysichthys

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hildae) is known only from a few specimens collected in the lower Buzi River. The Pungwe River is located in eastern Zimbabwe and western Mozambique. The Pungwe barbel (Barbus hondeensis) is known only from a single specimen collected from the upper Pungwe River. The Cunene River is located in south-western Angola and north-western Namibia. The Cunene citharine (Nannocharax fasciolaris) is known only from its original collection during the 1860s from the Cunene River drainage. The Berg River (Bergrivier in Afrikaans) is located in southwestern South Africa (Western Cape). The Berg redfin (Pseudobarbus burgi) is confined to the Berg River and its tributaries, where it is threatened by habitat destruction and introduced fish species. The Breede River (Breedeivier in Afrikaans) is located in south-western South Africa (Western Cape). Burchell’s redfin (Pseudobarbus burchelli) is confined to the Breede River drainage, where it is threatened by habitat destruction and introduced fish species.

Coasts and Satellite Islands This section is comprised of a small stretch of the West African coast along with the adjacent Cape Verde Islands, as well as all of the remainder of the African coastline and satellite islands from Angola to the Red Sea. The isolated South Atlantic islands of Ascension, Saint Helena, and Tristan da Cunha are also included here. The grey-headed thicket rat (Grammomys caniceps) is confined to dry coastal shrubland in eastern Kenya and south-eastern Somalia. Two species of golden mole (Cryptochloris) endemic to coastal dunes in western South Africa are seriously threatened by diamond mining. De Winton’s golden mole (C. wintoni) was historically known only from a small area of Northern Cape province. Not recorded for many decades, it is possibly extinct. Van Zyl’s golden mole (C. zyli) is known only from two localities in Western Cape and Northern Cape. The African penguin (Spheniscus demersus) breeds in scattered coastal colonies in Namibia and South Africa, with vagrants recorded as far north as Gabon and central Mozambique. In recent decades it has undergone a significant decline, most likely as a result of commercial fisheries and shifts in prey populations. The northern rockhopper penguin (Eudyptes moseleyi) breeds primarily in the Tristan da Cunha Archipelago in the South Atlantic, with the remainder being found on the Amsterdam and Saint Paul Islands in the southern Indian Ocean. The species has been declining for decades for reasons that are poorly known, although changes in sea-surface temperature, pollution, unsustainable levels of harvesting, fisheries by-catch, and introduced predators have all been implicated.

The African Region

The Cape cormorant (Phalacrocorax capensis) occurs along the southern African coast from Angola to Mozambique, although breeding only takes place within a much smaller stretch. Several of its key nesting colonies in South Africa and Namibia suffered serious declines due to collapsing fish stocks, oil pollution, human disturbance, and disease. The Bank cormorant (P. neglectus) is confined to Namibia and western South Africa, where its colonies face similar threats. The Cape gannet (Morus capensis) is a large seabird that ranges, during the non-breeding season, along the entire coast of Africa from the Gulf of Guinea to Kenya, but currently nests on on six small islands off Namibia and South Africa. Historically heavily hunted for food, today the main threats are collapsing fish stocks, storms and oil spills. The Damara tern (Sternula balaenarum) is a type of small seabird that breeds along the southern and south-western coast of Africa, from where it migrates north as far as Liberia. Its primary nesting colonies in Namibia are under threat by offroad vehicles, diamond mining, and residential development. Ash’s lark (Mirafra ashi) is known only from a single small stretch of coast in south-eastern Somalia, where it is threatened by development. The Obbia lark (Spizocorys obbiensis) is confined to a narrow coastal strip in south-eastern Somalia. The salt marsh leaf-toed gecko (Cryptactites peringueyi) is restricted to two subpopulations in coastal south-eastern South Africa (Eastern Cape).

Jozani-Chwaka Bay National Park, as well as in the adjacent agricultural areas to the south and the coral thickets and mangrove swamps of Uzi Island some 10 km to the southwest. Small numbers are also found along Zanzibar’s eastern coast and in some mangrove swamps in the west (including the uninhabited islet of Vundwe), with a small, translocated population in the Masingini Forest Reserve. The species was also introduced to the Ngezi Forest Reserve on Pemba Island in 1974, where it still marginally persists. Despite long being the focus of national as well as international conservation efforts it continues to be threatened by loss of habitat and degradation and, to a lesser extent, poaching for meat, persecution, road kills, and capture for use as pets. All told the population is thought to number less than 2000. The Zanzibar dwarf galago (Galagoides zanzibaricus zanzibaricus) historically occurred throughout Zanzibar but is now largely restricted to a few areas in the central and southern part of the island. The Zanzibar servaline genet (Genetta servalina archeri) first became known to science in 1995 when a specimen was killed near the Jozani Forest. It has since been recorded by camera traps within Jozani-Chwaka Bay National Park. The Zanzibar blind snake (Letheobia pallida) is a littleknown species confined to Zanzibar. The punctulated reed frog (Hyperolius puncticulatus) is confined to a few localities in the western part of the island. The Zanzibar running frog (Kassina jozani) is known only from the Jozani Forest and its surroundings on Zanzibar.

The Zanzibar Archipelago

Pemba Island A mainly low-lying island, Pemba is separated from both Zanzibar and the African mainland by deep channels. The Pemba flying fox (Pteropus voeltzkowi) had, by the early 1990s, been reduced to no more than a few hundred individuals, although numbers have increased in recent years. The Pemba scops owl (Otus pembaensis) is largely confined to two small areas of forest (Ngezi and Msitu Mkuu). The Pemba green pigeon (Treron pembaensis) is confined to Pemba and adjoining coral islets. The Pemba wolf snake (Lycophidion pembanum) is confined to forests and plantations on Pemba Island. The Pemba gracile blind snake (Letheobia pembana) is known only from a single specimen collected in 1905. The Ngezi reed frog (Hyperolius watsonae) is confined to the Ngezi Forest in the northern part of the island. Pakenham’s puddle frog (Phrynobatrachus pakenhami) is confined to northern Pemba Island.

The Zanzibar Archipelago consists of several islands lying off the coast of north-eastern Tanzania. There are four main islands, three primary islands with human populations and a fourth coral island that serves as an essential breeding ground for seabirds, plus a number of smaller islets. The Mrora toad (Mertensophryne howelli) is confined to Zanzibar and Mafia Island. Zanzibar Zanzibar (formally known as Unguja) is the largest and most populated island in the group and has a fauna reflecting its connection to the African mainland during the last Ice Age. Hilly and at one time heavily forested, most of the original forest cover has been replaced by plantations and the wildlife heavily hunted and trapped. Many species have been largely or wholly exterminated as a result, most notably the local population of African leopard (Panthera pardus). The Zanzibar red colobus (Piliocolobus kirkii) is a strikingly attractive species of monkey that was apparently already rare and confined to the island in 1868 when it was first investigated by its discoverer, Sir John Kirk. Historically it may have occurred on the East African mainland, but has long been extirpated there. On Zanzibar it lives mainly in the low, thin coastal scrub of the south-eastern part of the island in

The Mafia Archipelago The Mafia Archipelago consists of one large main island and several smaller ones, some of which are uninhabited. The Mafia killifish (Nothobranchius korthausae) is confined to a few swamps, pools, ditches, and small streams on Mafia Island.

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The Cape Verde Islands The Cape Verde Islands (Cabo Verde in Portuguese) is a horseshoe-shaped cluster of 10 volcanic islands and 8 islets approximately 570 km off the coast of West Africa, divided spatially into two island groups. All are arid for the most part with vegetation consisting mainly of tropical dry forest and shrubland, although in certain more humid mountainous areas a unique type of cloud forest, known as laurel forest, has evolved. Through sheer isolation the islands have developed a number of endemic vertebrates, many of which are threatened by human activities and the introduction of goats, monkeys, and mongooses. The Cape Verde warbler (Acrocephalus brevipennis) was long thought to be restricted to the island of Santiago, but in 1998 it was discovered in small numbers on São Nicolau. In 2004 another population was discovered on Fogo. It is threatened by habitat destruction and drought. The Cape Verde house gecko (Hemidactylus bouvieri) is endemic to the archipelago, where two subspecies are highly threatened. The nominate form (H. b. bouvieri) occurs on São Vincente, Santo Antão, and São Nicolau, while the Raso house gecko (H. b. razonensis) is known only from Santo Luzia and Raso, having been extirpated from Santiago and Brava. Vaillant’s mabuya (Chioninia vaillantii) is a type of lizard that occurs on Fogo and Santiago Islands and on the islet Ilhéu de Cima (one of the Rombos Islets). Subfossil evidence suggests that it was formerly found on Maio and Boa Vista as well, but the species is now absent from these islands. The Barlavento Islands The Barlavento Islands are the northern group of the Cape Verde Islands.

The Cape Verde giant skink (Chioninia coctei) is only known for certain from a few specimens collected on the islands of Branco and Raso, but subfossil materials indicate that it may have once been more widespread in the Barlavento Islands. In any case the species was last observed in 1940 and appears to be extinct, a victim of overhunting and drought. Raso (Ilhéu Raso in Portuguese) is a small, uninhabited volcanic islet. The Raso lark (Alauda razae) is confined to Raso, where the total population is stable at around 100. However, even a slight habitat change may prove fatal to such a small and specialized population. The giant wall gecko (Tarentola gigas) is confined to the islands of Raso and Branco, where it is threatened by loss of habitat and introduced species. The Raso giant wall gecko (T. g. gigas) is confined to a few areas on Raso. Branco (Ilhéu Branco in Portuguese) is another small, uninhabited islet located west of Raso. The Branco giant wall gecko (Tarentola gigas brancoensis) is confined to Branco. Boa Vista is the easternmost of the Cape Verde Islands. The Boa Vista wall gecko (Tarentola boavistensis) is confined to Boa Vista Island and Sal Rei islet. The Sotavento Islands The Sotavento Islands are the southern group of the Cape Verde Islands. Fogo (Ilha de Fogo in Portuguese) reaches the highest altitude of all the Cape Verde islands, rising to 2829 m at the summit of its active volcanco, Pico do Fogo. López-Jurado’s house gecko (Hemidactylus lopezjuradoi) was first described in 2008 from Fogo. Only a few specimens have been collected.

Figure 3.10 Illustration of a Cape Verde giant skink from 1885. (Credit: J. Terrier.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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The African Region

Santiago (Ilha de Santiago in Portuguese) is the largest of the Cape Verde Islands. The Santiago wall gecko (Tarentola rudis) is confined to the southern part of Santiago Island and on Santa Maria islet.

The Saint Helena plover (Charadrius sanctaehelenae) is the island’s only surviving endemic land bird. The species is mainly found on pastureland and seems to fluctuate in population over time, with the most recent estimate being around 500.

Ascension

Tristan da Cunha

Ascension is a tropical volcanic island rising out of the deep waters of the South Atlantic Ocean, midway between the horn of South America and Africa. Rats probably came ashore soon after the island’s discovery in 1501 and cats were introduced after 1815. By 1843 the island was barren with few plants, and today introduced plants and trees cover most of the slopes. In recent years the UK government has taken steps to restore the island, and in 2016 announced that the area around it was to become a huge marine reserve. The Ascension flightless crake (Mundia elpenor) was a type of flightless rail known only from subfossil bones and a brief account made by merchant and traveller Peter Mundy in 1656. It is thought to have become extinct after rats were introduced to the island in the eighteenth century, but it may have survived until the arrival of feral cats in 1815. The Ascension frigatebird (Fregata aquila) is a large seabird confined to Ascension and the seas that surround it. In the early nineteenth century it was very numerous on the island but was extirpated there by introduced cats, thereafter breeding only on Boatswainbird Islet, a flat-topped, steep-sided rock 250 m off the north-eastern coast. With the successful eradication of cats on Ascension in 2006, however, the birds have once again started to recolonize.

Tristan da Cunha and its smaller satellite islands of Inaccessible, Nightingale, and Gough lie roughly 2400 km west of the tip of South Africa. The vegetation has suffered from the heavy gales that sweep the islands and, of course, from humans. In sheltered places the growth is luxuriant but mostly stunted. The only woody tree is a buckthorn (Phylica arborea) and reed-like tussock grass (Spartina arundinacea) dominates the vegetation. Introduced pigs, goats, rabbits, and cats have long lived in a feral state, but these remote, volcanic islands nevertheless remain vitally important for seabirds in particular. Inaccessible and Nightingale have been uninhabited since the 1960s, with the former as well as Gough Island having since been set aside as wildlife reserves. The Tristan albatross (Diomedea dabbenena) breeds only on Gough and Inaccessible, having become extirpated on the main island of Tristan da Cunha. The Atlantic yellow-nosed albatross (Thalassarche chlororhynchos) breeds only on Inaccessible Island, Middle Island, Nightingale Island, Stoltenhoff Island, and Gough Island, from where it disperses at other times throughout the South Atlantic as far as south-western Africa and south-eastern South America. It is threatened by introduced species within their nesting colonies and by fisheries by-catch. The Atlantic petrel (Pterodroma incerta) breeds only on Gough Island and possibly Tristan da Cunha.

Saint Helena Saint Helena is another remote, tropical volcanic island in the South Atlantic Ocean. When first discovered in 1502 by the Portuguese it was uninhabited and entirely covered in dense tropical forest, with even the tremendous cliffs that overhang the sea colonized by trees. Today nothing of these forests remains. Goats had been introduced by 1513, and in 1588 a visitor reported there were flocks ‘a mile long’. Since then cattle, dogs, cats, donkeys, mice, rats, and sheep were introduced as well. Fire also destroyed large forests. Moreover, humans introduced a great number of exotic plants, in particular New Zealand flax, and these now outnumber the native ones about 10–1. At least two endemic species of tree have vanished along with many of the native flowering plants, and much of the island is barren. The native birds have also suffered, with a number of species having gone extinct since 1502. Among these are the Saint Helena flightless dove (Dysmoropelia dekarchiskos), Saint Helena flightless hoopoe (Upupa antaios), Saint Helena gadfly petrel (Pterodroma rupinarum), Saint Helena Bulwer’s petrel (Bulweria bifax), Saint Helena flightless crake (Atlantisia podarces), Saint Helena rail (Zapornia astrictocarpus), and Saint Helena cuckoo (Nannococcyx psix), all of which are known only from fossils.

Tristan da Cunha Tristan da Cunha is the main island in the archipelago. It features an active volcano rising to 2060 m. The Tristan moorhen (Gallinula nesiotis) was a flightless bird that was wiped out, presumably by rats and pigs, about 1872. Three subspecies of Tristan thrush (Turdus eremita) are to be found on each of the main islands within Tristan da Cunha. The nominate form (T. e. eremita) is confined to Tristan da Cunha itself, where the population is estimated at around ‘several hundred’. Inaccessible Island Inaccessible Island is located 45 km south-west of Tristan da Cunha. It is essentially a large, extinct volcano fringed with sheer sea cliffs and with only a few boulder beaches. Sailors have always been wary of its difficult landings and inhospitable terrain, and indeed the island has been without permanent inhabitants since 1873. The spectacled petrel (Procellaria conspicillata) breeds only the high western plateau of Inaccessible Island, although at other times ranges throughout the South Atlantic Ocean.

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The Afrotropical Realm Figure 3.11 An 1861 illustration of a Tristan moorhen. (Credit: J. Jury.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The Inaccessible rail (Atlantisia rogersi) is the world’s smallest flightless bird. It is endemic to Inaccessible where it is considered stable, thanks mainly to the fact that the island remains free from rats. Should they ever come ashore, however, the effect on this ground-nesting species would be catastrophic. The Inacessible thrush (Turdus eremita gordoni) is confined to the island. Three subspecies of Inaccessible finch (Nesospiza acunhae) are endemic to the island. The nominate form (N. a. acunhae) is found along the coastline, Dunne’s Inaccessible finch (N. a. dunnei) along the eastern edge of the island’s plateau and coastline, and Fraser’s Inaccessible finch (N. a. fraseri) atop the island’s plateau at 300–600 m. Wilkins’ finch (N. wilkinsi) is confined to Inaccessible and Nightingale, each with its own distinct subspecies. The Inaccessible Wilkins’ finch (N. w. dunnei) is confined to the island, where the total population is between 40 and 90 individuals. Nightingale Island Nightingale Island is an active volcanic island just 3 km2 in area. It is the main island in a small group that also includes Middle and Stoltenhoff islands. The Nightingale thrush (Turdus eremita procax) is confined to Nightingale, Middle, and Stoltenhoff Islands. The Nightingale finch (Nesospiza questi) and the Nightingale Wilkins’ finch (N. wilkinsi wilkinsi) are both confined to the island.

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Gough Island Gough Island is a rugged, rectangular-shaped volcanic island about 400 km south-east of the main Tristan da Cunha Archipelago. Surrounded by a number of smaller satellite islands and rocks, it is largely uninhabited apart from the crew of a weather station and is one of the most remote places on Earth. MacGillivray’s prion (Pachyptila macgillivrayi) is a type of seabird confined, as far as known, to Gough Island, although it may ultimately prove to be conspecific with the now-extirpated population from Amsterdam Island and an existing one on St. Paul, where a few hundred are confined to a single rat-free island. While still common on Gough, it remains vulnerable. The Gough moorhen (Gallinula comeri) was historically endemic to Gough, although in 1956 eight were released on Tristan da Cunha, which they subsequently managed to colonize (thereby replacing the now-extinct Tristan moorhen, G. nesiotis). The Gough finch (Rowettia goughensis) is confined to Gough Island.

Balance for the African Region After the evolution of modern humans (Homo sapiens) within Africa approximately 350,000–260,000 years ago the continent was mainly populated by groups of hunter-gatherers, some of whom migrated out around 50,000 years ago and proceeded to populate the rest of the globe. Other migrations within the African continent have been dated to that time, with evidence

The African Region

of early human settlement having been found in Southern Africa, North Africa, and in the Sahara. The domestication of cattle in Africa preceded agriculture. Pre-colonial Africa possessed perhaps as many as 10,000 different states and polities, ranging from small family groups of hunter-gathers such as the San people of southern Africa; larger, more structured groups such as the family clans of the Bantu of central, southern, and eastern Africa; heavily structured clans in the Horn of Africa; the large Sahelian kingdoms; and autonomous citystates and kingdoms of West Africa and the coastal trading towns of south-east Africa. Early European exploration of the African Region was dominated by the Portuguese. During the mid-fifteenth century they had begun to sail ever further along the North and West African coast, in the process discovering the Cape Verde Islands. In 1488 Bartolomeu Dias rounded what is now the Cape of Good Hope, at the southernmost tip of the continent, thereby opening up the important European sea route to India and the Far East. In the early 1490s the Portugese explorer Pêro da Covilhã visited the East African coast (what is now Kenya, Zanzibar, and Ethiopia). The remote South Atlantic island of Tristan da Cunha was first discovered in 1506 by the Portugese explorer Tristão da Cunha. In 1514–15 Antonio Fernandes explored inland, reaching present-day Zimbabwe. In 1618 the Spanish missionary Pedro Páez is believed to have been the first European to see and describe the source of the Blue Nile in Ethiopia. In 1826 the Scottish explorer Alexander Gordon Laing became the first European to reach the fabled city of Timbuktu, in what is now Mali. Sadly, he would be murdered upon leaving the city, but two years later the French explorer René Caillié succeeded in returning alive. In 1846 Candido José da Costa Cardoso discovered Lake Malawi, while Rodrigues Graça travelled from Angola to south-western Katanga, in what is now the Democratic Republic of Congo. Around 1847–48 António da Silva Porto reached the upper Zambezi River. In 1848 the German missionary Johannes Rebmann became the first European to sight Mount Kilimanjaro. In 1849 David Livingstone and William Cotton Oswell crossed the Kalahari Desert to Lake Ngami, in what is now Botswana. A few years later, between 1853 and 1856, Livingstone became the first to traverse Africa from west to east, travelling from Luanda in Angola to Quelimane in Mozambique. He also explored much of the upper Zambezi, and discovered and named Victoria Falls. In 1858 Richard Francis Burton and John Hanning Speke discovered Lake Tanganyika and Lake Victoria. In 1862 Speke would discover the source of the Nile at the northern end of Lake Victoria. In 1864 the English explorer Samuel Baker discovered Lake Albert where, in the distance, his sighted the Mountains of the Moon (the Ruwenzori Mountains). Between 1875 and 1877 Henry Morton Stanley circumnavigated Lake Tanganyika and

Lake Victoria, and sighted Lake George. Overall, however, European exploration of the African interior remained limited prior to the twentieth century, the latter being content merely to establish sporadic trading posts along the coast while they were actively discovering and colonizing the New World. Indeed, tropical Africa was one of the last areas of the world to be colonized and influenced by Europeans. Throughout this period human population remained sparse, with maps showing large areas of the continent labelled ‘unknown’. Slavery had long been practiced in Africa, at first exclusively by Arabs and Africans. Between the fifteenth and nineteenth centuries the Atlantic slave trade took an estimated 7–12 million West Africans to the New World. In addition, more than a million Europeans were captured by Barbary pirates and sold as slaves in North Africa during this period. The decline of the slave trade during the nineteenth century was actually hampered by African leaders themselves. Between 1808 and 1860 the British West Africa Squadron seized approximately 1600 slave ships and freed some 150,000 Africans. The history of European colonization of the African continent was driven by Britain, France, Germany, Portugal, Spain, the Netherlands, and Italy, and to this day nearly all countries speak the language imposed by their particular colonial power. By 1914 almost 90 per cent of Africa was under European control, with only Abyssinia (Ethiopia) and Liberia remaining independent, although the former would later be invaded in 1936. There was a gradual decolonization after World War II, and since that time these states have been hampered by political instability, corruption, violence, and authoritarianism. Border and territorial disputes have been commonplace, with the European-imposed boundaries of many nations having been widely contested by means of armed conflict. More critically, population has increased well beyond the carrying capacity of the continent, and continues to grow. As a result there has been an extraordinary increase in human population and a general devastation of the environment. In recent historical time (i.e. since A . D . 1500), the African Region has lost at least 56 species/5 subspecies of vertebrates. Among the extinct forms 1 species/5 subspecies are mammals, 9 species are birds, 3 species are reptiles, and 43 species are freshwater fishes. Another 12 species are possibly extinct. In addition, there are 808 species/68 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 113 species/50 subspecies are mammals, 115 species/16 subspecies are birds, 107 species/2 subspecies are reptiles, 138 species are amphibians, and 335 species are freshwater fishes.

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Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

1 species

3 species

~ species

113 species

5 subspecies

~ subspecies

~ subspecies

50 subspecies

6 taxa

3 taxa

~ taxa

163 taxa

9 species

2 species

~ species

115 species

~ subspecies

~ subspecies

~ subspecies

16 subspecies

9 taxa

2 taxa

~ taxa

131 taxa

3 species

2 species

~ species

107 species

~ subspecies

~ subspecies

~ subspecies

2 subspecies

3 taxa

2 taxa

~ taxa

109 taxa

~ species

4 species

~ species

138 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

4 taxa

~ taxa

138 taxa

43 species

1 species

~ species

335 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

43 taxa

1 taxon

~ taxa

335 taxa

56 species

12 species

~ species

808 species

5 subspecies

~ subspecies

~ subspecies

68 subspecies

61 taxa

12 taxa

~ taxa

876 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

Balance for the Afrotropical Realm In recent historical time (i.e. since A . D . 1500), the Afrotropical Realm as a whole has lost at least 56 species/6 subspecies of vertebrates. Among the extinct forms 1 species/6 subspecies are mammals, 9 species are birds, 3 species are reptiles, and 43 species are freshwater fishes. Another 15 species are possibly extinct, and 1 species is currently extinct in the wild. In addition, there are 1773 species/121 subspecies currently threatened with extinction (that is to say, either Critically

Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 260 species/79 subspecies are mammals, 182 species/24 subspecies are birds, 157 species/5 subspecies are reptiles, 281 species are amphibians, and 893 species/13 subspecies are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

1 species

4 species

~ species

260 species

6 subspecies

~ subspecies

~ subspecies

79 subspecies

7 taxa

4 taxa

~ taxa

339 taxa

9 species

3 species

~ species

182 species

~ subspecies

~ subspecies

~ subspecies

24 subspecies

9 taxa

3 taxa

~ taxa

206 taxa

3 species

3 species

~ species

157 species

~ subspecies

~ subspecies

~ subspecies

5 subspecies

3 taxa

3 taxa

~ taxa

162 taxa

Birds

Reptiles

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(cont.)

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Amphibians

~ species

4 species

~ species

281 species

~ subspecies

~ subspecies

~ subspecies

~ taxa

4 taxa

~ taxa

281 taxa

43 species

1 species

1 species

893 species

~ subspecies

~ subspecies

~ subspecies

13 subspecies

43 taxa

1 taxon

1 taxon

906 taxa

56 species

15 species

1 species

1773 species

6 subspecies

~ subspecies

~ subspecies

121 subspecies

62 taxa

15 taxa

1 taxon

1894 taxa

Freshwater fishes

Total vertebrates

~ subspecies

Note: ~, not applicable.

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Chapter

4

The Madagascan Realm

The Madagascan Realm includes Madagascar along with the Mascarene, Comoros, and Seychelles island groups of the south-western Indian Ocean and a few smaller islands. Biogeographically speaking it has long been included with Africa, although the growing weight of evidence argues for separation. Most are essentially old pieces of the ancient supercontinent of Gondwana that broke away from Africa millions of years ago, although some (such as the Comoros and Mascarenes) are volcanic islands that formed much more recently. The lesser yellow bat (Scotophilus borbonicus) was considered common on the island of Réunion in the Mascarenes during the early nineteenth century but soon disappeared. A single specimen was collected from southern Madagascar in 1868 (former Toliara province), but the species has not been recorded since. It is most likely extinct, a victim of habitat destruction. The Madagascar harrier (Circus macrosceles) is a bird of prey found across the whole of Madagascar and on Grande Comore, Mohéli, and Anjouan in the Comoro Islands, where it is primarily associated with wetlands. Despite its wide distribution the species occurs at extremely low densities, with the total population estimated at less than 250. The Madagascar sacred ibis (Threskiornis bernieri) is confined to coastal areas along the length of western Madagascar as well as in the Seychelles, where the total population is thought to be between 2000 and 3000. Humblot’s heron (Ardea humbloti) is largely restricted to western coastal Madagascar but is also recorded as a vagrant from areas further east (i.e. the Central Highlands, especially Lake Alaotra), the Anorontany Archipelago, and the Comoros Islands. The total population is thought to be no more than 1500. The Madagascar pond heron (A. idae) is widespread in freshwater wetland areas across east-central Africa, Madagascar, and the Seychelles, Comoros, and Mascarene Islands. It is everywhere rare, however, with a total population estimated at between 2000 and 6000. The Madagascar pratincole (Glareola ocularis) is a type of small, migratory wading bird found widely over centraleastern Africa, Madagascar, and the islands of the western Indian Ocean. Nevertheless, the total population is believed to be small and threatened by the loss and degradation of wetlands, particularly its breeding areas on the eastern coast of Madagascar.

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Madagascar The island of Madagascar is the world’s fourth largest and is, in many ways, a mini-continent. The seas around it are, in general, deep, with the wide Mozambique Channel which separates it from the East African coast reaching a depth of 3292 m. The prehistoric breakup of the supercontinent Gondwana separated the Madagascar–Antarctic–India landmass from the Africa–South America landmass around 135 million years ago. Madagascar later split from India about 88 million years ago, allowing plants and animals on the island to henceforth evolve in relative isolation. Biogeographically it is almost as peculiar as Australia, which has been isolated from Asia for more than 50 million years. In both cases evolution has followed a unique pattern, with prosimians in many ways taking over the role that marsupials serve in Australia. While much has been lost, much remains. Madagascar continues to be a major biodiversity hotspot, with over 90 per cent of its fauna and a similar percentage of its flora found nowhere else on Earth. Madagascar is tropical and chiefly mountainous, with a diverse range of habitats. A narrow and steep escarpment runs along the length of the eastern coast, which contains much of the island’s remaining lowland rainforest. To the west of this ridge lies a plateau in the centre of the island ranging in altitude from 750 to 1500 m. These central highlands are the most densely populated part of the island and are characterized by terraced, rice-growing valleys lying between grassy hills and patches of the subhumid forests that formerly covered the entire region. To the west of the highlands, the increasingly arid terrain gradually slopes down to the Mozambique Channel and to mangrove swamps along the coast. The western and southern sides are home to dry deciduous forests, spiny thickets, and xeric shrublands. Madagascar’s highest peaks rise from three prominent highland massifs, with Maromokotro (2876 m) in the Tsaratanana Massif the island’s highest point. Many of its endemic species are very specialized in terms of certain bioregions. Madagascar’s earliest human settlers found a rich prehistoric world very different from the one that we see today, one populated by an extraordinary megafauna including at least 17 species of lemur larger than any living species, some the size of a gorilla. Other highlights were at least two types of hippopotamus, gigantic crocodilians and tortoises, and an

Madagascar

extraordinary burrowing creature that must have looked like a cross between an aardvark and a cow. However, it is the various species of enormous ratites known as elephant birds that were the most spectacular. The largest birds that ever lived, one of them (Vorombe titan) stood fully 3 m high and weighed up to 730 kg. How much responsibility these early people had in wiping out these creatures is still open to debate, although it is clear that at least some may have survived up into historic times. Madagascar’s megafaunal extinctions were among the most severe for any continent or large island, with all endemic wildlife over 10 kg (at least two dozen species) disappearing even before the arrival of Europeans. The aye-aye (Daubentonia madagascariensis) is perhaps the most remarkable of all surviving lemurs. The world’s largest nocturnal primate, it resembles a witch’s cat in general appearance and is notable for its enormous ears, rodent-like, perpetually growing teeth and narrow middle fingers. All of these features are special adaptations for its unusual method of finding tree-boring insect larvae. Individuals tap on tree limbs while listening intently for the grubs burrowing within, then gnaw at the bark with their forward-slanting incisors in order to create a small hole into which they then insert their wire-like finger in order to extract it. A cryptic species, the aye-aye was rarely encountered after its discovery in the late eighteenth century, and by the 1930s was widely thought to be on the verge of extinction. It was rediscovered in 1957, however, and in 1966 nine individuals were transported to the island of Nosy Mangabe, in the Bay of Antongil off the north-eastern coast of Madagascar, in what was then considered a last-ditch effort to save it. By the 1980s sightings on the mainland became more frequent, and in the following decade a successful captive breeding programme was undertaken by a number of zoos around the world. While still rare and highly localized, the ayeaye is now known to be one of the most widespread of all lemurs, being found in both humid and dry forests across eastern, western, and northern Madagascar. Sadly, in many areas the species is shot on sight by local people in the erroneous belief that it is a threat to their crops, or because it is considered to be a harbinger of evil. The indri (Indri indri), the largest living lemur, was at the beginning of the nineteenth century still so common in eastern and north-eastern Madagascar that a traveller reported that no one could journey from Tamatave to Antanarivo without often hearing their eerie, whale-like calls as they passed through the great forests. The species declined along with the latter, but is still found widely, if very locally, from Anjanaharibe-Sud and Antohaka Lava (former Antsiranana province) south to the Anosibe An-ala Classified Forest (former Toamasina province). It is everywhere under threat, however, from habitat destruction and illegal hunting, and has never been kept in captivity apart from a brief period in Paris in 1939. Several species of sifaka (Propithecus) are found throughout the forested areas of Madagascar, where they are everywhere threatened by habitat destruction and, increasingly, subsistence hunting. Perrier’s sifaka (P. perrieri) is found in

Figure 4.1 Illustration of an indri. (Credit: Alfred Grandidier.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

both dry and humid forest fragments within a very small area in northernmost Madagascar (former Antsiranana province), where the total population is thought to number less than 500. The silky sifaka (P. candidus), one of the rarest and most threatened of all sifakas, has a very restricted range in northeastern Madagascar (former Antsiranana province), where it lives in both lower- and upper-elevation rainforest. The total population is thought to be less than 250. The diademed sifaka (P. diadema), second only to the indri (Indri indri) in size, occurs widely but patchily throughout the rainforests of eastern and north-eastern Madagascar (former Toamasina province) from the Mangoro and Onive rivers north to the Mananara River. Milne-Edwards’ sifaka (P. edwardsi) is confined to a small area of middle- and upper-elevation rainforest in central-eastern Madagascar (former Fianarantsoa province). Tattersall’s sifaka (P. tattersalli) is confined to a small area of north-eastern coastal Madagascar (former Antsiranana province) between the Loky River in the north and the Manambato in the south. The range centres on the town of Daraina, and consists of human-altered savanna, dry scrub,

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agricultural land, gallery forests, and forest fragments. In recent years gold-mining activities in the region have posed an additional threat. Coquerel’s sifaka (P. coquereli) is found patchily in the lowland dry forests as well as coastal mangroves of north-western Madagascar (former Mahajanga province) to the north and east of the Betsiboka River. The crowned sifaka (P. coronatus) is found over a relatively wide area of northwestern and central-western Madagascar (former Mahajanga and Antananarivo provinces) from the Mahavavy River to the Betsiboka River. Von der Decken’s sifaka (P. deckenii) occurs patchily in north-western and central-western Madagascar (former Mahajanga and Toliara provinces) between the Mahavavy and Manambolo rivers. Verreaux’s sifaka (P. verreauxi) occurs in a variety of habitats in southern and southwestern Madagascar (former Toliara province) including lowland and montane dry deciduous, spiny thicket, and even some lowland rainforest. It is, however, most associated with spiny woodlands, where they can be seen leaping recklessly between thorny trunks and bounding along the ground. Some hunting continues, but the main threat is habitat loss due to slash-andburn agriculture and charcoal and fuelwood production. The Sambirano woolly lemur (Avahi unicolor) is confined to two small, disjunct areas of the Sambirano region of northwestern Madagascar (former Antsiranana and Mahajanga provinces). Peyrieras’ woolly lemur (A. peyrierasi) is confined to a small area of lowland and montane rainforest in southeastern Madagascar (former Fianarantsoa province). Gmelin’s woolly lemur (A. laniger) remains widespread in eastern and north-eastern Madagascar, although all species are threatened by loss of habitat and subsistence hunting. The ruffed lemurs (Varecia) are large and garishly coloured. Three subspecies of pied ruffed lemur (V. variegata) are found discontinuously throughout the eastern rainforests. The black and white ruffed lemur (V. v. variegata) occurs in eastern Madagascar (former Toamasina and Fianarantsoa provinces) south of the Anove River from about Ambatovaky south to Betampona and Zahamena National Park. The whitebelted ruffed lemur (V. v. subcincta) occurs in north-eastern Madagascar (former Toamasina province) from the Antainambalana River south to the Anove River, including parts of Makira, Mananara-Nord, Atialanankorendrina, and Marotandrano. This subspecies was additionally introduced to the island of Nosy Mangabe in the Bay of Antongil in the 1930s and still occurs there. Hill’s black and white ruffed lemur (V. v. editorum) is only known with certainty from southeastern Madagascar (former Fianarantsoa province) from Mantadia southwards to the Manombo Special Reserve. The red ruffed lemur (V. rubra) is confined to the Masoala Peninsula and the region immediately north of the Bay of Antongil in north-eastern Madagascar (former Antsiranana province). All of these forms are seriously threatened by habitat destruction and hunting. The ring-tailed lemur (Lemur catta), an iconic symbol of Madagascar, remains relatively widespread in the dry forests and bush of the south and south-west (former Toliara and

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Fianarantsoa provinces). Nevertheless, its range is highly fragmented and population densities are low. Habitat destruction and hunting are the chief threats. Several species of so-called true lemur (Eulemur) are found throughout Madagascar, where they are threatened by habitat destruction, hunting, and capture for use as pets. The crowned lemur (E. coronatus) inhabits extreme northern Madagascar (former Antsaranana province), where it prefers semideciduous dry lowland and mid-altitude forest but may be found in practically all forest types, including high-altitude moist forest, wooded savanna, and even agricultural areas. The mongoose lemur (E. mongoz) is found in north-western Madagascar (former Mahajanga province) in the region of Ambato-Boéni and Ankarafantsika, with introduced populations also occurring on the islands of Mohéli, Anjouan, and Grande Comore in the Comoros Islands. Sanford’s lemur (E. sanfordi) has a restricted range in northern Madagascar (former Antsiranana province), with the southerly limit being the Manambato River. The range is centered on Ankarana, Anamalerana, and Montagne d’Ambre, with a disjunct population in Daraina to the south-east. It is found from sea level to 1400 m and thus inhabits both dry and subhumid forest, but seems to be more common and to occur at higher densities in the latter. The red-bellied lemur (E. rubriventer) is found over a wide area from the Tsaratanana Massif in northern Madagascar south along the eastern rainforests (excluding the Masoala Peninsula) to the Pic d’Ivobe and the Manampatrana River, although at one time it ranged further south. The white-fronted lemur (E. albifrons) is found throughout most of the remaining rainforest in north-eastern Madagascar (former Antsiranana and Toamasina provinces), from the Bemarivo River near Sambava south to the region of Mananara Nord, including the Masoala Peninsula. There is an isolated population in the Betampona Nature Reserve further south, and the species has also been introduced to Nosy Mangabe. Its distribution south of Mananara remains to be clarified as there is significant hybridization with the brown lemur (E. fulvus) over a wide area. The white-collared lemur (E. cinereiceps) is largely confined to a thin strip of lowland rainforest running from the Andringitra Massif south to the Mananara River in south-eastern Madagascar (former Fianarantsoa province). Isolated populations also occur in the Manombo Special Reserve and in the Mahabo Forest south of Farafangana, and possibly at Vohipaho, south of the Mananara River. The rufous lemur (E. rufus) occurs in north-western Madagascar (former Mahajanga province) from the Betsiboka River south to the Tsiribihina. The red-fronted lemur (E. rufifrons) occurs disjunctly in south-eastern and south-western Madagascar (former Toliara, Fianarantsoa, and Toamasina provinces). The red-collared lemur (E. collaris) is found in south-eastern Madagascar (former Toliara and Fianarantsoa provinces) roughly from Tolagnaro north to the Mananara River, with the western limits of the range being the forests of the Kalambatritra region. The black lemur (E. macaco) occurs in moist forests of the Sambirano region of

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north-eastern Madagascar (former Antsiranana and Mahajanga provinces) and on the islands of Nosy Be, Nosy Komba, and Nosy Tanikely. Sclater’s lemur (E. flavifrons) is confined to a small area in north-eastern Madagascar (former Mahajanga province) consisting of the Sahamalaza Peninsula along with a narrow stretch of forest on the adjacent mainland. The brown lemur (E. fulvus) is found widely but disjunctly in northern and eastern Madagascar, with an introduced population in the Comoros Islands. The greater bamboo lemur (Prolemur simus) is one of the world’s most critically endangered primates. Subfossil remains confirm that it was once widespread across northern, central and eastern Madagascar, but by the mid-twentieth century it was thought to be extinct, a victim of habitat destruction and extreme dietary specialization. A remnant population was discovered in 1986, however, and since then surveys in southern and central-eastern Madagascar have discovered around 500 individuals living in 11 subpopulations. Unfortunately, all live in areas of severely degraded and unprotected habitat. Several bamboo lemur species and subspecies of the genus Hapalemur are threatened by habitat destruction, subsistence hunting, and capture for use as pets. The golden bamboo lemur (H. aureus) was first discovered in 1986 in what is now Ranomafana National Park, in central-east Madagascar (former Fianarantsoa province). It is now known to also occur in Andringitra National Park and in a forest corridor connecting the two, as well as in the region of Betsakafandrika. The northern bamboo lemur (H. occidentalis) is found in a number of discontinuous pockets in northern Madagascar (former Antsiranana, Mahajanga, and Toamasina provinces). It is known (at least formerly) from the forests of Ankarana and Analamerana in the far north and possibly on the Ankarana Massif as well, as well as from the Ampasindava Peninsula, and the Sambirano and Sahamalaza regions in the north-west; the Tsiombikibo, Baie de Baly; Tsingy de Namaroka and Bongolava regions in the central-west; Masoala, Maroansetra, and Ile Roger (Aye aye Island) in the north-east; and as far south as Zahamena and Marovohangy near Lake Alaotra. Three subspecies of grey bamboo lemur (H. griseus) are found disjunctly in eastern and western Madagascar. The Mantadia grey bamboo lemur (H. g. griseus) is restricted to east-central Madagascar (former Toamasina province), roughly from the Onibe River south to the Onive River. The Beanamalao bamboo lemur (H. g. gilberti) is known from a small area of central-eastern Madagascar (former Toamasina province) between the Onive and Nosivolo rivers. The Ranomafana bamboo lemur (H. g. ranomafanensis) occurs in two widely separated populations in east-central and west-central Madagascar. In the east it is known from the forests to the south of the Mangoro and Onive rivers in former Fianarantsoa province, while in the west it has been reported from the forests of Tsingy de Bemaraha, probably as far as the Betsiboka River in former Mahajanga province. The southern bamboo lemur (H. meridionalis) is found in south-eastern Madagascar (former Toliara and Fianarantsoa

provinces), roughly from Andohahela north to the Mananara River. The Sahamalaza sportive lemur (Lepilemur sahamalazensis) is restricted to the three remaining humid and dry forest blocks on the Sahamalaza Peninsula in north-eastern Madagascar (former Mahajanga province). Only some of its range lies within protected areas. The Daraina sportive lemur (L. milanoii) and the Ankarana sportive lemur (L. ankaranensis) are both confined to forest fragments in far northern Madagascar (former Antsiranana province). Seal’s sportive lemur (L. seali) is confined to a small area of north-eastern Madagascar (former Antsiranana province). The weasel sportive lemur (L. mustelinus) is found disjunctly in north-eastern coastal Madagascar (former Toamasina province). All are threatened by habitat destruction and hunting. The Montagne d’Ambre fork-marked lemur (Phaner electromontis) is found sporadically in the regions of Montagne d’Ambre and Ankarana in far northern Madagascar (former Antsaranana province), with a further population at Daraina likely representing a distinct species. The Masoala forkmarked lemur (P. furcifer) is found over a relatively wide area of north-eastern Madagascar (former Antsiranana and Toamasina provinces). Both are threatened by loss of habitat. Three species of dwarf lemur (Cheirogaleus) are threatened by habitat destruction and hunting. The Ankarana dwarf lemur (C. shethi) is found patchily in forest and shrubby areas of far northern Madagascar (former Antsiranana province). Crossley’s dwarf lemur (C. crossleyi) is found disjunctly in eastern and north-eastern Madagascar (former Antsiranana and Toamasina provinces). The Lavasoa dwarf lemur (C. lavasoensis) is known disjunctly from forest patches in the Lavasoa–Ambatotsirongorongo Mountains and from the Kalambatritra (Sahalava) Forest in south-eastern Madagascar (former Toliara and Fianarantsoa provinces). The hairy-eared dwarf lemur (Allocebus trichotis) is a tiny, enigmatic species. Long known from only a few museum specimens collected during the late nineteenth century, it was not seen again until 1966. It was thereafter feared to be extinct until rediscovered in 1989 in lowland rainforest in northeastern Madagascar. Individuals have since been found living in highland rainforest at Andasibe in eastern Madagascar and in other localities as well, indicating a more widespread, if patchy, distribution. The northern giant mouse lemur (Mirza zaza) is confined to an area of north-eastern Madagascar centered on the Ampasindava Peninsula (former Antsiranana province). Several species of mouse lemur (Microcebus) are threatened by loss of habitat. The Tavaratra mouse lemur (M. tavaratra) and Arnhold’s mouse lemur (M. arnholdi) are both found patchily in far northern Madagascar (former Antsiranana province). Mittermeier’s mouse lemur (M. mittermeieri) is found patchily in both lowland and highland rainforest in northeastern Madagascar (former Antsiranana province). The Sambirano mouse lemur (M. sambiranensis) is known only from the few localities in north-eastern Madagascar (former

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Antsiratana and Mahajanga provinces). Margot Marsh’s mouse lemur (M. margotmarshae) is found mainly in the Antafondro Classified Forest of north-eastern Madagascar (former Mahajunga province), and perhaps also in highelevation areas of the Tsaratanana Special Reserve. The Anosy mouse lemur (M. tanosi) is confined to lowland and montane rainforests in the south-eastern corner of Madagascar (former Toliara province). Peters’ mouse lemur (M. myoxinus) has a disjunct distribution in central-western Madagascar (former Mahajanga and Toliara provinces) between the northern banks of the Tsiribihina River north to Baie de Baly, the forests of Belo sur Tsiribihina and Aboalimena, and in the Tsingy de Bemaraha and Tsingy de Namoroka national parks. The fossa (Cryptoprocta ferox), the largest carnivore of Madagascar, is found throughout the country except for the Central Plateau, but is everywhere threatened by habitat destruction, hunting, and human persecution. The eastern falanouc (Eupleres goudotii) is a mongooselike carnivore that is widespread throughout the length of eastern Madagascar, but rare and extensively hunted. The spotted fanaloka (Fossa fossana) is a small, civet-like species found over much of eastern and north-eastern Madagascar, where it is vulnerable to hunting and habitat destruction. The broad-striped vontsira (Galidictis fasciata) is a type of mongoose found from Marojejy National Park throughout the eastern rainforest as far south as Andohahela, although always at very low densities. The web-footed tenrec (Limnogale mergulus) is a semiaquatic insectivore known only from a small number of stream and river habitats across a wide area of east-central Madagascar. The northern shrew tenrec (Microgale jobihely) is known from two highly disjunct populations in eastern and northeastern Madagascar (former Mahajanga and Toamasina provinces). Nasolo’s shrew tenrec (M. nasoloi) is known only from a few specimens collected from three disjunct localities in south-western Madagascar (former Toliara province). Petter’s tuft-tailed rat (Eliurus petteri) was, until 2003, known only from three specimens, but has since been discovered in a few localities in central-eastern Madagascar (former Toamasina province). Ellerman’s tuft-tailed rat (E. ellermani) is known only from two specimens, one collected some 40 km north-west of Maroantsetra in north-eastern Madagascar and the other in the vicinity of Lohariandava in the central-east (former Toamasina province). The whitetipped tuft-tailed rat (E. penicillatus) was originally described in the early twentieth century from Ampitambe Forest near Ambositra in east-central Madagascar (former Fianarantsoa province). It was not recorded again until 2000 when it was collected at Anerana, about 35 km northeast of Fandriana. The Madagascan flying fox (Pteropus rufus) is widespread on Madagascar, with the highest density of roost sites being

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in the coastal regions, especially from Morombe in the south-west to Antsiranana in the north. It is intensively hunted for food. The Madagascan fruit bat (Eidolon dupreanum) is found over much of Madagascar with the exception of the eastern rainforests. It is everywhere rare, however, and dependent upon suitable caves for roosting sites, where it too is frequently hunted by local people for food. The Comoros bent-winged bat (Miniopterus griveaudi) is known from Grande Comore and Anjouan as well as from a few localities in northern and western Madagascar. The western yellow bat (Scotophilus tandrefana) is similarly known only from two specimens, one from near Parc National Tsingy de Bemaraha and the second at Sarodrano (former Mahajanga and Toliara provinces). The Madagascar slit-faced bat (Nycteris madagascariensis) is known only from two specimens collected in 1910, in the Irodo River Valley of northern Madagascar near Analamera (former Antsiranana province). It has not been found since, despite surveys. Racey’s pipistrelle bat (Pipistrellus raceyi) is known only from a few specimens collected at two lowland rainforest localities in the central-east and two more, lowland dry forest sites in the central-west. The Madagascar serpent eagle (Eutriorchis astur) is a rare inhabitant of the eastern rainforests from Marojejy in the north to Zahamena in the south, as well as the Central Highlands at Anjozorobe and Bemanevika. The total population likely numbers less than 1000. The Madagascar fish eagle (Haliaeetus vociferoides) survives in low numbers within wooded areas adjacent to mangroves and shallow bays along the western coast of Madagascar (former Antsiranana, Mahajanga, and Toliara provinces). Its numbers have declined dramatically in recent years, and it is now one of the world’s rarest birds of prey. Only about 120 breeding pairs survive. The Madagascar red owl (Tyto soumagnei) was long feared to be extinct but is now known to be fairly widespread throughout the eastern rainforests. However, the total population is small and severely fragmented. Bernier’s teal (Anas bernieri) is found in a narrow strip along almost the whole of the west coast and extreme northeast of Madagascar (former Antsiranana, Mahajanga, and Toliara provinces), in both mangrove and freshwater wetlands. It is threatened by habitat destruction and disturbance, and is frequently hunted. The Madagascar rail (Rallus madagascariensis) occupies suitable wetland habitat in the east up to 1800 m, where it is generally rare if locally not uncommon. The white-breasted mesite (Mesitornis variegatus) is a raillike ground-dweller found at five localities in northern and central-western Madagascar (Menabe Forest, the Ankarafantsika complex, the Ankarana Special Reserve, Analamera Special Reserve, and the Daraina forests), as well as in the Ambatovaky Special Reserve in the east. Surveys have demonstrated its genuine

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absence from many intervening areas, including some with apparently suitable habitat. The helmeted vanga (Euryceros prevostii) is a shrike-like bird found patchily in the rainforests of eastern and northeastern Madagascar (former Antsiranana and Toamasina provinces), where it is threatened by loss of habitat. The short-legged ground-roller (Brachypteracias leptosomus) is still fairly common in eastern Madagascar from Daraina Forest in the north to Andohahela in the south, where it is restricted to undisturbed primary forest. Appert’s greenbul (Xanthomixis apperti) is a small, terrestrial bird known only from four tiny forest patches in southwestern Madagascar (former Toliara province). The spider tortoise (Pyxis arachnoides) is confined to the arid region of south-western Madagascar (former Toliara province) from the coast up to 10–50 km inland going as far north as Morombe. There are three subspecies. The nominate form (P. a. arachnoides) occurs in the region of the Onilahy River near Toliara, Brygoo’s spider tortoise (P. a. brygooi) occurs south of the Mangoky River, and the oblong spider tortoise (P. a. oblonga) along the southern coast. All are threatened by habitat destruction and fragmentation, as well as by hunting for food. Petter’s chameleon (Furcifer petteri) is known from a few localities over a relatively wide area of northern Madagascar (former Antsiranana and Mahajanga provinces). The twobanded chameleon (F. balteatus) is known from two widely disjunct areas in the central south-east and far south-east (former Fianarantsoa and Toliara provinces), where it is rarely encountered. Laborde’s chameleon (Furcifer labordi) is found in both dry deciduous and spiny succulent forest in western and south-western Madagascar (former Toliara province). All are threatened by habitat destruction and overcollection for the exotic pet trade. Two subspecies of elongated leaf chameleon (Palleon nasus) inhabit areas of higher-elevation humid forest in south-eastern Madagascar (former Fianarantsoa and Toliara provinces). Boulenger’s elongated leaf chameleon (P. n. nasus) is known only from its type locality of Manjarivolo, while Paulian’s elongated leaf chameleon (P. n. pauliani) is fairly widespread. Both are threatened by habitat destruction. The leaf chameleons (Brookesia) are a group of dullcoloured, mostly terrestrial species that include some of the world’s smallest reptiles. The northern leaf chameleon (B. ebenaui) is known from a few small areas of both dry and humid lowland forest in far northern Madagascar (former Antsiranana and Mahajanga provinces). The mossy leaf chameleon (B. vadoni) is confined to a small area of north-eastern Madagascar (former Antsiranana and Toamasina provinces). Ramanantsoa’s leaf chameleon (B. ramanantsoai) is known only from a few localities in east-central Madagascar (former Antananarivo and Toamasina provinces). Peyrieras’ leaf chameleon (B. peyrierasi) is confined to the Masoala Peninsula and the island of Nosy Mangabe in north-eastern Madagascar (former Antsiranana and Toamasina provinces).

The Nosy Be leaf chameleon (B. minima) is confined to lowland forest fragments on Nosy Be and Nosy Komba and a few other localities in the north-west including Manongarivo and, most probably, Sahamalaza. All are threatened by loss of habitat. Seipp’s day gecko (Phelsuma seippi) is known from a few localities in the Sambirano region of northern Madagascar (former Antsiranana and Mahajanga provinces), and from the islands of Nosy Be and Nosy Komba. It is threatened by loss of habitat. Boivin’s velvet gecko (Blaesodactylus boivini) is known from two disjunct areas of northern Madagascar (former Antsiranana and Mahajanga provinces) along with some offshore islands. The leaf-tailed geckos (Uroplatus) are nocturnal and arboreal, with highly cryptic colouration used as camouflage. The Malahelo leaf-tailed gecko (U. malahelo) has been recorded from widely scattered localities in southern Madagascar including Ambatotsirongorongo, Andohahela, Kalambatritra, and Analavelona. The Malama leaf-tailed gecko (U. malama) has been recorded from a few localities in south-eastern Madagascar (former Fianarantsoa and Toliara provinces), including Andohahela, Ampamakiesiny, Kalambatritra, Ranomafana-Sud, Ivorona, Farafara, and Pic Ivohibe. Ebenau’s leaf-tailed gecko (U. ebenaui) is found in far northern Madagascar (former Antsiranana province), including the island of Nosy Be. The giant leaf-tailed gecko (U. giganteus), the world’s second largest living species of gecko, is found in north-eastern Madagascar, where it has been reported from Montagne d’Ambre and Marojejy south into the Masoala Peninsula (former Antsiranana province). All are highly sought after for the international pet trade. The four-striped plated lizard (Zonosaurus quadrilineatus) is confined to a small area of south-western coastal Madagascar (former Toliara provinces), where it lives in dry forest patches and sand dunes. Boettger’s plated lizard (Z. boettgeri) is found widely but patchily in north-eastern Madagascar (former Antsiranana province), including the island of Nosy Be. Peters’ keeled plated lizard (Tracheloptychus petersi) is found in a few mostly arid and sandy localities in southwestern coastal Madagascar (former Toliara province). Millot’s skink (Paracontias milloti) is known only from museum specimens collected from the small islands of Nosy Mitsio and Nosy Mamoko off the north-western coast of Madagascar, and possibly from Marojejy in the north-east. It is possible that the species was confined to areas of now-lost lowland humid forest and has disappeared. O’Shaughnessy’s skink (Brachyseps gastrostictus) has only been recorded from two widely separated localities in eastern Madagascar (Anjanaharibe-Sud in the north-east and Andrangoloaka further south), which suggests that the species is likely to occur between them as well. The Anosy skink (B. anosyensis) is known from a few scattered localities across a relatively wide area of south-eastern Madagascar (former

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Fianarantsoa and Toliara provinces). The splendid skink (B. splendidus) is a rare species known only from two extremely disjunct areas of central-western and southeastern Madagascar. Alluaud’s skink (Flexiseps alluaudi) is only known from Montagne d’Ambre and a few localities within the Sambirano region of northern Madagascar (former Antsiranana province), as well as from the island of Nosy Be. The Mandokava skink (F. mandokava) is confined to a few localities in northern Madagascar (former Antsiranana and Mahajanga provinces). Bocage’s skink (Pseudoacontias madagascariensis) is a large, fossorial species with severely reduced limbs known only from a single specimen collected from an undefined locality in northern Madagascar during the late nineteenth century. Angel’s skink (P. angelorum) has been recorded at two localities in north-eastern Madagascar (former Antsiranana province). The Betsileo mabuya skink (Trachylepis betsileana) is known only from a single specimen said to have been collected during the early nineteenth century from east-central Madagascar, but which may have possibly originated from somewhere in mainland Africa. Günther’s tree snake (Lycodryas guentheri) is known only from a few specimens collected in disjunct areas of the Central Highlands and south-eastern Madagascar. The lemon tree snake (L. citrinus) is known only from a few localities within a small area of western coastal Madagascar (former Mahajanga and Toliara provinces). Both are threatened by loss of habitat and collection for the international pet trade. The yellow-striped water snake (Thamnosophis stumpffi) is confined to the island of Nosy Be and adjacent mainland areas of suitable habitat in north-western Madagascar (former Antsiranana province). The Madagascar burrowing snake (Pararhadinaea melanogaster) is divided into two subspecies. The nominate form (P. m. melanogaster) is known only from a few localities across a relatively wide area of northern Madagascar (former Antsiranana province) and the island of Nosy Be. The small-headed blind snake (Madatyphlops microcephalus) is a burrowing species known from the lowland humid and dry forests of Montagne d’Ambre, Ampombofofo, and Montagne des Français in north-eastern Madagascar (former Antsiranana province). Trueb’s Madagascar tree frog (Madecassophryne truebae) is known only from a few localities in south-eastern Madagascar (former Toliara province). The Maharipeo cophyline tree frog (Cophyla maharipeo) is known only from a small population living in an abbey garden at Joffreville, and in the nearby Fontenay Private Nature Park in far northern Madagascar (former Antsiranana province). The four-spotted tree cophyline tree frog (C. tetra) is known only from a few localities in northeastern Madagascar (former Antsiranana province). Millot’s cophyline tree frog (C. milloti) is confined to a small area of

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northern Madagascar (former Antsiranana province), including the island of Nosy Be. All are threatened by loss of habitat. The Ranomafana climbing frog (Anodonthyla moramora) is known only from a small area of south-eastern Madagascar (former Fianarantsoa province), centered on Ranomafana National Park. A large and unique group endemic to the Madagascan Realm are the so-called skeleton frogs (Boophis), so named for their almost translucent skin that allows their bones and internal organs to be observed. They are arboreal and notably ‘tree frog-like’ in terms of their appearance and mode of life. Many are highly threatened by loss of habitat. Sandra’s skeleton frog (B. sandrae) is known only from a few localities in the region of Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). Andreone’s skeleton frog (B. andreonei) is known from a few localities within a small area of northern Madagascar (former Mahajanga and Antsiranana provinces). Jaeger’s skeleton frog (B. jaegeri) occurs on Nosy Be and the adjacent Sahamalaza Peninsula in north-eastern Madagascar (former Antsiranana and Mahajanga provinces). The Sambirano skeleton frog (B. sambirano) is known from the Manongarivo Special Reserve and in a few localities in and around the Tsaratanana Reserve in northern Madagascar (former Antsiranana province). Blommers’ skeleton frog (B. blommersae) is confined to two disjunct areas separated by unsuitable habitat in northern Madagascar (former Antsiranana province). Ulftunn’s skeleton frog (B. ulftunni) is known from a few localities in northern Madagascar (former Antsiranana province). The banded skeleton frog (B. vittatus) is found widely but disjunctly in northern Madagascar (former Anstiranana province). The Haingana skeleton frog (B. haingana) is known from two localities in the region of the Andohahela National Park in south-eastern Madagascar (former Toliara province), but likely occurs a little more widely. The golden-flanked skeleton frog (B. andohahela) is found widely but patchily in south-eastern Madagascar (former Fianarantsoa and Toliara provinces). Böhme’s skeleton frog (B. boehmei) is found widely but patchily in eastern and south-eastern Madagascar (former Toamasina, Fianarantsoa and Toliara provinces). Boettger’s skeleton frog (B. brachychir) is found patchily over a wide area of northern and western Madagascar. The green mantella (Mantella viridis) is confined to a small area of far northern Madagascar (former Antsiranana province) and the island of Nosy Hara. It is threatened by loss of habitat and collection for the international pet trade. Angel’s Madagascar frog (Boehmantis microtympanum) is confined to forest fragments in far south-eastern Madagascar (former Toliara province). Boettger’s stump-toed frog (Stumpffia psologlossa) is known only from a small area of north-eastern Madagascar (former Antsiranana province), including the island of Nosy Be. A number of grainy frogs (Gephyromantis) are threatened by loss of habitat. Klemmer’s grainy frog (G. klemmeri) is

Madagascar

known only from Marojejy, Anjanaharibe-Sud, and Betaolana in north-eastern Madagascar (former Antsiranana province). Webb’s grainy frog (G. webbi) and the Silvanus grainy frog (G. silvanus) are both known only from a few localities in the region of Antongil Bay and on Nosy Mangabe in north-eastern Madagascar (former Fianarantsoa province). Boettger’s grainy frog (G. horridus) is known only from three localities in northern Madagascar (Montagne d’Ambre, Tsaratanana and, at least historically, the island of Nosy Be, all in former Antsiranana province). The Isalo grainy frog (G. corvus) is known only from the Isalo and Makay massifs of southwestern Madagascar (former Toliara province). The Ambohitra grainy frog (G. ambohitra) is known from Montagne d’Ambre, Tsaratanana, and Manongarivo in northern Madagascar (former Antsiranana province). The Zavona grainy frog (G. zavona) and the Tandroka grainy frog (G. tandroka) are both confined to a few localities in northern Madagascar (former Antsiranana and Mahajanga provinces). Enk’s grainy frog (G. enki) is confined to the region of Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). The Salegy grainy frog (G. salegy) and the striated grainy frog (G. striatus) are both found relatively widely in northern Madagascar (former Antsiranana and Toamasina provinces). The greater spiny grainy frog (G. spiniferus) is known from a few localities in south-eastern Madagascar (former Fianarantsoa and Toliara provinces). The marbled rain frog (Scaphiophryne marmorata) is known from a few lowland and highland localities in centraleastern Madagascar (former Toamasina province), where it is threatened by loss of habitat. The obscure rain frog (S. obscura) is known only from an undefined locality somewhere on the north-western coast of Madagascar. The warty rain frog (S. verrucosa) is known only from its original collection in the early twentieth century from coastal south-western Madagascar (former Toliara province). Mass’ stream frog (Spinomantis massi) is known only from a few localities in northern Madagascar (former Mahajanga and Antsiranana provinces). The Tavaratra stream frog (S. tavaratra) is confined to a small area of northern Madagascar centered on the Marojejy Massif (former Antsiranana and Mahajanga provinces).

Mountains and Highlands The most important highland regions of Madagascar include a narrow and steep escarpment running along the eastern coast, containing much of the island’s remaining tropical rainforest. To the west of this ridge lies a plateau in the centre of the island known as the Central Highlands and, north of that, the Northern Highlands. Madagascar’s highest peaks rise from three prominent massifs (Tsaratanana, Andringitra, and Ankaratra). They provide a range of habitats from subhumid forest, grassland and woodland mosaics, and montane heathlands.

Sibree’s dwarf lemur (Cheirogaleus sibreei) is known only from a few small but widely separated pockets of high-altitude rainforest in eastern Madagascar. The yellow-bellied asity (Neodrepanis hypoxantha) is a type of sunbird that was long known only from specimens collected prior to 1933 and thought to be extinct. It has since been found in scattered higher-elevation forests throughout the length of eastern Madagascar. O’Shaughnessy’s chameleon (Calumma oshaughnessyi) is found widely but patchily along the eastern and southern highlands of Madagascar (former Toamasina, Antananarivo, Fianarantsoa, and Toliara provinces), where it is threatened by loss of habitat. The Makira skink (Paracontias vermisaurus) is known only from two localities within the Makira Reserve in north-eastern Madagascar (former Toamasina province). The Kankana skink (P. kankana) is known only from a single locality near Lake Alaotra, but likely occurs in a few other areas in northeastern Madagascar. Coulanges’ tree snake (Brygophis coulangesi) was long known only from a single specimen collected in 1968 at Fierenana in central-eastern Madagascar (former Toamasina province). In the mid 1990s a second was discovered at Andasaribe-Sud in north-eastern Madagascar (former Antsiranana province), suggesting that the species is widespread but very rare and localized. Grandidier’s water snake (Liopholidophis grandidieri) is a rare inhabitant of high-altitude rainforest and arid shrubland in east-central Madagascar (former Fianarantsoa province). Known localities include Ranomafana National Park and surroundings, as well as the Tsinjoarivo region. The rounded rain frog (Scaphiophryne boribory) is known from two disjunct localities in north-eastern and northern Madagascar (former Toamasina province). Guibe’s stream frog (Spinomantis guibei) is known only from a few localities in south-eastern Madagascar (former Toliara and Fianarantsoa provinces).

The Northern Highlands The Northern Highlands are a mountainous region in northeastern Madagascar (former Antsiranana province) that includes the Tsaratanana Massif and smaller nearby massifs such as Marojejy, Anjanaharibe-Sud, and Manongarivo. It is separated from the Central Highlands by a valley known as the Mandritsara Window, which acts as a barrier to dispersal between the two. They are noted for their extremely high level of species endemism. The montane shrew tenrec (Microgale monticola) is known from only four localities within a very narrow elevation range (1500–1950 m) on two massifs in north-eastern Madagascar (former Antsiranana province), the latter encompassing Marojejy National Park and the Anjanaharibe-Sud Special Reserve. The hairy-tailed tree rat (Brachytarsomys villosa) is a poorly known nocturnal and arboreal species known only

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from five localities in the Northern Highlands (former Antsiranana, Mahajanga, and Toamasina provinces). The bizarre-nosed chameleon (Calumma hafahafa) was originally known from a single male collected in 2003 from a forest at the Bemanevika Lakes in north-western Madagascar (former Mahajanga province). A handful of additional specimens have subsequently been taken from the same general area, as well as from a forest fragment near the Tsaratanana Massif (former Antsiranana provinces). The lined leaf chameleon (Brookesia lineata) is known only from two localities at Manongarivo in north-eastern Madagascar (former Mahajanga province) along with a third at Tsaratanana (former Antsiranana provinces). The Bekolosy leaf chameleon (B. bekolosy) is known only from a single specimen collected in 1992 from the Bekolosy Plateau, within the Manongarivo Special Reserve in north-eastern Madagascar (former Antsiranana province). The Tsaratanana cophyline tree frog (Cophyla tsaratananaensis) was long known only from three specimens collected in 1949 from the Tsaratanana Massif in north-eastern Madagascar, but has since been found at two additional montane localities. The yellowish cophyline tree frog (C. mavomavo) is known only from a small area of the Northern Highlands, but may occur more widely. The Anjanaharibe skeleton frog (Boophis anjanaharibeensis) is known only from the Anjanahribe-Sud Special Reserve, Ambolokopatrika, and Tsararano, and Marojejy National Park (former Antsiranana province). The Tahotra grainy frog (Gephyromantis tahotra) and Schilf’s grainy frog (G. schilfi) are both known from a few localities in northern Madagascar (former Antsiranana and Mahajanga provinces). Both are threatened by loss of habitat. The chubby diamond frog (Rhombophryne botabota) is known only from a few localities within the Northern Highlands, including Marojejy National Park and the Makira Plateau. Günther’s digging frog (Plethodontohyla guentheri) is known only from Marojejy National Park and the Anjanharibe-Sud Special Reserve. Montagne d’Ambre (Amber Mountain) is an isolated volcanic massif located near the northern tip of Madagascar (former Antsiranana province). It contains a significant pocket of subhumid forest surrounded at lower elevations by dry deciduous forest. Known for its high levels of endemism, it has long been protected within Montagne d’Ambre National Park. The Montagne d’Ambre dwarf lemur (Cheirogaleus andysabini) is confined to Montagne d’Ambre National Park and a few areas near the town of Joffreville. The Montagne d’Ambre rock thrush (Monticola erythronotus) is confined to the subhumid forests of Montagne d’Ambre, where the total population is thought to be less than 5000. The Montagne d’Ambre leaf chameleon (Brookesia tuberculata) is confined to Montagne d’Ambre. The Forêt d’Ambre

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leaf chameleon (B. desperata) is known only from the Forêt d’Ambre Special Reserve. Puellars’ cophyline tree frog (Cophyla puellarum) and Noromalala’s cophyline tree frog (C. noromalalae) are both confined to high-elevation rainforest in Montagne d’Ambre National Park. The Matavy diamond frog (Rhombophryne matavy) is known only from a single lowland forest locality within the Forêt d’Ambre Special Reserve. The Montagne d’Ambre stump-toed frog (Stumpffia madagascariensis) is confined to Montagne d’Ambre. The Marojejy Massif is a mountain chain located in northeastern Madagascar (former Antsiranana province). Access to the area around the massif was restricted to research scientists when it was set aside as a strict nature reserve in 1952. In 1998 it was opened to the public when it was converted into a national park. Originally seen as a transition zone between the eastern rainforests and the central highlands, Marojejy is now recognized as having its own unique features, with some of the richest biodiversity on the island. The wide range of elevations and rugged topography create diverse habitats that transition quickly with changes in altitude. Warm, dense rainforest can be found at lower elevations, followed by subhumid and cloud forest, and topped near the peaks with the only remaining, undisturbed montane shrubland in Madagascar. Unfortunately poaching, selective logging, and wood collection are still persistent problems within the park, particularly since the start of the 2009 political crisis. Moreover, a single outbreak of fire could result in the loss of large parts of the montane eroicoid habitat, as it already has at other sites. Two chameleons of the genus Calumna from Marojejy are threatened by habitat destruction. The Marojejy peak chameleon (C. jejy) is confined to the higher elevations between 1800 and 2130 m. Peyrieras’ chameleon (C. peyrierasi) is known only from a few specimens. The Marojejy leaf chameleon (Brookesia karchei) is confined to the Marojejy Massif. The Marojejy burrowing snake (Pararhadinaea melanogaster marojejyensis) is known only from a single specimen collected within Marojejy National Park. The Marojejy cophyline tree frog (Cophyla rava) is known only from high elevations within Marojejy National Park. The Ranjomavo grainy frog (Gephyromantis ranjomavo) is known only from the Marojejy Massif. The Vaventy diamond frog (Rhombophryne vaventy), tiny diamond frog (R. minuta), and Guibé’s diamond frog (R. serratopalpebrosa) are all known only from the Marojejy Massif, where they are threatened by loss of habitat. The Marojejy stump-toed frog (Stumpffia roseifemoralis) is known only from the Marojejy Massif. The Sorata Massif is located in north-eastern Madagascar (former Antsiranana province). The Sorata diamond frog (Rhombophryne longicrus) is confined to the Sorata Massif.

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The Manongarivo Massif is located in north-eastern Madagascar (former Antsiranana province). The Manongarivo leaf chameleon (Brookesia valerieae) is known only from two isolated, low-elevation localities at Manongarivo. The Tsaratanana Massif is located in north-central Madagascar (former Antsiranana province). It features the highest mountain in Madagascar, Maromokotro. The Tsaratanana chameleon (Calumma tsaratananense) is confined to the high-elevation heathland of Tsaratanana. The Tsaratanana skink (Paracontias manify) is confined to the Tsaratanana Massif. Olga’s cophyline tree frog (Cophyla olgae) and Guibé’s cophyline tree frog (C. alticola) are both known only from a few specimens collected from the highest parts of the Tsaratanana Massif. Nussbaum’s stream frog (Spinomantis nussbaumi) is known only from its type locality on the Befosa River, Antetikalambazaha, Tsaratanana Massif. The ornate diamond frog (Rhombophryne ornata), Guenther Peters’ diamond frog (R. guentherpetersi), and the Tany diamond frog (R. tany) are all confined to the Tsaratnana Massif, where they are threatened by loss of habitat.

The Central Highlands The Central Highlands or Central High Plateau is comprised of the contiguous part of the island’s interior above 800 m in the east and 600 m in the west. It is separated from the Northern Highlands by a low-lying valley known as the Mandritsara Window, which has apparently acted as a barrier to dispersal between the two. It is thought that subhumid forest originally extended over most of it, although this has long since been replaced by grassland and agriculture, with actual forest reduced to fragmented patches (most notably at Anjozorobe and Ambohitantely). A number of massifs rise out of the highlands and represent some of Madagascar’s highest mountains. Wright’s sportive lemur (Lepilemur wrightae) is known only from the Kalambatritra Special Reserve and the unprotected Beakora Forest in south-eastern Madagascar (former Fianarantsoa province). Goodman’s mouse lemur (Microcebus lehilahytsara) is confined to forest fragments in east-central Madagascar (former Toamasina province). The eastern voalavo (Voalavo antsahabensis) is a type of rodent known only from montane forest surrounding Anjozorobe in east-central Madagascar (former Antananarivo province). The four-toed rice tenrec (Oryzorictes tetradactylus) is a semi-fossorial species that appears to be restricted to the south-central highlands and the flank of the Eastern Escarpment (former Fianarantsoa province). Several chameleons of the genus Calumma are threatened by loss of habitat. Hillenius’ chameleon (C. hilleniusi) is

known from three disjunct localities in central Madagascar (Ankaratra, Andringitra, and Ivohibe in the central-west, with an isolated record as well from near Ambohijanahary in the south-east). The globe-horned chameleon (C. globifer) appears to be restricted to a relatively small area in the central-east highlands, having been reported from Ambohitantely and the Angavo-Anjozorobe corridor (former Antananarivo province). The Andringitra chameleon (C. andringitraense) is known from a number of localities around the Andringitra Massif and at Kalambatritra in south-eastern Madagascar, and may be present as well at Andohahela (former Fianarantsoa and Toliara provinces). The blunt-nosed chameleon (C. tsycorne) is found disjunctly in a few highland areas of southeastern Madagascar (former Fianarantsoa and Toliara provinces). The jewelled chameleon (Furcifer campani) is found patchily in montane savanna areas from Andringitra National Park in the south to Ankaratra in the north (former Antananarivo and Fianarantsoa provinces). The species was formerly collected in large numbers for the international pet trade, but in recent years exports have fallen off considerably. The whitelipped chameleon (F. minor) is confined to a small area of the Central Highlands (former Antananarivo and Fianarantsoa provinces), where it is threatened by loss of its Tapia forest habitat. Pronk’s day gecko (Phelsuma pronki) and the yellowthroated day gecko (P. flavigularis) are both confined to isolated rainforest fragments bordering the Central Highlands in central-eastern Madagascar (former Antananarivo and Toamasina provinces), where they are highly threatened by habitat destruction and overcollection for the international pet trade. Gould’s day gecko (P. gouldi) is known only from a type specimen collected from the Anja Reserve, 13 km south of Ambalavao in south-central Madagascar (former Fianarantoa province), along with a few additional photographs from the same locality. Three species of dwarf gecko (Lygodactylus) are threatened by loss of habitat. Blanc’s dwarf gecko (L. blanci) is confined to Mount Ibity in central Madagascar (former Antananarivo province). The ornate dwarf gecko (L. ornatus) is known only from a single locality in north-eastern Madagascar (former Mahajanga province). Paulian’s dwarf gecko (L. pauliani) is known only from a handful of specimens collected from a small area of central Madagascar (former Fianarantsoa province). The rusty skink (Madascincus macrolepis) is a leaf litterdependent species known from two disjunct localities in eastcentral Madagascar (former Toamasina and Fianarantsoa provinces). The Ankafina brook snake (Pseudoxyrhopus ankafinaensis) is known only from a single specimen collected during the nineteenth century in eastern Madagascar (former Fianarantsoa province). Numerous dedicated searches have failed to find it, and the species is most likely extinct. The Andrangoloaka skeleton frog (Boophis andrangoloaka) is known only from two small, disjunct localities in

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north-central Madagascar (former Antananarivo province). The spinophis skeleton frog (B. spinophis) is known only from two specimens collected from Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province) and from the Ambohitantely Special Reserve in north-central Madagascar (former Antananarivo province). It likely occurs patchily between these two disjunct localities. The bright-eyed skeleton frog (B. rhodoscelis) is found widely but patchily in central-eastern Madagascar (former Mahajanga, Toamasina, and Fianarantsoa provinces). All are threatened by loss of habitat. Cowan’s mantella (Mantella cowanii) is only known for certain from three small, disjunct localities in east-central Madagascar (former Antananarivo and Fianarantsoa provinces), all of which are embedded within a heavily modified agricultural landscape. The species may possibly occur more widely, but this has yet to be confirmed. Vallan’s climbing frog (Anodonthyla vallani) is only reliably known from a single high-elevation locality within the Ambohitantely Special Reserve, north-central Madagascar (former Antananarivo province). The dotted Madagascar frog (Guibemantis punctatus) is confined to the vicinity of the Ambohitantely Special Reserve in north-central Madagascar (former Antananarivo province). The Ambohimitombi Madagascar frog (Mantidactylus ambohimitombi) is known only from a small area of montane grassland in east-central Madagascar (former Antananarivo province). Delorme’s Madagascar frog (M. delormei) is known only from the Andringitra Massif and from the Maharira Forest in Ranomafana National Park in southeastern Madagascar (former Fianarantsoa province). The Alaotran grainy frog (Gephyromantis mafy) is confined to a single forest fragment near Lake Alaotra in centraleastern Madagascar (former Toamasina province). Helen’s diamond frog (Anilany helenae) is known only from two small forest fragments in the vicinity of Ambohitantely, in central Madagascar (former Antananarivo province). The Eastern Escarpment is located along the eastern edge of the Central Highlands in central-eastern and south-eastern Madagascar (former Toamasina, Antananarivo, and Fianarantsoa provinces). The Betsileo woolly lemur (Avahi betsileo) is known only from the Bemosary Classified Forest in central-eastern Madagascar (former Toamasina, Antananarivo, and Fianarantsoa provinces). The Betsileo sportive lemur (Lepilemur betsileo) is known only from a small area of central-eastern Madagascar (former Toamasina, Antananarivo, and Fianarantsoa provinces). The small-toothed sportive lemur (L. microdon) is known for certain only from a small area of south-eastern Madagascar (former Fianarantsoa province). Both are threatened by habitat destruction and hunting. Groves’ dwarf lemur (Cheirogaleus grovesi) is known only from a few specimens collected within Ranomafana and

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Andringitra national parks (former Toamasina, Antananarivo, and Fianarantsoa provinces). The rufous mouse lemur (Microcebus rufus) is confined to central-eastern and south-eastern Madagascar (former Toamasina, Antananarivo, and Fianarantsoa provinces), roughly from Andringitra National Park to Ranamafana National Park. The Tarzan chameleon (Calumma tarzan) is known only from two small patches of degraded forest in central-eastern Madagascar (former Antananarivo province). Glaw’s chameleon (C. glawi) is known from a few localities in south-eastern Madagascar (former Toamasina and Antananarivo provinces). Lamberton’s leaf chameleon (Brookesia lambertoni) is known only from a small area of central-eastern Madagascar (former Toamasina province). The cork bark leaf-tailed gecko (Uroplatus pietschmanni) has been reported from a small area of central-eastern Madagascar (former Toamasaina province) that includes Ambatovy and Analamay. The delightful brook snake (Pseudoxyrhopus oblectator) is known only from the forest of Ranomafana, Anjozorobe, and Ambatovy in central-eastern Madagascar (former Toamasina and Fianarantsoa provinces). The Betsileo cophyline tree frog (Cophyla cowanii) is known only from a few specimens collected during the nineteenth century from an undefined locality in east-central Madagascar (former Toamasina and Fianarantsoa provinces). Emile’s climbing frog (Anodonthyla emilei) is known only from two localities within Ranomafana National Park in south-eastern Madagascar (former Toamasina province). The Anamalazoatra climbing frog (A. pollicaris) is known only from a few localities in central-eastern Madagascar (former Toamasina province). Several species of skeleton frog (Boophis) endemic to the Eastern Escarpment are threatened by loss of habitat. The Mandraka skeleton frog (B. mandraka) is known for certain only from a single locality in central-eastern Madagascar (former Toamasina province). The peppered skeleton frog (B. piperatus), Schuboe’s skeleton frog (B. schuboeae), Boppa’s skeleton frog (B. boppa), and Narin’s skeleton frog (B. narinsi) are all known only from the region of Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). Liam’s skeleton frog (Boophis liami) is known only from Vohidrazana and near Andasibe in central-eastern Madagascar (former Toamasina province), although it may occur more widely. Burger’s skeleton frog (B. burgeri) is known only from a few specimens collected from a small area around Andasibe in east-central Madagascar (former Toamasina province). Lilian’s skeleton frog (B. lilianae) is known only with certainty from a few adults and tadpoles collected in southern east-central Madagascar (former Toamasina province). Pop’s skeleton frog (B. popi) and Major’s skeleton frog (B. majori) are both found widely but patchily in eastern-central Madagascar (former Toamasina and Fianarantsoa provinces).

Madagascar

Several species of mantella (Mantella) endemic to the Eastern Escarpment are threatened by loss of habitat and collection for the international pet trade. The black-eared mantella (M. milotympanum) was previously known only from a number of fragmented localities in east-central Madagascar covering a small area south of Fierenana (former Toamasina province). It has since been discovered in a few areas to the south-east along the Ivohitra River and around Ranomainty. The golden mantella (M. aurantiaca) has a very restricted distribution in east-central Madagascar (former Toamasina province) centered on the Torotorofotsy area and the Andromena Forest, with populations occurring in several other small forest fragments north and south of Moramanga. The eastern yellow mantella (M. crocea) is known only from a few scattered localities in east-central Madagascar (former Toamasina province). The painted mantella (M. madagascariensis) is known only from a few localities within a relatively wide area of east-central Madagascar (former Toamasina and Fianarantsoa provinces). The white-moustached Madagascar frog (Mantidactylus albofrenatus) and the Andasibe Madagascar frog (M. zolitschka) are each known only from a small area of eastcentral Madagascar (former Toamasina province). The Ranomafana Madagascar frog (M. paidroa) is known only from the region of Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). All are threatened by loss of habitat. The Kibomena stump-toed frog (Stumpffia kibomena) is known only from three localities within a small area of centraleastern Madagascar (former Toamasina province). Runewsweek’s grainy frog (Gephyromantis runewsweeki) is only reliably known from near the summit of Mount Maharira, within Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). The Thelen grainy frog (G. thelenae) and Eiselt’s grainy frog (G. eiselti) are each known only from a small area in central-eastern Madagascar (former Toamasina province). The horned grainy frog (G. cornutus) is known from two small, disjunct localities in central-eastern Madagascar (former Toamasina province). The Betsileo digging frog (Plethodontohyla brevipes) is known only from a few individuals collected at Betsileo and Ranomafana in east-central Madagascar (former Toamasina and Fianarantsoa provinces). The Ankaratra Range (also known as the Ankaratra Massif ) is an extinct/dormant volcanic range located in eastcentral Madagascar (former Antananarivo province). The highest peak in Ankaratra is Tsiafajavona, at 2644 m. Some trace of activity remains in the southernmost part of the range, creating hot springs in the area around Antsirabe. The Ankaratra dwarf gecko (Lygodactylus mirabilis) is confined to montane grassland within the Ankaratra Range. Williams’ skeleton frog (Boophis williamsi) is known from streams at between 2100 and 2381 m on Mount Tsiafajavona, an isolated peak in the Ankaratra Massif and the second highest in Madagascar. One of the most threatened

amphibians in Madagascar, it lives in a severely degraded area where its habitat is under constant threat from fire, illegal logging, overgrazing by livestock, and expanding potato farming. Paulian’s Madagascar frog (Mantidactylus pauliani) is known only from a small area of the Ankaratra Range. The Andringitra Massif is located in south-eastern Madagascar (former Fianarantsoa province). It is protected within Andringitra National Park. The Andringitra dwarf gecko (Lygodactylus intermedius) is confined to rocky heathland on the Andringitra Massif. The Andringitra skeleton frog (Boophis laurenti) is confined to montane heathland and some higher-elevation forest within Andringitra National Park. The Andringitra Madagascar frog (Mantidactylus madecassus) and Bourgat’s Madagascar frog (M. bourgati) are both confined to the higher elevations of Andringitra. The mountain climbing frog (Anodonthyla montana) is known only from a single locality in Andringitra, between 1900 and 2650 m. The Anosy Chain (Chaînes Anosyennes in French) is located in south-eastern Madagascar (former Fianarantsoa and Toliara provinces). Capuron’s chameleon (Calumma capuroni) is known only from Andohahela National Park. The striped brook snake (Pseudoxyrhopus sokosoko) is found only in far south-eastern Madagascar (former Toliara province), with all known specimens having been found in the lower-elevation forests of the Anosy and Vohimena Mountains. The Andohahela skeleton frog (Boophis miadana) is known only from a single locality within Andohahela National Park (former Toliara province). The Andohahela climbing frog (Anodonthyla jeanbai) is known only from a single locality within Andohahela National Park. The Anosy climbing frog (A. rouxae) has a very small range close to the summit in the Anosy Mountains in south-eastern Madagascar, as well as in Andohahela National Park. The Andohahela stream frog (Spinomantis microtis) is confined to the Anosy Chain. The Isalo Massif is located in south-central Madagascar (former Fianarantsoa and Toliara provinces). It is protected within Isalo National Park. The Isalo serotine bat (Neoromicia malagasyensis) is confined to the Isalo Massif region. Nora Lott’s Madagascar frog (Mantidactylus noralottae) is currently known only from the vicinity of Isalo National Park, where it lives on canyon walls and beside rock pools. The blue-legged mantella (Mantella expectata) is confined to a small area centered on Isalo National Park. The Isalo grainy frog (Gephyromantis azzurrae) is known only from three localities on the Isalo Massif. Gottlebe’s rain frog (Scaphiophryne gottlebei) is known only from a few localities in the Isalo Massif area.

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Lowland Rainforests Lowland rainforests occupy a relatively narrow strip between the eastern coast and the mountainous Central Highlands, from sea level to 800 m elevation. They extend from Marojejy in the north-east to the south-east corner of the island. At the northern edge around Vohemar, these moist forests transition to the dry deciduous forest characteristic of the western and far northern coast, and to higher-elevation subhumid forests to the east. The southern end lies at the crest of the Anosy Mountains, where a narrow belt of dry forest marks the transition to the spiny thickets beyond. While the lowland rainforests have been preserved generally better than have the original woodlands of the high Central Plateau, there has still been considerable loss. Although it has nearly been destroyed, the moist coastal forest of Madagascar may also be included in the lowland rainforest category. Taken together, the lowland rainforests represent a great reservoir of diversity and endemism. Nearly all of Madagascar’s endemic mammal genera are represented here, including all five families of lemurs. Three species of woolly lemur (Avahi) from lowland rainforest areas are threatened by loss of habitat and subsistence hunting. The Masoala woolly lemur (A. mooreorum) is known only from Masoala National Park in north-eastern Madagascar (former Antsiranana province). The Manombo woolly lemur (A. ramanantsoavanai) is confined to a few areas of southeastern Madagascar (former Toliara and Fianarantsoa provinces). The southern woolly lemur (A. meridionalis) is confined to a small area of lowland rainforest and littoral forest in the south-eastern corner of Madagascar (former Toliara province). Several species of sportive lemur (Lepilemur) are found in lowland rainforest areas, where they are threatened by habitat destruction and subsistence hunting. The Masoala sportive lemur (L. scottorum) is known only from Masoala National Park in north-eastern Madagascar (former Antsiranana province). The Mananara-Nord sportive lemur (L. hollandorum) is confined to a small area of coastal north-eastern Madagascar (former Toamasina province) centered on the Mananara-Nord Biosphere Reserve. The Manombo sportive lemur (L. jamesorum) is confined to the region of the Manombo Special Reserve in south-eastern Madagascar (former Fianarantsoa province) where the total population in 2010 was estimated at around 1400 and declining. Madame Fleurette’s sportive lemur (L. fleuretae) is confined to three small rainforest fragments in the south-eastern corner of Madagascar (former Toliara province). In recent years a great many species of mouse lemur (Microcebus) have been described throughout Madagascar, each confined to a relatively small area where they are often threatened by habitat destruction and hunting. MacArthur’s mouse lemur (M. macarthurii) is known only from Makira National Park and the island of Nosy Mangabe in northeastern Madagascar (former Toamasina province). Simmons’

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mouse lemur (M. simmonsi) is known only from the Betampona, Zahamena, and Tampolo regions of north-eastern Madagascar (former Toamasina province). The Sahafina mouse lemur (M. gerpi) is known only from the Sahafina Forest, near Mantadia National Park in central-eastern coastal Madagascar (former Toamasina province). The Marohita mouse lemur (M. marohita) is currently known only from the Marohita Forest in central-eastern coastal Madagascar (former Toamasina province). Jolly’s mouse lemur (M. jollyae) is confined to the Mananjary and Kianjavato regions of south-eastern Madagascar (former Fianarantsoa province). The brown-tailed vontsira (Salanoia concolor) is a rare mongoose-like species confined to north-eastern Madagascar (former Antsiranana and Toamasina provinces). The dryad shrew tenrec (Microgale dryas) is only known from a few specimens collected in north-eastern Madagascar (former Toamasina province). Peterson’s long-fingered bat (Miniopterus petersoni) appears to be restricted to the lowland rainforests of the south-east corner of Madagascar (former Toliara province). Bernier’s vanga (Oriolia bernieri) is a shrike-like bird that is rare and patchily distributed throughout its fairly wide range in eastern Madagascar. The red-tailed newtonia (Newtonia fanovanae) is a type of passerine bird that was long known only from a single specimen collected in 1931 near Fanoyana, in east-central Madagascar. It was almost simultaneously rediscovered in Andohahela National Park in 1989 and in the Ambatovaky Special Reserve in 1990 and is now known from a number of low-elevation rainforest blocks, but is everywhere rare. The dusky tetraka (Crossleyia tenebrosa) is a rare type of warbler confined to eastern and north-eastern Madagascar (former Antsiranana and Toamasina provinces). The scaly ground-roller (Geobiastes squamiger) is found throughout the lowland rainforests of eastern Madagascar, but is everywhere declining due to loss of habitat. The brown mesite (Mesitornis unicolor) is a grounddwelling bird with a thin and patchy distribution down the length of the eastern rainforests from Marojejy and the Masoala Peninsula as far south as Taolañaro (former Fort Dauphin). It is everywhere uncommon and rarely seen. Several chameleons of the genus Calumma are threatened by habitat destruction. The Vatosoa chameleon (C. vatosoa) and Vences’ chameleon (C. vencesi) are both known only from a small area of north-eastern Madagascar (former Antsiranana province). The Zahamena chameleon (C. furcifer) is known only from a few localities in central-eastern Madagascar (former Toamasina province). The lance-nosed chameleon (C. gallus) is found patchily in central-eastern Madagascar (former Toamasina province). The hooded chameleon (C. cucullatum) is found widely but patchily over a relatively wide area of north-eastern and eastern Madagascar. The Zahamena ground gecko (Paroedura masobe) is restricted to low-elevation primary rainforest in the north of the Zahamena–Ankeniheny corridor, in central-eastern

Madagascar

Madagascar (former Toamasina province). It is rarely encountered and difficult to find. Spannring’s leaf-toed gecko (Matoatoa spannringi) was long known only from a few microhabitats at Fiadanana in south-eastern Madagascar (former Fianarantsoa province), although more recently a single specimen was recorded from Sainte Luce in the far south-east (former Toliara province). Boettger’s dwarf gecko (Lygodactylus madagascariensis) is found patchily in north-eastern Madagascar (former Antsiranana province). The tiny-scaled dwarf gecko (L. bivittis) is found in eastern Madagascar (former Antsiranana and Toamasina provinces). The Roavolana dwarf gecko (L. roavolana) is found patchily in the south-eastern corner of Madagascar (former Toliara province). All are threatened by loss of habitat. Gabriellà’s gecko (Paragehyra gabriellae) is known from a few localities in the south-eastern corner of Madagascar (former Toliara province). The Tsararano Forest skink (Paracontias tsararano) is known only from a single locality in north-eastern Madagascar (former Antsiranana province), but may be more widespread. The banded skink (Brachyseps spilostichus) is known only from the Tsararano Forest in north-eastern Madagascar (former Antsiranana province). The Masoala skink (Flexiseps stylus) is known only from the Masoala Peninsula in north-eastern Madagascar (former Antsiranana province), where it is threatened by loss of habitat. The Mandady skink (Brachyseps mandady) is known only from the Masoala Peninsula in north-eastern Madagascar (former Antsiranana province), where it is threatened by loss of habitat. The dwarf skink (Madascincus nanus) is known only from north-eastern Madagascar (former Mahajanga province). Three species of plated lizard (Zonasaurus) are threatened by loss of habitat. Boettger’s plated lizard (Zonosaurus subunicolor) is known from two disjunct mainland localities in north-eastern Madagascar along with a few offshore islands (former Antsiranana province). The giant plated lizard (Z. maximus) and the Anelanelany plated lizard (Z. anelanelany) are both known only from a few localities in south-eastern Madagascar (former Fianarantsoa and Toliara provinces). The Iaraka tree snake (Phisalixella iarakaensis) is known only from a single specimen collected at Iaraka in northeastern Madagascar (former Antsiranana province). The Zeny forest snake (Compsophis zeny) is known only from a few specimens collected over a relatively wide area of south-eastern Madagascar (former Fianarantsoa and Toliara provinces). Vincke’s forest snake (C. vinckei) is known from two specimens, the first collected at Andasibe and the second at An’ala, in east-central Madagascar (former Toamasina province). Blanc’s night snake (Ithycyphus blanci) is a venomous species known only from a single specimen collected from lowland rainforest within Marojejy National Park, northeastern Madagascar (former Antsiranana province). Most of

this habitat has since been destroyed as a result of rosewood logging. The small brook snake (Pseudoxyrhopus kely) is a naturally rare species known with certainty only from five localities in the south-eastern corner of Madagascar (former Toliara province). Specimens have also been recorded from the dry forests of Bemaraha in western Madagascar, but will almost certainly prove to be a separate taxon. The Andasibe blind snake (Madatyphlops andasibensis) is known only from Andasibe in east-central Madagascar (former Toamasina province). Domergue’s blind snake (M. domerguei) and Rajery’s blind snake (M. rajeryi) are both known only from Ranomafana National Park in east-central Madagascar (former Fianarantsoa province). Given their cryptic lifestyle some or all of these species may prove to be more widespread than is currently understood. Theo’s climbing frog (Anodonthyla theoi) is known only from a two small patches of lowland rainforest in the Manombo Special Reserve of coastal south-eastern Madagascar (former Fianarantsoa province). Hutchison’s climbing frog (A. hutchisoni) is known only from two localities within the Masoala Peninsula of northern Madagascar (former Antsiranana and Toamasina provinces). The blackthroated climbing frog (A. nigrigularis) is confined to a small area of extreme south-eastern Madagascar (former Toliara province). All are threatened by loss of habitat. Several species of skeleton frog (Boophis) are threatened by loss of their lowland forest habitat. Fay’s skeleton frog (B. fayi) is known only from a few localities in northern Madagascar (former Toamasina province). Englaender’s skeleton frog (B. englaenderi) is known only from a few localities in northern Madagascar (former Antsiranana and Toamasina provinces). The Andasibe skeleton frog (B. feonnyala) is known only from a small area of central-eastern Madagascar (former Toamasina province). The secret skeleton frog (B. arcanus) is known only from a few localities within a small area of south-eastern Madagascar (former Fianarantsoa province). The Vohidrazana skeleton frog (B. solomaso) is known for certain only from the Vohidrazana Forest and nearby Analambalo in central-eastern Madagascar (former Toamasina province), but may also occur within the Betampona Reserve and in other localities between. The tiny skeleton frog (B. miniatus) is known only from a few localities within a relatively wide area of south-eastern Madagascar (former Fianarantsoa and Toliara provinces). The Toliara skeleton frog (B. haematopu) is known only from a few localities within a small area of extreme south-eastern Madagascar (former Toliara province). The Tiarano web-footed frog (Paradoxophyla tiarano) is a semi-aquatic species that to date has only been collected from one area on the Masoala Peninsula in north-eastern Madagascar (former Antsiranana province), but is suspected to occur more widely. The Savaka diamond frog (Rhombophryne savaka) is confined to lowland rainforest within Marojejy National Park in north-eastern Madagascar (former Antsiranana province).

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The Ranomafana stump-toed frog (Stumpffia miery) is confined to a small area of south-eastern Madagascar (former Fianarantsoa province). Bernhard’s mantella (Mantella bernhardi) is known only from a few localities within a relatively wide area of southeastern Madagascar (former Fianarantsoa province). The Antsiranana mantella (M. manery) is known from a few scattered localities in northern Madagascar (former Antsiranana province). Harald Meier’s mantella (M. haraldmeieri) is known from a few localities in extreme south-eastern Madagascar (former Toliara province). All are threatened by loss of habitat and collection for the international pet trade. The Anosy stream frog (Spinomantis brunae) is known from Andohahela National Park and Manantantely in far south-eastern Madagascar (former Toliara province). The stream grainy frog (Gephyromantis rivicola) is known from a few localities over a relatively wide area of northern Madagascar (former Antsiranana and Toamasina provinces). Hintelmann’s grainy frog (G. hintelmannae) is confined to a small area of coastal south-eastern Madagascar (former Toamasina province). Both are threatened by loss of habitat. Watterson’s Madagascar frog (Guibemantis wattersoni) is confined to a few localities in extreme south-eastern Madagascar (former Toliara province). The white-flanked Madagascar frog (G. tasifotsy) is known from a few localities over a relatively wide area of south-eastern Madagascar (former Fianarantsoa province). Kathrin’s Madagascar frog (G. kathrinae) is known only from two small, highly disjunct localities in northern and central-eastern Madagascar. It is not known if it occurs in the intervening areas or if the two populations instead represent distinct species. All are threatened by loss of habitat. The three-banded Madagascar frog (Mantidactylus tricinctus) is known only from a few localities in south-eastern Madagascar (former Toliara and Fianarantsoa provinces).

Littoral Forests and Scrub The moist coastal forests of eastern Madagascar are among the most threatened ecoregions in the country. Thomas’ dwarf lemur (Cheirogaleus thomasi) is confined to littoral forest fragments along the south-eastern corner of Madagascar (former Toliara province). The Vohibola chameleon (Calumma vohibola) is confined to littoral forest fragments between Ivoloina and Vohibola in eastern Madagascar (former Toamasina province). Several species of day gecko (Phelsuma) are threatened by loss of habitat. The flat-tailed day gecko (P. serraticauda) is known from a few localities in coastal eastern and northeastern Madagascar (former Antsiranana and Toamasina provinces). The Masoala day gecko (P. masohoala) is known only from three specimens collected at Cap Est on the Masoala Peninsula of north-eastern Madagascar (former Antsiranana province). Extensive searches have failed to record it at any other sites. The Ampitambe day gecko (P. kely) is known only from the area around Lake Ampitambe in east-central coastal

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Madagascar (former Toamasina province). Hoesch’s day gecko (P. hoeschi) has so far only been recorded from Brickaville (Ampasimanaolotra) and Anosibe An’Ala in eastcentral coastal Madagascar (former Toamasina province). Additional subpopulations are likely to occur between Ampasimanolotra and Vatomandry. The southern day gecko (P. antanosy) is known from a few coastal forest fragments in south-eastern coastal Madagascar (former Toliara province), specifically Ambatotsirongorongo and Sainte Luce (former Toliara province). A third population, at Petriky, was extirpated before 1994 as a result of habitat destruction. The rock skink (Flexiseps decaryi) was long known only from a single locality, but has since been reported from a few other areas on the south-eastern coast of Madagascar (former Toliara province). The Manombo Madagascar frog (Guibemantis diphonus) is confined to the Manombo Special Reserve in coastal southeastern Madagascar (former Fianarantsoa province). The ringwearing Madagascar frog (G. annulatus) is confined to three isolated pockets of low-elevation littoral forest (Mandena, Saint Luce, and the Agnalaro Forest) in south-eastern Madagascar (former Toliara province), where it is highly dependent upon stands of Pandanus plants. The southern jumping frog (Aglyptodactylus australis) is known only from a small area of extreme south-eastern Madagascar (former Toliara province), where it is threatened by loss of habitat.

Lowland Dry Deciduous Forests The tropical dry deciduous forests that are to be found in Madagascar are among the richest and most distinctive of their kind in the world, with high numbers of plant and animal species. There are two separate areas within this bioregion: the western side of Madagascar from the Ampasindava Peninsula in the north-east to Belo-sur-Tsiribihina and Maromandia in the central-west; and the far northern tip of the island (apart from the high elevations of Montagne d’Ambre). Together, these dry deciduous forests span the coastal plain with its limestone plateaus (known as tsingy) emanating virtually at sea level to altitudes of roughly 600 m. The area also includes wetlands and grasslands (the latter mostly created by forest clearance for agriculture). The dry forests themselves are characterized by a deciduous canopy extending to a height of 10–15 m. Rainfall is more abundant than in the succulent woodlands and spiny thickets to the south, but lower by far than in the eastern lowland rainforests. The trees have adapted to the dry climate by shedding leaves in the winter months to limit fluid loss. Some, such as the baobabs, have adapted by evolving the ability to store copious amounts of water in their large, bulbous trunks. While human population densities are lower in these regions, the dry deciduous forests have nevertheless been largely cleared for slash-and-burn agriculture, pasture, firewood, and logging. Lorenz Von Liburnau’s woolly lemur (Avahi occidentalis) is found in two disjunct areas of north-western Madagascar

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(former Mahajanga province) where it is threatened by habitat destruction and hunting. Several dry forest species of sportive lemur (Lepilemur) are threatened by habitat destruction and hunting. The Sahafary sportive lemur (L. septentrionalis) is arguably the most endangered lemur species of them all. Only about 50 individuals survive in a few small patches of forest north of the Irodo River in far north-eastern Madagascar (former Antsiranana province), near the villages of Madirobe and Ankarongana. Gray’s sportive lemur (L. dorsalis) is confined to the Sambirano region of north-eastern Madagascar (former Mahajanga province). Mittermeier’s sportive lemur (L. mittermeieri) is confined to the Ampasindava Peninsula in northeastern Madagascar (former Antsiranana province). MilneEdwards’ sportive lemur (L. edwardsi), Antafia sportive lemur (L. aeeclis), Grewcock’s sportive lemur (L. grewcockorum), Ahmanson’s sportive lemur (L. ahmansonorum), and Otto’s sportive lemur (L. otto) are all confined to small areas of coastal north-western Madagascar (former Mahajanga province). The Bemaraha sportive lemur (L. randrianasoloi) is confined to a small area of central-western coastal Madagascar (former Mahajanga province). The red-tailed sportive lemur (L. ruficaudatus) is found patchily in central-western coastal Madagascar (former Toliara province). The pale fork-marked lemur (Phaner pallescens) is patchily but widely distributed along a narrow strip of western Madagascar (former Mahajunga and Toliara provinces). The northerly populations appear isolated and may possibly represent distinct taxa. The Sambirano fork-marked lemur (P. parienti) is confined to the Sambirano region of north-eastern Madagascar (former Antiranana and Mahajanga provinces), specifically the Ampasindava Peninsula south to the Andranomalaza River. Both are threatened by loss of habitat. Coquerel’s giant mouse lemur (Mirza coquereli) is found patchily throughout western Madagascar (former Mahajunga and Toliara provinces), where it is threatened by habitat destruction. Owing to their small size the mouse lemurs (Microcebus) of the western dry forests are less vulnerable to hunting pressure, although habitat destruction is a serious threat. The Bongolava mouse lemur (M. bongolavensis) is known only from three small dry forest fragments in the area around PortBergé in north-western Madagascar (former Mahajanga province). The Ambarijeby mouse lemur (M. danfossi) is only known from a few dry forest fragments between the Sofia and Maevarano rivers in north-western Madagascar (former Mahajanga province). The golden-brown mouse lemur (M. ravelobensis) occurs in the forests of Ankarafantsika as well as in the Bongolava and Mariarano Classified Forests in northwestern Madagascar (former Mahajanga province). Madame Berthe’s mouse lemur (M. berthae), the world’s smallest primate, is confined to the Menabe region south of the Tsiribihina River in western coastal Madagascar (former Toliara province). The total range is quite small and highly fragmented.

The western falanouc (Eupleres major) is a mongoose-like mammal that occurs patchily in north-western Madagascar (former Antsiranana and Mahajanga provinces) from near Baly Bay National Park north through the Ankarafantsika and Analalava forests to the Sambirano, east to the foothills of Tsaratanana, the Sahamalaza Peninsula, and possibly as far as the lower elevations of Montagne d’Ambre. It is rarely seen and presumably rare. Jenkin’s shrew tenrec (Microgale jenkinsae) is known only from two specimens collected in the Mikea Forest of southwestern Madagascar (former Toliara province). The giant jumping rat (Hypogeomys antimena) is a rabbitsized, kangaroo-like species restricted to a small area of coastal western Madagascar (former Toliara province) from the Tomitsy River north to the Tsiribihina River. Habitat destruction, hunting and introduced species threaten its survival. The lowland red forest rat (Nesomys lambertoni) is a littleknown species confined to a small area of dry forest in western Madagascar (former Mahajanga province). The greater big-footed mouse (Macrotarsomys ingens) is a nocturnal species known only from the vicinity of Ankarafantsika National Park in north-western Madagascar (former Mahajanga province), where it is threatened by feral cats and dogs as well as by wildfires. Petter’s big-footed mouse (M. petteri) is known only from a single specimen collected in 2003 in the Mikea Forest of south-western coastal Madagascar (former Toliara province). Grandidier’s trident bat (Paratriaenops auritus) is an obligate cave-roosting species with a restricted range in the northeastern tip of the island (former Antsiranana province). Van Dam’s vanga (Xenopirostris damii) is a rare shrikelike bird currently known only from two small, disjunct areas in north-western and far northern Madagascar (Ankarafantsika and Analamera), where it is under threat by fire and other pressures. However, there is still much suitable habitat between the two sites that have yet to be surveyed. The flat-tailed tortoise or kapidolo (Pyxis planicauda) is restricted to dry forest fragments within a small area of western coastal Madagascar (former Toliara province). The ploughshare tortoise or angonoka (Astrochelys yniphora), one of the world’s rarest tortoises, is confined to two localities around Baly Bay in north-western Madagascar (former Mahajanga province) which have recently been declared a national park. Nevertheless, the species may soon be extinct in the wild due to illegal collection for use in the international pet trade. It is well established in captivity. The rhinoceros chameleon (Furcifer rhinoceratus) is known only from a few localities in north-western coastal Madagascar (former Mahajanga province). The Antimena chameleon (Furcifer antimena) is known only from a small area of south-western coastal Madagascar (former Toliara province), but may be more widespread. Both are threatened by loss of habitat. Decary’s leaf chameleon (Brookesia decaryi) and the toothed leaf chameleon (B. dentata) are both known only

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from Ankarafantsika National Park in north-western Madagascar (former Mahajanga province). The Bemaraha day gecko (Phelsuma borai) is known for certain only from a single specimen collected from the Bemaraha region of central-western Madagascar (former Mahajanga province). However, published photos indicate that the species may occur to the north at Ankarafantsika, suggesting that it may occur in intervening areas as well. Van Heygen’s day gecko (P. vanheygeni) is known only from three localities on the Ampasindava Peninsula of northern Madagascar (former Antsiranana province). The yellowheaded day gecko (P. klemmeri) is known only from the Ampasindava Peninsula in northern Madagascar and from an area of dry forest near Mandrozo Lake some 600 km to the south. Surveys between these two localities as well as on coastal islands have failed to detect this highly conspicuous species, and the two populations are likely to be genuinely isolated from one another. Henkel’s leaf-tailed gecko (Uroplatus henkeli) and Günther’s leaf-tailed gecko (U. guentheri) are both found widely but patchily in northern and western Madagascar (former Antsiranana, Mahajanga, and Toliara provinces). The Vazimba Madagascar ground gecko (Paroedura vazimba) is known from Ankarafantsika, Anjiamangirana, and Bora in north-western Madagascar (former Mahajanga province). The Ambonihazo velvet gecko (Blaesodactylus ambonihazo) is known only from a few specimens collected from Ankarafantsika National Park in north-western Madagascar (former Mahajanga province). Boettger’s skink (Voeltzkowia mira) is confined to a few localities in north-western Madagascar (former Mahajanga province). The Tavaratra mabuya skink (Trachylepis tavaratra) is confined to far northern Madagascar (former Antsiranana province). Yamagishi’s mermaid skink (Sirenoscincus yamagishi), sonamed for its lack of hindlimbs, is known from Belambo and Ankarafantsika in north-western Madagascar (former Mahajanga province). The Menamainty skink (Pseudoacontias menamainty) is known only from a single specimen collected in north-western coastal Madagascar (former Mahajanga province). The yellow skink (Flexiseps ardouini) is confined to far northern Madagascar (former Antsiranana province). Petter’s short skink (Pygomeles petteri) is a burrowing species confined to the lowland dry forests of Ankarafantsika National Park in north-western Madagascar (former Mahajanga province). The variable tree snake (Phisalixella variabilis) appears to be restricted to a few localities in far north-eastern Madagascar (former Antsiranana province), where it is relatively rare. Marta’s water snake (Thamnosophis martae) is confined to lowland dry forests in the vicinity of Montagne des Français and Forêt d’Ambre in far north-eastern Madagascar (former Antsiranana province).

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The Maintikibo snake (Liophidium maintikibo) is known only from two specimens collected from the Kirindy Forest and the Andranomena Special Reserve in western coastal Madagascar (former Toliara province). Therezien’s snake (L. therezieni) is known only from a few specimens collected from widely spaced localities in north-eastern and north-western Madagascar (former Antsiranana and Mahajanga provinces). The Berara cophyline tree frog (Cophyla berara) is known only from a few areas of the Sahamalaza Peninsula in northeastern Madagascar (former Antsiranana province). The Madinika Madagascar frog (Wakea madinika) is known only from specimens collected in 2001 from a temporary pond on a cacao plantation in north-eastern Madagascar (former Antsiranana province). It may occur more widely in surrounding areas, but extensive surveys have failed to locate it. The Fôret d’Ambre skeleton frog (Boophis baetkei) is confined to a single locality on the eastern edge of the Fôret d’Ambre Special Reserve in far northern Madagascar (former Antsiranana province). The Berara skeleton frog (B. tsilomaro) appears to be restricted to in and around Berara Forest in north-eastern Madagascar (former Anstiranana province). The Ankarafa skeleton frog (B. ankarafensis) is confined to a single forest fragment on the Sahamalaza Peninsula, north-western Madagascar (former Antsiranana province). The Antsohihy stump-toed frog (Stumpffia analamaina) is known only from a small fragment of dry deciduous forest along the road between Antosohihy and Mandritsara in northwestern Madagascar (former Mahajanga province). It may be more widespread within a small range of suitable habitat north and north-east of Antsohihy. The dry forest jumping frog (Aglyptodactylus laticeps) is well known from a few widely separated localities in western Madagascar (former Mahajanga province), where it can be locally abundant but is thought to be declining overall by an increasing lack of suitable habitat.

Tsingy Forest Tsingy forests are dry deciduous forest areas located on karstic limestone massifs and plateaus rising from sea level upwards to around 600 m. The tsingy tuft-tailed rat (Eliurus antsingy) is known only from a few specimens collected in disjunct areas of limestone tsingy forests and other formations in western Madagascar (former Mahajanga province). The Maramaintso plated lizard (Zonosaurus maramaintso) is known only from a single undefined locality in western Madagascar (though most likely the Tsingy de Bemaraha region). The Tanjaka Madagascar ground gecko (Paroedura tanjaka) is found in a few rocky forested localities in western Madagascar (former Mahajnga province) at Namoroka and near Bemaraha. The unexpected tree snake (Lycodryas inopinae) is known only from a few specimens collected from far northern Madagascar (former Antsiranana province).

Madagascar

Montagne des Français is located in far northern Madagascar (former Antsiranana province). The Montagne des Français leaf chameleon (Brookesia tristis) is known only from the Montagne des Français Massif in north-eastern Madagascar (former Antsiranana province). The Lohatsara Madagascar ground gecko (Paroedura lohatsara) is confined to the lower elevations of Montagne des Français in far northern Madagascar (former Anstiranana province). The stout dwarf snake (Heteroliodon fohy) is known only from four specimens collected or observed at Montagne des Français and near the village of Ampombofofo in far northeastern Madagascar (former Antsiranana province). Stafford’s stump-toed frog (Stumpffia staffordi) is confined to a karstic cave and its environs on Montagne des Français. The Ankarana Massif is located in far northern Madagascar (former Antsiranana province). The important Ankarana Special Reserve was created in 1956. Rösler’s day gecko (Phelsuma roesleri) is known only from a few specimens collected from lower elevation dry forests on the Ankarana Massif. Mocquard’s keeled snake (Alluaudina mocquardi) is known only from karst outcrop caves at Ankarana, where it is thought to be threatened by sapphire mining operations. The Ankarana tsingy frog (Tsingymantis antitra) is known only from the Ankarana Special Reserve where it lives in lowland limestone karst formations. The Ankarana stump-toed frog (Stumpffia be) is known only from the eastern side of the Ankarana Special Reserve in far northern Madagascar (former Antsiranana province). Tsingy de Namoroko is located in north-western Madagascar (former Mahajanga province). The Namoroka leaf chameleon (Brookesia bonsi) is confined to an area of Tsingy de Namoroka Strict Nature Reserve. Tsingy de Bemaraha is located in central-western coastal Madagascar (former Mahajanga province). It is protected within Tsingy de Bemaraha National Park. The Bemaraha woolly lemur (Avahi cleesei) is known only from the Tsingy de Bemaraha region. Nicosia’s chameleon (Furcifer nicosiai) is known only from the Tsingy de Bemaraha region. The Antsingy leaf chameleon (Brookesia perarmata) and the dwarf leaf chameleon (B. exarmata) are both known only from Tsingy de Bemaraha National Park. The Bemaraha digging frog (Plethodontohyla fonetana) is endemic to the Tsingy de Bemaraha National Park and adjacent Beanka Classified Forest. The Antsingy grainy frog (Gephyromantis atsingy) is confined to Tsingy de Bemaraha.

Semi-Deserts and Arid Shrublands Semi-deserts and shrublands predominate in south-western and southern Madagascar as a result of rainshadow from the Central Highlands.

Petter’s sportive lemur (Lepilemur petteri) is relatively widespread in south-western and southern Madagascar (former Toliara province), but is everywhere threatened by loss of habitat. Standing’s day gecko (Phelsuma standingi) is known only from a few localities in south-western Madagascar (former Toliara province). It is extensively collected for the international pet trade. The Toliara day gecko (P. breviceps) is known only from a few localities in south-western coastal Madagascar (former Toliara province). Mocquard’s leaf-toed gecko (Matoatoa brevipes) is a rare species confined to areas of intact spiny thicket in coastal south-western Madagascar (former Toliara province), where it is known from Ambolisaka near Morombe, Tsivanoa near Belalanda, Ranobe, Mikea, Tsimanampetsotsa, and Cap St. Marie. Decary’s dwarf gecko (Lygodactylus decaryi) is known only from specimens collected from Andrahamana, Angavo, and Behara in southern Madagascar (former Toliara province) between the 1930s and 1950s. Dumas’ mabuya skink (Trachylepis dumasi) is found widely but very patchily throughout south-western and southern coastal Madagascar (former Mahajanga and Toliara provinces). The Andranovaho skink (Flexiseps andranovahensis) is known only from three widely separated localities in coastal south-western Madagascar (former Toliara province). Appert’s snake (Liophidium apperti) is known only from a single specimen collected during the 1980s from south-western Madagascar (former Toliara province). The forest where it was discovered has since been destroyed, and the species may be extinct.

Succulent Spiny Woodlands The succulent spiny woodlands are xeric shrubland region found in south-central and south-western Madagascar (former Toliara and Fianarantsoa provinces). The climate is tropical and dry, with a marked dry season from May to October. To the north this bioregion transitions into dry deciduous forest, while to the south it borders the even drier spiny thickets. Subhumid forests lie to the east. The vegetation is similar to the dry deciduous forest but includes more dry-adapted, xerophytic species. Forests reaching 15 m in height contain a mixture of trees and shrubs, among them two species of baobab and succulents of the genus Pachypodium. This bioregion is threatened by burning, logging, and cattle and goat farming. Hunting is also a problem. Protected areas include the Zombitse-Vohibasia National Park, Kirindy Mitea National Park and Kirindy Forest, and the Andranomena Reserve. Hubbard’s sportive lemur (Lepilemur hubbardi) are both confined to small areas of south-western Madagascar (former Toliara province). The narrow-striped mongoose or bokiboky (Mungotictis decemlineata) is divided into two subspecies. The northern

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narrow-striped mongoose (M. d. decemlineata) inhabits the succulent spiny woodlands of south-western coastal Madagascar (former Toliara province), where it is threatened by loss of habitat, hunting, and predation by feral dogs. The subdesert mesite (Monias benschi) is a rail-like terrestrial bird confined to a narrow coastal strip in south-western Madagascar (former Toliara province). Although still common in suitable habitat, it is threatened by habitat destruction. The long-tailed ground-roller (Uratelornis chimaera) is a largely terrestrial bird restricted to a narrow coastal strip in south-western Madagascar (former Toliara province) between the Fiherenana and Mangoky rivers. Owing to its weak flight it is particularly vulnerable to hunting and predation by feral animals. The Belalanda chameleon (Furcifer belalandaensis) is confined to two small localities near Toliara in south-western Madagascar (former Toliara province). Tuzet’s chameleon (F. tuzetae) is known only from a single specimen collected in 1972 at Andrenalamivola in south-western Madagascar (former Toliara province). It is possibly confined to gallery forest along the Mangoky River. Hielsher’s day gecko (Phelsuma hielscheri) is known only from a few localities in the south-western and south-central Madagascar (former Toliara province). The unadorned tree snake (Lycodryas inornatus) occurs in south-western coastal Madagascar (former Toliara province).

Arid Spiny Bush The arid spiny bush (also known as southern spiny thickets) is a bioregion in coastal southern and south-western Madagascar (former Toliara province). Around 95 per cent of the original flora is endemic, making it one of the most unique places on Earth, and many of these plants show extreme adaptation to the arid environment. The fauna is equally remarkable. As with other areas of Madagascar this habitat is under pressure from human activities such as conversion to grazing land, harvesting for charcoal and firewood, and logging. At present only about 3 per cent is protected in national parks and reserves. The white-footed sportive lemur (Lepilemur leucopus) is confined to a few localities within a small area of southeastern Madagascar. Grandidier’s vontsira (Galidictis grandidieri) is a type of mongoose that was only described to science in 1986 and has a restricted range around Lake Tsimanampetsotsa in southwestern Madagascar. The southern narrow-striped mongoose (Mungotictis decemlineata lineata) is confined to arid spiny bush areas of south-western Madagascar, where it is threatened by loss of habitat, hunting, and predation by feral dogs. The red-shouldered vanga (Calicalicus rufocarpalis) is a shrike-like bird that was originally described from two specimens collected in 1947 near Toliara in south-western Madagascar. It was not seen again until being photographed in 1992 and then in 1997, when a number of individuals were

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seen in the same area. It has since been located in a few other localities, but the total population is thought to be under 1000. The radiated tortoise (Astrochelys radiata) is found primarily in the dry spiny bush of southern and south-western Madagascar, from the area of Amboasary in the south across the Karimbola and Mahafaly plateaus north of Toliara to Morombe. It is usually found in a narrow band within 50–100 km of the coast. Historically this species was quite common, often being found along roadways, although this is no longer the case. However, they may still be locally abundant in certain areas. The main threats area habitat destruction and collection, both for the international pet trade and for local consumption. The Tsimanampetsotsa clawless gecko (Ebenavia maintimainty) is restricted to areas of dry limestone spiny bush in south-western Madagascar. Most of the known population is protected within Tsimanampetsotsa National Park. Grandidier’s Madagascar ground gecko (Paroedura androyensis) is found in scattered localities throughout southern coastal Madagascar, including Ambatotsirongorongo, Andohahela, Antafoky, Cap Sainte Marie, Petriky, Sarodrano, Sept Lacs, Zombitse, and Tsivanoa. The Lavenambato mabuya skink (Trachylepis vezo) is currently known only from a few specimens collected at a single locality near the mouth of the Onilahy River in south-western Madagascar. However, it likely also occurs on the Mahafaly Plateau and elsewhere. The Madagascar three-lined snake (Liophidium trilineatum) is known from very few records from the region of Lake Tsimanampetsotsa, as well as Andranobe and Benonoka in south-western and southern Madagascar.

Lakes, Rivers, and Marshes Madagascar has one of the most distinctive freshwater ecosystems in the world, with many endemic species. From the island’s Central Plateau a number of short, swift rivers flow over a narrow strip of land as they descend to the Indian Ocean to the east. In the west, longer rivers flow down into the Mozambique Channel. Due to climate variability throughout the island, the freshwater systems of the eastern side contrast greatly with those of the west, although they all experience seasonal flooding. Madagascar’s freshwater fishes are ‘living fossils’ and belong to the most primitive of catfish, herrings, cichlids, killifish, silversides, and the like. The Madagascar grebe (Tachybaptus pelzelnii), an inhabitant of small lakes and forested wetlands, ranges across Madagascar and is still reasonably common in a few areas of the Central Highlands and west, although in small numbers. In 2007 the total population was estimated at between 1500 and 2500. Meller’s duck (Anas melleri) is found mainly in the wetlands of the eastern and northern Central Highlands. There are additional, isolated populations on isolated massifs on the western edge of the plateau, and (at least formerly) an

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introduced population on the island of Mauritius that is now most likely extinct. The species has been in decline since the beginning of human colonization and is now rare throughout its range, with the exception of forested areas of the north-west and in the wetlands around Lake Alaotra. The Madagascar pochard (Aythya innotata) is a type of diving duck that was considered to be relatively common at Lake Alaotra in central-eastern Madagascar (former Toamasina province) during the 1930s, but declined dramatically over the subsequent two decades. There were a few unconfirmed sightings thereafter, with the last certain record taking place in 1960. The species was presumed extinct until 1991, when a single male was captured alive, although subsequent searches failed to discover others. The species was once more considered to be extinct until 2006, when it was rediscovered in very small numbers in a small volcanic lake some 330 km north of Lake Alaotra in north-eastern Madagascar (former Mahajanga province). The Madagascar snipe (Gallinago macrodactyla) is found throughout the eastern half of the island but is much more common over 700 m elevation. It is threatened by the increasing conversion of wetland-edge habitat for rice cultivation. The Sakalava rail (Zapornia olivieri) is known from several widely separated wetland areas in lowland western Madagascar, where it is rare and localized. Specimens were taken in 1930 and 1962, and all records since have been field sightings. A single bird was seen in 1995 at Lake Bemamba and another in 1999; two were reported at Amboropotsy Marsh near Bekopaka in 2001; and a small group were observed over the course of several days at Lake Kinkony south-west of Mahajanga in 2003. Other sightings have occurred since at Lake Kinkony, Lake Ampandra, Lake Amparihy, Lake Sahapy, and Lake Mandrozo, all between the Betsiboka and Mangoky rivers. The total population may be no more than 250. The slender-billed flufftail (Sarothrura watersi) is a small, secretive rail of isolated mid-altitude rainforest wetland areas across much of eastern Madagascar. It has been recorded from near Analamazaotra in 1928, near Andapa in 1930, and, in more recent years, from Anjozorobe, Torotorofotsy Marsh, Ranomafana/Vohiparara, Andringitra National Park, and from localities between the latter two areas. The total population is thought to be less than 1000. The Madagascar big-headed turtle (Erymnochelys madagascariensis) inhabits slow-moving rivers, lakes, and swamps in western Madagascar (former Antsiranana, Mahajanga, and Toliara provinces), where it is known to be in serious decline. Its large size makes it an attractive food source for local people. The black diamond cichlid (Paratilapia polleni) is a popular aquarium fish with a restricted range in north-eastern Madagascar (former Antsiranana and Mahajanga provinces), where it is threatened by deforestation and irrigation projects. It is still relatively common on the island of Nosy Be and in a few lakes on the mainland. The Itasy cichlid (Ptychochromoides itasy) was last seen in Lake Itasy, central Madagascar (former Antananarivo

province) in the 1970s, having been extirpated by a combination of habitat degradation and introduced species. It was rediscovered in 2010 in the Sakay River, a tributary of the Tsiribihina River. The Fort Dauphin cichlid (Ptychochromis mainty) is known only from forested streams in extreme south-eastern Madagascar (former Toliara province). The Diana cichlid (P. oligacanthus) is found in northern Madagascar (former Antsiranana and Mahajanga provinces) and in crater lakes on the island of Nosy Be. The Antsirabe Nord cichlid (P. loisellei) is known from the Mahanara River and its tributaries in north-eastern coastal Madagascar (former Antsiranana province), with further subpopulations in Lake Nosiarina in the Bemariovo River drainage and from Lake Mahatsara near Vohemar. All are threatened by overfishing, loss of habitat, and invasive species. Cichlids of the genus Paretroplus are largely restricted to lakes and rivers in north-western Madagascar, where they have declined drastically due to habitat destruction, overfishing, and invasive species. The damba cichlid (P. maculatus) lives in shallow floodplain lakes and small rivers in the Betsiboka and Mahajamba drainages, where it has suffered a severe decline in recent years. The Maromandia cichlid (P. maromandia) is known only from the Andanomalaza and Maevarano River drainages. It was present in Lake Andrapongy until the mid1990s, but has since been extirpated there. The Calico cichlid (P. kieneri) is still relatively widespread in north-western Madagascar but is everywhere rare. Van Dam’s cichlid (P. damii) is also found over a large area of western and far northern Madagascar, but has disappeared from areas where it formerly occurred, as for instance the island of Nosy Be. Petit’s cichlid (P. petiti) is known only from a juvenile type specimen of unknown origin. The Tsimoly cichlid (P. tsimoly) is known from two north-bank tributaries of the Betsiboka (Ankalimilotrabe and Boinakely rivers), with an additional population located in an unnamed lake east of the Kalamilotra River. Similar-looking specimens have also been collected from the upper reaches of the Kamoro River near the village of Tsimoly. Loiselle’s cichlid (P. loisellei) is so far known from the Mahanara River in northern coastal Madagascar (former Antsiranana province), but may be more widespread. Nourissat’s cichlid (P. nourissati) is known from the Amboaboa and Mangarahara rivers in northern Madagascar (former Mahajanga and Toamasina provinces). The Lamenbe cichlid (P. lamenabe) is confined to the lower reaches of the Mahajamba River near the town of Androka and to Lake Tseny in the Sofia River drainage (former Mahajanga province). The elongated grunter (Mesopristes elongata) is known from several rivers along the eastern coast of Madagascar, where it is threatened by overfishing and habitat destruction. Sauvage’s flagtail (Kuhlia sauvagii) is a perch-like species that occurs in a number of rivers draining eastern coastal Madagascar (former Toamasina, Fianarantsoa, and Toliara provinces). Although relatively widespread and common, it is

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vulnerable to deforestation as it prefers relatively pristine forested rivers and streams. The Amboaboa round herring (Sauvagella robusta) is a small fish known from the middle and upper reaches of the Amboaboa, Mangarahara, and Anjombony rivers in northern Madagascar (former Mahajanga province), with additional populations in Lake Tseny and, at least formerly, Lake Sarodrono. The tricolour rainbowfish (Bedotia tricolor) is only known from a single tributary of the Faraony River in south-eastern Madagascar (former Fianarantsoa province), whose watershed has been almost entirely stripped of forest cover. The species must also cope with an introduced predator, the spotted snakehead (Channa punctata). Geay’s rainbowfish (B. geayi) is known only from small forest streams associated with the Mananjary River drainage in south-central Madagascar (former Fianarantsoa province). The white-margined rainbowfish (B. albomarginata) is known only from small forested to moderately degraded tributaries in the upper to middle reaches of the Mananara and Rienana rivers in south-eastern Madagascar (former Fianarantsoa and Toliara provinces). The Rianila rainbowfish (B. leucopteron) is known from the middle reaches of the Iaroka–Rianila system in central-eastern Madagascar (former Toamasina province), but may be present in adjacent drainages. The long-finned rainbowfish (B. longianalis) is confined to a few rivers in north-eastern coastal Madagascar (former Toamasina province) and the island of Nosy Boraha. Pellegrin’s rainbowfish (Rheocles pellegrini) was long known only from a single specimen collected in 1931 from an undefined area of northern Madagascar (former Antsiranana and/or Toamasina provinces). Further specimens have been collected from the same general region within the past couple of decades. Wright’s rainbowfish (R. wrightae) is known from a few rivers in central-eastern coastal Madagascar (former Toamasina province). The Alaotran rainbowfish (R. alaotrensis) appears to be confined to Lake Alaotra, a few smaller surrounding lakes and to the Maningory River in central-eastern Madagascar (former Toamasina province). Waterlot’s silverside (Teramulus waterloti) is known from the Mananjeba and Anjingo rivers in north-western Madagascar (former Antsiranana and Mahajanga rivers). The range is notably fragmented, with numerous localities having been sampled without finding the species. The Masoala goby (Sicyopterus punctissimus) is restricted to a few isolated localities on the Masoala Peninsula of northeastern Madagascar (former Antsiranana province), where it is uncommon. The Pangalana sleeper goby (Eleotris vomerodentata) is known only from a single specimen collected during the 1980s from Pangalana Lagoon in eastern coastal Madagascar (former Toamasina province). The Malagasy mountain mullet (Acentrogobius therezieni) is a small goby found in the Andranomavokely River and the associated tsingy pools and springs in the region of the Namoroka Strict Nature Reserve, north-western Madagascar (former Mahajanga province).

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Blind and pigment-free gobies of the genus Typhleotris are endemic to subterranean habitats (caves, sinkholes, and wells) of south-western coastal Madagascar (former Toliara province), where they are seriously threatened by human disturbance. Petit’s cavefish (T. madagascariensis) is found in the Mahafaly Plateau karst formation, as well as in isolated localities along the coastal plain below and to the west. The Mararybe cavefish (T. mararybe) is confined to a single cave system near the town of Itampolo. Paulian’s cavefish (T. pauliani) inhabits coastal regions to the south of Morombe in the vicinity of Andalambezo. Several killifish of the genus Pachypanchax are threatened by loss of habitat and invasive species. The powder-blue killifish (P. omalonota) is found in the Sambirano River drainage of northern Madagascar (former Antsiranana and Mahajanga provinces) and on the island of Nosy Be. Patricia Yazgi’s killifish (P. patriciae) occurs in the Mananjeba, Mahavavy du Nord, Ifasy, Manehoko, and Ampandra rivers of northern Madagascar (former Antsiranana province). Holly’s killifish (P. sakaramyi) is known from the Sakaramy and Antongombato rivers and possibly a few crater lakes as well in far northern Madagascar (former Antsiranana province). The Varatraza killifish (P. varatraza) is known from the Menambery, Fanambana, and Ampanobe rivers of northern Madagascar (former Antsiranana province), but may be more widespread. Arnoult’s killifish (P. arnoulti) is found patchily in the lakes, rivers, and streams of north-western Madagascar (former Mahajanga province). The Madagascar lampeye (Pantanodon madagascariensis) was originally known from forest streams on the east-central slopes between Mahavelona and Fenoarivo in eastern coastal Madagascar (former Toamasina province). No specimens have been collected since the early 1960s, and the species is now believed to be extinct, a victim of conversion of its preferred swamp habitat into rice fields and competition from introduced Gambusia species.

Lake Alaotra Lake Alaotra (Lac Alaotra in French) is located on the northern central plateau (former Toamasina province). The largest lake in Madagascar, it serves as an important habitat for waterfowl but is increasingly threatened owing to the destruction of the surrounding marshes by burning and conversion to rice fields. A 42,478-ha protected area was created around the lake in 2007. The Alaotran bamboo lemur (Hapalemur alaotrensis) is confined to the papyrus and reed beds in and around the lake. Its unique, semi-aquatic habitat long provided it with protection, but during the mid-twentieth century the population began to decline dramatically due to habitat destruction and hunting by local fishermen (who employ a variety of cruel methods including harpoons, dogs or simply shake the reeds until the animals fall helplessly into their boats). It is also frequently captured for use as pets. By the early 1990s, when an international conservation programme began, the species

Madagascar

Figure 4.2 The only known photograph of an Alaotran grebe in the wild. (Credit: Alaotran grebe Tachybaptus rufolavatus on Lake Alaotra September 1985. BirdLife International, University of London Union and BP Conservation Programme Zahamena, Madagascar Expedition 1985. © Paul Thompson) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

was already on the verge of extinction. Numbers continued to drop in any case, reaching a low of just 2500 in 2013. Durrell’s vontsira (Salanoia durrelli) is a mongoose-like species known only from the marshlands of Lake Alaotra. The Alaotran grebe (Tachybaptus rufolavatus) was a nearly flightless waterbird known mainly from Lake Alaotra, where the species was last recorded in 1985. Thorough surveys since have failed to find any evidence of its existence, and it is now considered to be extinct.

The Mananara du Nord River The Mananara du Nord River is located in north-eastern Madagascar (former Toamasina province). Ernest Magnus’ cichlid (Ptychochromis ernestmagnusi) is known only from a few specimens collected from the Mananara du Nord River. The Mananara du Nord catfish (Gogo atratus) is confined to the Mananara du Nord River.

The Lokoho River The Lokoho River is located in north-eastern Madagascar (former Antsiranana province). The Lokoho rainbowfish (Rheocles vatosoa) is confined to the upper reaches of the Lokoho River. The Manentenina River is an effluent of the Lokoho River within Marojejy National Park. The Manentenina rainbowfish (Bedotia marojejy) is confined to the Manantenina River.

The Sofia River The Sofia River is located in north-western Madagascar (former Mahajanga province), where it rises on the Tsaratanana Massif.

The Mangarahara River is a major tributary of the Sofia River. The Mangarahara cichlid (Ptychochromis insolitus) is known only from the Amboaboa River, (a tributary of the Mangarahara River), near the town of Marotandrano. It was previously also found in the Mangarahara River itself, but seems to have been extirpated there. The species was feared extinct, with only three males surviving in captivity (London Zoo and Berlin Aquarium), until a small number were rediscovered in the Amboaboa in 2013. The latter were taken into captivity in order to establish a breeding programme. The Mandritsara cichlid (Paretroplus gymnopreopercularis) is known only from the Mangarahara River and its tributary, the Amboaboa River. Derham’s rainbowfish (Rheocles derhami) is confined to the Amboaboa and Mangarahara rivers within the Sofia River drainage. The Amboaboa catfish (Arius festinus) is known only from the Amboaboa River near its confluence with the Mangarahara River, and possibly Lake Tseny. The Bemarivo River is a major tributary of the Sofia River. The pinstripe cichlid (Paretroplus menarambo) was originally found in the floodplain lakes of the Bemarivo River, where it was at one time an important food fish. It was last reported from Lake Sarodrano in the late 1990s, but not seen again in the wild for many years. A remnant population was later discovered in Lake Tseny. Captive breeding populations exist in Europe, North America, and Madagascar. The Ankofia River is a tributary of the Sofia River. The Ankofia cichlid (Ptychochromis inornatus) is confined to the Ankofia River drainage, including Lake Andrapongy. The Andrapongy catfish (Arius uncinatus) is confined to Lake Andrapongy within the Ankofia River drainage. Sparks’ killifish (Pachypanchax sparksorum) is known only from streams flowing into the Ankofia River and its principal tributary, the Anjingo River.

The Betsiboka River The Betsiboka River is located in west-central Madagascar (former Mahajanga and Antananarivo provinces). The Ikopa River is located in the former Antananarivo province. Bleeker’s cichlid (Paratilapia bleekeri) is known only from specimens collected during the nineteenth century from the Ikopa River. The Mangoro River The Mangoro River is located on the central-eastern coast of Madagascar (former Toamasina province). The ornate catfish (Gogo ornatus) is confined to the Mangoro and Nosivolo rivers, where it is threatened by loss of habitat and introduced fish species. The Nosivolo River is the primary tributary of the Mangoro River.

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The songatana cichlid (Oxylapia polli) is known only from the Marolambo Rapids of the Nosivolo River, where it is threatened by habitat destruction and sedimentation caused by deforestation. The Katria cichlid (Katria katria) is confined to the Nosivolo River. The Mandolotra rainbowfish (Rheocles lateralis) is confined to the headwaters of the Mandolotra River, a tributary stream of the Nosivolo River.

The Onilahy River The Onilahy is located in south-western Madagascar (former Toliara province). The Onilahy cichlid (Ptychochromis onilahy) was first described in 2006 based on five preserved specimens in the Museum of Natural History, Paris that were originally collected from the Onilahy River in 1962. It is most likely extinct, although the possibility exists of small remnant populations in remote areas. The trondo cichlid (Ptychochromoides betsileanus) was originally described from specimens collected in the Betsileo region of east-central Madagascar. It was subsequently recorded from the Ilanana River (a tributary of the Onilahy River), the Manantanana River (a tributary of the Mangoky), and the upper reaches of the Matsiatra, especially its tributary the Mandranofotsy, as well as in some waterfalls in streams near Ampamaherana and in the Manantanana and Zomandao rivers of the Ambalavao region. However, no more extant populations have been reported since the discovery of the one at Ilanana in 1997 despite numerous field surveys. It is now thought to be extinct owing to a combination of overfishing, declining water quality and competition with introduced Tilapia. The Onilahy catfish (Ancharius griseus) is known only from the Ihazofotsy and Ilanana rivers within the Onilahy River drainage. Miscellaneous Lakes, Rivers, and Marshes Lake Kinkony (Lac Kinkony in French) is located in northwestern Madagascar (former Mahajanga province). It is the country’s second largest. The Kinkony cichlid (Paretroplus dambabe) was formerly very common in the Mahavavy du Sud drainage but now appears to be mostly (or entirely) confined to Lake Kinkony, where it has been reduced to low numbers. The Ankavanana River is located in north-eastern Madagascar (former Antsiranana province), on the Masoala Peninsula. The Ankavanana rainbowfish (Bedotia masoala) is known only from a short stretch of the Ankananana River. The Antainambalana River is located in north-eastern Madagascar (former Toamasina province). The Makira cichlid (Ptychochromis makira) is known only from two specimens collected from the Antainambalana River. However, the river has not yet been extensively surveyed for freshwater fish, and it is possible that the species is much more widespread.

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The Sambirano River is located in north-western Madagascar (former Antsiranana province). The Sambirano frillfin goby (Bathygobius samberanoensis) is a little-known species of questionable taxonomic validity, apparently confined to the Sambirano River. The Manambolo River is located in eastern Madagascar (former Toamasina province). Sikora’s rainbowfish (Rheocles sikorae) is known only from the Manambolo River, where it was long thought to be extinct until rediscovered in 1996. It is now known from a few forested streams near the town of Marolambo. The Ivoloina River is located in eastern Madagascar (former Toamasina province). The Ivoloina rainbowfish (Bedotia madagascariensis) is confined to the lower Ivoloina River drainage, where it is threatened by habitat destruction. The Mananjary River is located in south-eastern Madagascar (former Fianarantsoa province). The Mananjary catfish (Gogo brevibarbis) is known only from a single specimen collected from the Mananjary River. The Mananara du Sud River is located in south-eastern Madagascar (former Fianarantsoa province). The Vondrozo cichlid (Ptychochromoides vondrozo) is known only from the Mananara du Sud River and its tributaries. The Andriambondro River is located in south-eastern Madagascar (former Fianarantsoa province). The Sandrananta catfish (Gogo arcuatus) is known only from the Sandrananta River, within the Andriambondro River drainage.

Coasts and Satellite Islands Madagascar features still extensive areas of coastal mangroves, wetlands, and littoral forests, although in general they receive little legal protection and are everywhere under threat by development. The black-banded plover (Charadrius thoracicus) is found discontinuously in western and southern coastal Madagascar (former Mahajanga and Toliara provinces). Petit’s gecko (Paragehyra petiti) is confined to cliffs along the Onilahy River near Lavenombato, in south-western coastal Madagascar (former Toliara province). The small area where this species occurs is potentially at risk from a proposed limestone mining operation which, if it goes ahead, may result in the complete destruction of its habitat. The Fasika skink (Paracontias fasika) is known only from a single specimen collected from a small patch of coastal forest just 10 m above sea level in far northern Madagascar (former Antsiranana province). The minimal skink (P. minimus) is similarly known only from a small area of coastal shrub in far northern Madagascar (former Antsiranana province). Rothschild’s skink (P. rothschildi) is a rare sand-dweller from coastal dunes and degraded dry forests near Sakalava Bay in far northern Madagascar (former Antsiranana province). The Antsiranana skink (Madascincus arenicola) is a fossorial or semi-fossorial skink known only from a small area of far northern coastal Madagascar (former Antsiranana

The Comoros Islands

province), where it has been recorded from Baie des Sakalava and Baie des Dunes in the Forêt d’Orangea region, and at an unnamed research site in the Ampombofofo region. Grandidier’s blind snake (Xenotyphlops grandidieri) is known only from coastal forest and shrubby dunes of Sakalava Bay, in far northern Madagascar (former Antsiranana province). Mocquard’s blind snake (X. mocquardi) is known only from a single locality in far northern coastal Madagascar (former Antsiranana province). The Maromandia diamond frog (Rhombophryne tetradactyla) is known only from the islands of Nosy Mangabe and Nosy Boraha (Ile St. Marie) off the north-eastern coast of Madagascar. Megson’s stump-toed frog (Stumpffia megsoni) is known only from a few small caves in far northern coastal Madagascar (former Antsiranana province).

Nosy Hara Nosy Hara is located off the far northern coast of Madagascar (former Antsiranana province). The Nosy Hara leaf chameleon (Brookesia micra), the world’s smallest lizard, is confined to Nosy Hara. The Nosy Hara diamond frog (Rhombophryne hara) is confined to Nosy Hara.

Nosy Mangabe Nosy Mangabe is located in Antongil Bay off the north-east coast of Madagascar (former Toamasina province), near the town of Maroantsetra. Covered by dense rainforest, it has been designated a special reserve and there are no permanent settlements on the island. The Nosy Mangabe diamond frog (Rhombophryne mangabensis) is known only from 13 specimens collected in 1991 and 2005.

Nosy Boraha Nosy Boraha (Île Sainte-Marie in French) is located off the north-eastern coast of Madagascar (former Toamasina province). Delalande’s coua (Coua delalandei) was a type of cuckoo restricted to Nosy Boraha. There it was often trapped for food and feathers, and the introduction of rats may have also contributed to its demise. Ultimately, however, it was the complete destruction of the island’s forests that was the cause of the extinction. It was last collected in 1834 (a report of a sighting from around 1930 appears to be unfounded).

Nosy Be and Nosy Komba Nosy Be is a small volcanic island located off the north-eastern coast of Madagascar (former Antsiranana province). One of Madagascar’s largest and busiest tourist resorts, it has been heavily deforested. One of the only remaining natural areas for the island’s many endemic species is the small Lokobe Strict Nature Reserve. The smaller islet of Nosy Komba is located to the south-east.

The Nosy Be sportive lemur (Lepilemur tymerlachsoni) is confined to Nosy Be. The Mamiratra mouse lemur (Microcebus mamiratra) is confined to Nosy Be. The Nosy Be skink (Pseudoacontias unicolor) appears to be confined to the Lokobe Strict Nature Reserve on Nosy Be. The Nosy Be mabuya skink (Trachylepis lavarambo) is confined to Nosy Be. Reuter’s blind snake (Madatyphlops reuteri) and Boettger’s blind snake (M. madagascariensis) are both known only from Nosy Be. The mystery cophyline tree frog (Cophyla occultans) appears to be confined to Nosy Be, with populations in north-eastern Madagascar formerly included within its range now thought to represent an as-yet undescribed species. The Andoany diamond frog (Rhombophryne pygmaea) and Boettger’s diamond frog (R. testudo) are both known only from Nosy Be and Nosy Komba.

The Glorioso Islands The Glorioso Islands (Îles Glorieuses in French) are a group of two small, uninhabited islands and three rocky islets located about 160 km north-west of Madagascar. The climate is lush and tropical, and the terrain low and flat. In 2012, France founded the Parc Naturel Marin des Glorieuses, a marine protected area that covers all flora and fauna. The Glorioso snake-eyed skink (Cryptoblepharus gloriosus) is thought to be at risk, as its restricted range makes it vulnerable to stochastic events and sea level rise due to climate change.

The Comoros Islands The Comoros Islands (Les Comores in French) are an archipelago of four main volcanic islands situated at the north end of the Mozambique Channel, north-west of Madagascar. Several smaller islands and submerged reefs are also included in the chain. Livingstone’s flying fox (Pteropus livingstonii) is one of the world’s largest bats. Considered common at the time it was first discovered by the explorer David Livingstone in 1863, it is today restricted to the islands of Anjouan and Mohéli where the population stands at about 1200. In 1992 the Durrell Wildlife Conservation Trust sent an expedition to the Comoros in order to bring some of the bats back to Britain to establish a captive breeding population. This has proved successful and the species is now established in several other zoos. Its status in the wild, however, remains perilous due to habitat destruction, hunting, and the threat of stochastic events. The Comoros rousette fruit bat (Rousettus obliviosus) is confined to the islands of Grande Comore, Anjouan, and Mohéli, where it is dependent upon caves for roosting. The Comoros green pigeon (Treron griveaudi) is currently confined to Mohéli, although it is thought to have been historically present on Grand Comoro and Anjouan.

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The Comoros cuckoo-shrike (Ceblepyris cucullatus) is found on the islands of Grande Comore and Mohéli, where the total population is thought to be less than 2500. The Comoros ground gecko (Paroedura sanctijohannis) occurs on all four main islands, where with the exception of Mayotte it is confined to small, high-elevation areas. The Comoros blind snake (Madatyphlops comorensis) is a burrowing species known only from a few specimens collected on Grande Comore and, in one instance, Anjouan.

Grande Comore Grand Comore is the largest of the Comoro Islands and is comprised of two shield volcanoes. The Comoro blue vanga (Cyanolanius comorensis) is divided into two subspecies. The Grande Comore blue vanga (C. c. bensoni) is a shrike-like bird known only from sightings in 1974 and 1981, and is possibly extinct. The Grande Comore bulbul (Hypsipetes parvirostris) is confined to Grande Comore, where it remains reasonably common.

Mount Karthala Mount Karthala, a highly active volcano, is the highest point in the Comoros. Nominally protected within Mount Karthala National Park, it provides a last refuge for several species of bird but continues to be threatened by deforestation. The Grande Comore scops owl (Otus pauliani) is confined to Mount Karthala. The Grande Comore drongo (Dicrurus fuscipennis) is confined to Mount Karthala, where in 1985 the total population was estimated at around 100. Humblot’s flycatcher (Humblotia flavirostris) is confined to Mount Karthala. The Karthala white-eye (Zosterops mouroniensis) is confined to Mount Karthala.

Anjouan Anjouan is the easternmost of the Comoros Islands. The Anjouan mouse-eared bat (Myotis anjouanensis) is a little-known species confined to Anjouan. The Anjouan scops owl (Otus capnodes) was long feared to be extinct. Between 1884 and 1897 it was collected in large numbers for museums, but not seen after that. In 1992, however, it was rediscovered on Anjouan. The species remains highly threatened, with an estimated population of 50–100 pairs. The Anjouan sparrowhawk (Accipiter francesiae pusillus) is confined to Anjouan, where it is seriously threatened. It was thought to be extinct until searches in the 1980s and in 2005 confirmed that it is still extant.

Mayotte Mayotte consists of a large main island (Grande-Terre), a smaller one (Petite-Terre), and several islets.

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The Mayotte drongo (Dicrurus waldenii) is a type of passerine bird confined to Mayotte, where it occurs widely but sparsely. The black-striped day gecko (Phelsuma nigristriata) and Robert Mertens’ day gecko (P. robertmertensi) are both confined to the island. Peters’ bright snake (Liophidium mayottensis) is found in very low numbers throughout the island.

Mohéli Mohéli is the smallest of the four major Comoros Islands. The Mohéli scops owl (Otus moheliensis) is confined to Mohéli. The Mohéli bulbul (Hypsipetes moheliensis) is a type of passerine bird confined to Mohéli. The Mohéli blue vanga (Cyanolanius comorensis comorensis) is confined to Mohéli.

The Seychelles The Seychelles comprise four main islands and hundreds of smaller islands and islets about located 950 km north of Madagascar. There are two main divisions: the larger granitic Seychelles and the smaller, outlying coralline Aldabras. The natural forest has been largely destroyed throughout, resulting in a steppe-like vegetation, serious erosion, and a sinking water table. Habitats are deteriorating as well due to invasion by introduced plants, principally cinnamon (Cinnamomum verum), as well as the drying effects of climate change. This, together with introduced cats and dogs and uninvited rats, has become a great threat to many endemic species. The Seychelles sheath-tailed bat (Coleura seychellensis) was historically found throughout the Seychelles, but declined rapidly with the destruction of its lowland forests in the late nineteenth and early twentieth centuries. Introduced predators and disturbance of its cave roosting sites have since made it one of the world’s rarest species, with the total population in 2016 estimated at less than 100. Two subspecies are recognized. The nominate form (C. s. seychellensis) is now confined to Mahé, having been extirpated from Praslin in the 1980s. The subspecies C. s. silhouettae is now confined to Silhouette, having been extirpated from La Digue in the 1980s. The Seychelles parakeet (Psittacula wardi) was endemic to Mahé and Silhouette islands, and possibly Praslin as well. It became extinct as a result of hunting pressure. The last known individuals were shot in 1893, and none were found during a survey in 1906. The Seychelles kestrel (Falco araeus) once occurred on almost all of the granitic islands but began to decline sharply after 1940, likely due to pesticide use and loss of habitat due to cinnamon cultivation and logging. By 1959 it was found only on Mahé, where there were less than 30 birds in 1964–65. Since then, it has made a remarkable recovery and now numbers around 800. The majority still live on Mahé (along with a few of its satellite islands Sainte Anne, Cerf, Conception, and

The Seychelles

Thérèse), but the species is once again to be found as well on Silhouette, Ile du Nord, Praslin, and occasionally La Digue and Félicité. The Seychelles turtledove (Nesoenas picturatus rostrata) was formerly distributed over all of the main islands but was largely absorbed by the introduced Madagascar turtledove (N. p. picturatus). Pure populations of the Seychelles turtledove now survive only on Cousin and Cousine Islands. The Seychelles paradise flycatcher (Terpsiphone corvina), which formerly occurred on several islands, has long been largely confined to the Veuve Nature Reserve on the western side of La Digue. A few birds have been found on neighbouring Marianne and Praslin but are not established there. Occasional sightings on Félicité have also been reported. On La Digue the population has been increasing steadily from an estimated 28 birds in 1965 to a current, stabilized 150–200. A translocation of 23 birds to Denis Island was conducted in 2008, with the aim of establishing a permanent population. The Seychelles magpie-robin (Copsychus sechellarum) once ranged throughout the islands but had been reduced to just 12–15 individuals on Frégate by the mid 1960s. A small, introduced population on Alphonse survived until about 1960, when it was wiped out by feral cats. By the late 1970s the Frégate population numbered around 40 birds but had been reduced to half that by 1981. Numbers remained around this level until the 1990s. In 1994, with the advent of a recovery programme, the population had increased to 48, with two birds translocated to Aride. In 2000, following further translocations, the population reached 86 (comprising 46 on Frégate, 23 on Cousin, 15 on Cousine, and 2 on Aride). Since then the population has continued to climb, reaching 283 by 2015, with the species now established on Denis Island as well. The Seychelles warbler (Acrocephalus sechellensis) was historically present on several of the inner granitic islands, until human disturbance in the twentieth century reduced the species to a single population of a hundred birds or less on Cousin Island by the 1960s. Conservation efforts have since helped it to recover on Cousin Island, and the species has been translocated to the islands of Aride, Cousine, Denis, and Frégate. The Seychelles swiftlet (Aerodramus elaphrus) occurs on the islands of Mahé, Praslin and La Digue, with one known breeding site on each island. A colony on Félicité has disappeared, as has a second one on Mahé. The Seychelles fody (Foudia sechellarum) is a small songbird that was reduced, by the early 1970s, to between 400 and 500 birds living on the islands of Frégate, Cousin, and Cousine, due mainly to a combination of human persecution (it was erroneously thought to consume crops) and habitat destruction. It has since recovered there and has been introduced to the predator-free islands of Aride, D’Arros, and Denis. The total population now stands at around 3500. The Seychelles white-eye (Zosterops modestus) was long thought to survive only in three tiny areas on Mahé, with only 25–35 individuals known in 1996. That year, however, a

previously unknown population of at least 250 was discovered on Conception. Since that time the species has been translocated to Frégate Island, North Island and Cousine. The Seychelles giant tortoise (Aldabrachelys gigantea), one of the world’s largest, was formerly common throughout the Seychelles and Aldabra islands. Four subspecies have been identified, among them the Aldabra giant tortoise (A. g. gigantea), which is discussed separately in this volume. Günther’s giant tortoise (A. g. hololissa) historically lived on Cerf, Cousine, Frégate, Mahe, Praslin, Round, and Silhouette. Thought to have been exterminated by 1840, a few captive and semi-wild specimens of unknown provenance still survive. The Seychelles black mud turtle (Pelusios subniger parietalis) occurs in the lowland marshes of Mahé, Cerf, Praslin, La Digue, Fregate, and Silhouette islands, where it has already lost most of its available habitat due to drainage. The total population is thought to be around 250. The Seychelles tiger chameleon (Archaius tigris) is an arboreal species from the forests of Mahé, Silhouette, and Praslin. It is threatened by habitat degradation and overcollection for the international pet trade. The giant bronze gecko (Ailuronyx trachygaster) is a canopy-dwelling species endemic to Silhouette and Praslin. Brauer’s burrowing skink (Janetaescinus braueri) is confined to Mahé and Silhouette. Vesey-Fitzgerald’s burrowing skink (J. veseyfitzgeraldi) is known from Mahé, Silhouette, Curieuse, Félicité, La Digue and Frégate. Both are considered vulnerable to habitat destruction and degradation, as well as to predation by introduced tailless tenrecs (Tenrec ecaudatus). Wright’s mabuya skink (Trachylepis wrightii) is confined to islands with seabird colonies (i.e. Aride, Cousin, Cousine, St. Pierre, Mammelles, Recifs, and Frégate). The Seychelles wolf snake (Lycognathophis seychellensis) is confined to Mahé, Silhouette, Praslin, Aride, La Digue, and Frégate. The Seychelles house snake (Lamprophis geometricus) is confined to Mahé, Silhouette, Praslin, and Frégate. The Seychelles pygmy frog (Sooglossus sechellensis) is found on the islands of Mahé, Silhouette, and Praslin, where it is still relatively common. Thomasset’s frog (S. thomasseti) is confined to Mahé and Silhouette, where it lives in wet rocky areas. Gardiner’s Seychelles frog (Sechellophryne gardineri), one of the world’s smallest frogs, is confined to high- and midaltitude rainforests on Mahé and Silhouette. Cooper’s black caecilian (Praslinia cooperi) is a burrowing amphibian known from a handful of higher-elevation localities on Mahé and Silhouette, with a possible old record from Praslin as well. It has not been collected since the early 1990s. The golden killifish (Pachypanchax playfairii) is a freshand brackish-water species endemic to streams and ponds in the granitic Seychelles. Little is known of its current status, although an introduced population on the island of Zanzibar is now believed to have been extirpated. Popularly kept as an aquarium fish, little is known of its status in the wild.

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Mahé Mahé is the largest island in the Seychelles. Considerable development and habitat alteration have taken place during the twenty-first century. Fortunately, several protected areas have been set aside, the largest of which is Morne Seychellois National Park, which covers almost a quarter of the island. Arnold’s giant tortoise (Aldabrachelys gigantea arnoldi) and Daudin’s giant tortoise (A. g. daudinii) were both historically confined to Mahé, where they were thought to have been exterminated by 1840. Captive and semi-wild specimens of the former subspecies still survive; however, of unknown provenance. The Seychelles scops owl (Otus insularis) is confined to Mahé, where the population appears to be stable. The Mahé caecilian (Grandisonia brevis) is found in only a few scattered localities on Mahé and perhaps Silhouette. Rare and hard to detect, it has not been observed in many years.

Praslin Lying 44 km north-east of Mahé, Praslin still has substantial tracks of tropical forest, with a large area in the south of the island having been designated as a national park. The Seychelles black parrot (Coracopsis barklyi) is largely confined to Praslin Island, with occasional reports from Curieuse 1 km to the north. Less than 50 were estimated there in 1965, although today that number is nearer to 100000.

Silhouette Silhouette Island is located 20 km north-west of Mahé and is the third largest in the Seychelles. Lush and mountainous, almost 95 per cent of it is protected within Silhouette National Park. The Seychelles palm frog (Sechellophryne pipilodryas) is confined to the forests of Silhouette Island, where it is closely associated with the thief palm (Phoenicophorium borsigianum).

Marianne Marianne is a small, nowadays uninhabited granitic islet that formerly served as a coconut plantation. The Marianne white-eye (Zosterops semiflavus) was a type of passerine bird only known with certainty from this island, although it may have occurred as well on Praslin, La Digue, Silhouette, and Mahé as well. It became extinct around 1888.

The Aldabra Islands The remote Aldabra Islands are a group of four raised coral atolls along with a number of islets located south of the Seychelles and north-west of Madagascar.

Aldabra Atoll Aldabra is the world’s second largest coral atoll, and consists of four main islands along with some 40 small islets. During the 1960s there were plans to transform it into a military airbase, which would probably have doomed most of the native species.

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Fortunately, the project was eventually cancelled after strong opposition from conservationists. The Aldabra flying fox (Pteropus aldabrensis) is confined to Aldabra Atoll, where the total population is reported to consist of only a few hundred animals. The Aldabra trident bat (Paratriaenops pauliani) is littleknown and apparently confined to Picard Island. The Aldabra brush warbler (Nesillas aldabrana) was discovered only in 1967. Confined to Ile Malabar, its population was already extremely small and vulnerable to predation and habitat alteration by invasive species. It was last recorded in 1983, and searches in 1986 confirmed its extinction. The Aldabra white-throated rail (Dryolimnas cuvieri aldabranus) is the last surviving flightless rail of the Indian Ocean region. While limited in its range and vulnerable to feral cats, it does not appear to be threatened at present. The Aldabra giant tortoise (Aldabrachelys gigantea gigantea) is endemic to the islands of the Aldabra Atoll, where for many centuries it was heavily exploited by European sailors but ultimately saved by some early established protected areas. Today over 100,000 there, with additional populations in the Sainte Anne Marine National Park on Moyenne Island in the Seychelles and on the island of Changuu, near Zanzibar. Populations have also been introduced to protected areas on Mauritius and Réunion.

Assumption Island Located about 27 km south of Aldabra Atoll, Assumption is a coral island with extensive sandy dunes. Due to the disruptive effect of guano mining, which lasted until 1983, it is dominated by expanses of bare rock and caves and sparsely covered with low-growing vegetation. The Assumption white-throated rail (Dryolimnas cuvieri abbotti) became extinct sometime during the early twentieth century. The Assumption day gecko (Phelsuma abbotti sumptio) is confined to the island, where it appears to be fairly common.

The Amirante Islands The Amirante Islands (Les Amirantes in French) are a group of 29 coral islands and atolls that belong to the outer islands of the Seychelles. The endemic Amirante turtledove (Nesoenas picturatus aldabrana) is now extinct due to hybridization with the introduced Madagascar turtledove (N. p. picturatus).

The Mascarene Islands The Mascarene Islands (Les Mascareignes in French) are located east of Madagascar, and consist of three large volcanic islands (Mauritius, Réunion, and Rodrigues) along with a number of remnants. The islands share a common geological origin and are notable for a unique, if sadly devastated, flora and fauna. Indeed, since the settlement of these islands by Europeans in the seventeenth and eighteenth centuries, no less than 45 endemic species and subspecies have become extinct. Many of these were birds that,

The Mascarene Islands

living unmolested, evolved to flightlessness and were doomed when man and his introduced animals ravaged these islands. The explosion in human population, partly explained by labour imported from India after 1848, has also been fatal to wildlife. Cultivation spread up the volcanic slopes, destroying vegetation and leading to the most serious erosion. What little wilderness remains, however, has been well protected in recent decades. Three species of flying fox (Pteropus) were historically found throughout the Mascarenes. The lesser Mascarene flying fox (P. subniger) is believed to have died out on Réunion in the 1860s and on Mauritius before 1873. The greater Mascarene flying fox (P. niger) was extirpated from Réunion in the early eighteenth century but has survived on Mauritius, where it is fairly common. The Rodrigues flying fox (P. rodricensis) was formerly found on Mauritius and Round Island but is today confined to Rodrigues. The small population is vulnerable to cyclones. The Mascarene petrel (Pseudobulweria aterrima) is known to breed only on Réunion, where it is very rare, although subfossil evidence has been discovered on Rodrigues and a dead bird found on Mauritius. The Mascarene grey parrot (Psittacula bensoni) was found on both Mauritius and Réunion. Easily hunted, the capture of one specimen would result in its calling out to summon the entire flock. It was exterminated on Réunion by the 1730s and by the 1760s on Mauritius. The Mascarene coot (Fulica newtonii) is known only from early traveller’s reports and bones collected on Mauritius and Réunion. It was extinct before 1700.

Mauritius Mauritius (Île Maurice in French) is a relatively large island surrounded by the world’s third largest coral reef and a ring of about 50 satellite islets, many of which have been set aside as protected areas. When early navigators first visited in the sixteenth century it was still uninhabited. Dense evergreen forests descended from the mountain peaks to the sea, and in drier areas there were palm savannas. Its position today is best described by mentioning that the population density is one of the highest on Earth. As a result, only about 17 per cent of the island remains forested, although of this only about 2 per cent can accurately be described as ‘natural forest’. Indeed, the remaining forests are nowhere entirely natural, because exotic, introduced plants have invaded the native vegetation. Severe loss and degradation of habitat has been compounded by introduced long-tailed macaques (Macaca fascicularis), sambar deer (Rusa unicolor), small Indian mongoose (Herpestes auropunctatus), rats, and feral cats. Frequent cyclones are another problem. All of these factors have served to threaten not only the indigenous flora but also the animals, of which Mauritius has already lost 11 species of endemic birds, and 3 reptiles. The Mauritius free-tailed bat (Mormopterus acetabulosus) is confined to a few lava tube caves where it is vulnerable to disturbance.

The Mauritius owl (Mascarenotus sauzieri) was last recorded in 1837, and was certainly extinct by 1859. The Mauritius kestrel (Falco punctatus) was historically widespread but had been reduced, by 1974, to just four wild birds (including a single breeding pair) due to loss of habitat, pesticide use, and introduced predators. It has since made a spectacular comeback with the help of a recovery programme, and by the early twenty-first century numbered between 600 and 800. The Mauritius night-heron (Nycticorax mauritianus) is known only from subfossil bones. Last recorded in 1693, it was probably exterminated by hunters before 1700. The Mauritius sheldgoose (Alopochen mauritiana) was last recorded in 1693, when it was said to be rare, and could not be found in 1698. The cause of its extinction is thought to have been hunting. Theodor’s duck (Anas theodori) is known only from explorer’s accounts and bones, and was last recorded in 1696. It or a similar species may have also lived on Réunion. The Mauritius red rail (Aphanapteryx bonasia) is known from traveller’s accounts, illustrations, and bones. It went extinct around 1693 due to hunting and perhaps predation by feral cats. The broad-billed parrot (Lophopsittacus mauritianus) was probably also flightless to judge from traveller’s accounts as well as from a 1638 sketch and some subfossil bones. Heavily hunted, the last records date from 1673 to 1675. It was absent in 1693. The Mauritius echo parakeet (Psittacula eques echo) is the only extant parrot of the Mascarene Islands, all others having become extinct due to human activity. It is also one of the most remarkable success stories in the history of conservation. By the early 1980s it appeared to be doomed. The roughly 10 individuals still surviving had hardly ever bred successfully due to a lack of suitable trees for nesting, nest predation, and competition from the introduced ring-necked parakeet (P. krameri). However, a dedicated research and captive-breeding programme was launched by the Durrell Wildlife Conservation Trust and Mauritian Wildlife Foundation to save the species. By the mid-1990s the numbers had risen to around 50–60. Since then the recovery has continued, with the total wild population sitting at 280–300. Perhaps the most famous extinct species of them all is the dodo (Raphus cucullatus), which has become a symbol for the meaningless and short-sighted extermination of animals by man. A flightless, heavy pigeon weighing somewhere between 13 and 23 kg, it was driven to extinction as a result of hunting by settlers and nest predation by introduced pigs. The last individuals are thought to have been killed on the offshore islet of Ile d’Ambre in 1662. All that remains today are numerous bones and specimen fragments, along with contemporary reports and paintings. The Mauritius wood pigeon (Columba thiriouxi) is known only from subfossil remains, and was extinct around 1730. The Mauritius blue pigeon (Alectroenas nitidissimus) was last reported in 1832, and is thought to have become extinct a few years later.

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The Madagascan Realm Figure 4.3 Sketch of two broad-billed parrots from a ship’s journal, 1601. (Credit: Joris Joostensz Laerle.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

Figure 4.4 Illustration of a dodo by Frederick William Frohawk, 1907. (Credit: Frederick William Frohawk.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The Mauritius pink pigeon (Nesoenas mayeri) was at one time one of the world’s rarest birds, having been reduced to just 12 wild individuals in 1986. A successful captive breeding and reintroduction programme has since raised the population to around 400 but the total range remains small, consisting as it does of the Black River Gorges of south-west Mauritius and the Ile aux Aigrettes just off the eastern coast. The Mauritius turtledove (N. cicur) is known only from subfossil remains. It was extinct around 1730.

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The Mauritius cuckoo-shrike (Lalage typica) is a rare endemic species with a total population of around 1000. The Mauritius black bulbul (Hypsipetes olivaceus) had been reduced to around 200 pairs in the mid1970s. Today the population remains dangerously small but stable. The Mauritius paradise flycatcher (Terpsiphone bourbonnensis desolata) is endemic to the island, where the population is estimated at around 250. The Mauritius olive white-eye (Zosterops chloronothos) had been reduced to around 200 pairs in the early 1990s. It has since recovered somewhat, but appears to be confined to the wettest native upland forests. The Mauritius fody (Foudia rubra) is restricted to southwestern Mauritius, where it declined from around 250 pairs in the mid-1970s to 108–122 pairs by 2001. In 2005, 45 handreared chicks were released onto Ile aux Aigrettes (on which the species apparently occurred historically), where by 2014 the population rose to between 180 and 200. Mauritius had at least two endemic species of giant land tortoise (Cylindraspis). Both were historically common, but by the beginning of the eighteenth century ships on their way to and from India were slaughtering or capturing thousands of them for food and export, particularly to hospitals. In addition, introduced species such as rats, cats, and pigs ate the eggs and hatchlings. As a result, both the domed Mauritius giant tortoise (C. triserrata) and the saddle-backed Mauritius giant tortoise (C. inepta) were likely extinct on the main island by about 1700, and on most of the surrounding islets by 1735. At least one of the species may have survived on Round Island until much later. An expedition there in 1844 found several very large specimens, although by that point the island was

The Mascarene Islands

already overrun with introduced rabbits and all the tortoises appear to have been gone by 1893. The Mauritius giant skink (Leiolopisma mauritiana) is known only from subfossil bones. It is thought to have gone extinct around 1600. Bojer’s skink (Gongylomorphus bojerii) was historically widespread in the Mascarenes, where it was divided into two subspecies. The Mauritius Bojer’s skink (G. b. bojerii) was found on Mauritius and surrounding islands, but was extirpated from the former due to the introduction of predatory snakes and shrews and now survives only a handful of predator-free islands. Hoffstetter’s blind snake (Madatyphlops cariei) was last recorded during the seventeenth century, and is thought to have been exterminated by introduced species.

Round Island Round Island (Île Ronde in French) is located about 22 km off the north-eastern coast of Mauritius. The largest of a group of volcanic islets that serve as the last refuge of a remarkable reptilian fauna, it has never been inhabited and is almost inaccessible, rising steeply out of the sea. Only about 150 years ago it was covered by forest, but owing to monumental stupidity goats and rabbits were introduced there about 1850. The result was that, by the 1970s, there was no forest left and indeed very little vegetation at all, with only some small groups of endemic bottle palms and most of the soil eroded away. Insect life had all but disappeared and the endemic lizards that feed on them vanished too, along with the snakes that preyed upon the lizards. Fortunately, since the mid-1970s the island has been the subject of a concerted conservation programme. Goats and rabbits have been eliminated, the forest has begun to regenerate and the surviving reptiles, all highly threatened, have been safeguarded by a successful captive-breeding programme. More recently, Aldabra giant tortoises (Aldabrachelys gigantea gigantea) have been introduced. The Round Island day gecko (Phelsuma guentheri), which historically occurred on Mauritius itself, was reduced by the 1970s to a few hundred individuals on Round Island. The population is now fairly stable. Telfair’s skink (Leiolopisma telfairii) was near extinction by the late 1960s, but has made a remarkable recovery in recent years thanks to conservation efforts. The Round Island burrowing boa (Bolyeria multocarinata) was last reported in 1975, and is now certainly extinct. The Round Island keel-scaled boa (Casarea dussumieri) was thought to be extinct in the 1970s when it was unexpectedly rediscovered. Today the species is established on Round Island as well as on Gunner’s Quoin, Flat Island, and Ile de la Pas. Subfossil remains have shown that it was historically present on Mauritius itself.

Serpent Island Serpent Island (Île Serpent in French) is located close to Round Island.

The Serpent Island night gecko (Nactus serpensinsula) and Durrells’ night gecko (N. durrellorum) are both confined to Serpent Island.

Gunner’s Quoin, Pigeon Rock, Ile aux Vacoas, and Flat Island Gunner’s Quoin, Pigeon Rock, Ile aux Vacoas, and Flat Island are small islets located off the northern coast of Mauritius. The lesser night gecko (Nactus coindemirensis) is confined to Gunner’s Quoin, Pigeon Rock, Ile aux Vacoas, and Flat Island.

Réunion Réunion (La Réunion in French, and formerly Île Bourbon), located about 175 km south-west of Mauritius, is the westernmost and largest of the Mascarene Islands. Although the evergreen forests that originally covered the island have been destroyed and altered by humans, it has more undisturbed vegetation than the other Mascarene Islands. Nevertheless, at least 1 mammal, 11 birds, and 2 reptiles have become extinct since the arrival of Europeans, mainly due to hunting pressure and/or the predations of introduced rats and cats. Barau’s petrel (Pterodroma baraui) is effectively endemic to Réunion for breeding purposes, but disperses widely throughout the tropical Indian Ocean at other times. The Réunion owl (Mascarenotus grucheti) is known only from subfossil bones. It was exterminated in the early seventeenth century. The Réunion kestrel (Falco duboisi) went extinct after 1672. The Réunion harrier (Circus maillardi) is currently estimated to number between 200 and 560 mature individuals. The Réunion sacred ibis (Threskiornis solitarius) is known from bones collected on Réunion, and is likely synonymous with the ‘solitaire’ known from numerous early accounts. If so, the last report dates from 1763. The Réunion night heron (Nycticorax duboisi) was last recorded in 1674. It was probably driven to extinction by hunters before 1700. The Réunion purple gallinule (Porphyrio caerulescens) was an inhabitant of montane forest, where it was described by early settlers. It was likely hunted to extinction around 1730. The Réunion sheldgoose (Alopochen kervazoi) is known only from subfossil bones collected in 1974. It was exterminated after 1672. The Réunion rail (Dryolimnas augusti) was described from fossils and thought to correspond to a rail species mentioned by Dubois in 1674. Likely flightless, it was exterminated in the late seventeenth century. The Réunion parrot (Mascarinus mascarin) was a unique species with no near relatives. First mentioned in 1674, several live specimens were brought back to France in the late eighteenth century. The last reports of wild birds date from the 1770s, and the species may have been extinct by 1804 (although tradition has it that a single individual survived in

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The Madagascan Realm

the King of Bavaria’s menagerie until 1834). Two museum specimens are all that remain today. The Réunion echo parakeet (Psittacula eques eques) has been extinct since about 1770. The Réunion pink pigeon (Nesoenas duboisi) was last recorded in 1674. It is thought to have been extinct by the early eighteenth century. The Réunion crested starling (Fregilupus varius) was apparently common as late as the 1830s, but was thereafter greatly affected by introduced diseases and hunting. The last known specimen was shot in 1837, and the species became extinct some time during the 1850s. The Réunion cuckoo-shrike (Lalage newtoni) is restricted to two very small areas in the north-west of the island, where the total population is estimated at fewer than 30 pairs. The Réunion fody (Foudia delloni) was a type of passerine bird that was once so abundant it was considered a crop pest. Last seen shortly after 1672, it was likely driven extinct by introduced rats. The Réunion giant tortoise (Cylindraspis indica) was common up until the early eighteenth century, but was easy prey for European sailors and killed or collected in vast numbers. In addition, introduced pigs, cats, and rats destroyed the eggs and hatchlings. By the beginning of the nineteenth century only a few still survived in highland areas, where they were finally driven extinct during the 1840s.

The Réunion Bojer’s skink (Gongylomorphus bojerii borbonica) was observed and collected many times during the nineteenth century, but has not been recorded since.

Rodrigues Rodrigues (Île Rodrigues in French) is located about 560 km east of Mauritius. The smallest of the Mascarene Islands, it is notably hilly with a high central ridge cut with deep valleys. A large fringing reef surrounds it, forming a lagoon within which lie 18 small islets. In 1691, when Europeans first settled here, it had been covered with forests, of which only patches remain on the reef. There is hardly any native vegetation left at all on Rodrigues and a large percentage of the island’s vertebrates have vanished. The Rodrigues owl (Mascarenotus murivorus) is known only from bones and early reports. It was last recorded in 1726. The Rodrigues night-heron (Nycticorax megacephalus) is known only from bones and early reports. It was last recorded in 1726, and mentioned as absent in 1761. The Rodrigues flightless rail (Erythromachus leguati) was driven to extinction by heavy hunting. It was last recorded in 1726, with its absence being noted by Alexandre Guy Pingré in 1761. The Rodrigues parrot (Necropsittacus rodricanus) was last seen about 1761, and presumably became extinct soon after.

Figure 4.5 The only known life drawing of a Réunion crested starling, by Paul Philippe Sauguin de Jossigny, early 1770s. (Credit: Paul Philippe Sauguin de Jossigny.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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The Rodrigues parakeet (Psittacula exsul), long reduced by habitat destruction and hunting, is thought to have ultimately gone extinct as a result of vicious cyclones in 1875. The Rodrigues solitaire (Pezophaps solitaria), a flightless relative of the dodo, is known only from bones and the accounts of travellers. Reported in 1761, it had become extinct by 1778. The Rodrigues blue pigeon (Alectroenas payandeei) is known only from a single subfossil bone. It was not reported by early travellers to the island and likely disappeared by the 1690s. The Rodrigues turtledove (Nesoenas rodericanus) is known from subfossil bones. It went extinct between 1726 and 1761 as a result of predation by rats. The Rodrigues starling (Necropsar rodericanus) was last recorded in 1726, and not found during a visit to the island in 1761. The Rodrigues warbler (Acrocephalus rodericanus) was historically quite common but dwindled to very low numbers by the 1970s. Fortunately, it proved highly adaptable to introduced vegetation and has made a considerable comeback in recent years. The Rodrigues fody (Foudia flavicans) was reduced to just five or six pairs in 1968. It has made a remarkable recovery in the decades since owing to the expansion of native and exotic woodland, and is no longer considered to be immediately threatened. Two species of giant land tortoise (Cylindraspis) once roamed this island in huge numbers. When the French traveller François Leguat visited there in 1691, he reported ‘such a plenty of land turtles in this isle that sometimes you see two or three hundred of them in a flock, so that you may go above a hundred paces on their backs’. In 1759 one ship alone slaughtered 6000 tortoises and within 18 months 30,000 tortoises were captured. By the end of the century both the domed Rodrigues giant tortoise (C. peltastes) and the saddle-backed Rodrigues giant tortoise (C. vosmaeri) were on the point of extinction. One or the other species appears to have survived at least until 1802, when a few were reported killed in large fires used to clear the island’s vegetation for agriculture. Two species of day gecko (Phelsuma) historically endemic to Rodrigues are now thought to be extinct. The Rodrigues giant day gecko (P. gigas) was common on Rodrigues in 1691–93, but seems then to have disappeared rapidly. In any case it was gone before 1874. The Rodrigues blue-dotted day gecko (P. edwardnewtoni) was probably extinct on Rodrigues before 1874, but subsisted on Ile aux Frégates at least until 1917. In 1963 no individuals were found there in spite of an intensive search.

Balance for the Madagascan Realm While the precise timeline is still debated Madagascar was certainly one of the last major landmasses on Earth to be settled by humans, predating only that of Iceland and New Zealand. The earliest archaeological evidence for human presence there may be

as much as 10,000 years old, although actual settlement by Austronesian peoples first began between 550 and 350 B . C . The latter are believed to have arrived in successive waves by outrigger canoe from Borneo. These early settlers practiced slash-and-burn agriculture to clear the coastal rainforests for cultivation and hunted the island’s megafauna, wiping out most of them in the process. By A . D . 600 groups had begun to clear the forests of the central highlands as well. Arab traders first reached the island between the seventh and ninth centuries, and a wave of Bantu-speaking migrants crossed the Mozambique Channel from south-eastern Africa around A . D . 1000. Others continued to reach on the island over time, including South Indian Tamil merchants who introduced the zebu (Bos primigenius indicus), a type of long-horned humped cattle, which they kept in large herds. Irrigated paddy fields were developed in the highlands and were later extended with terraced paddies. By the seventeenth century the rising intensity of land cultivation and the ever-increasing demand for zebu pasturage had largely transformed the central highlands from a forest ecosystem to grassland. European contact with Madagascar began in 1500–01 with the Portuguese sea captain Diogo Dias first sighting the island. In 1528 Diogo Rodrigues explored what are now the Mascarene Islands, naming what are now Réunion, Mauritius, and Rodrigues. The first small Portuguese settlements began to appear along the coasts of Madagascar a few years later for the purposes of trade and missionary work. By the late seventeenth century the French, too, were establishing trading posts along the eastern coast, and the island began to gain prominence among European pirates and slave traders as well. Indeed, the small island of Nosy Boroha, off the northeastern coast, has been proposed by some historians as the site of the legendary pirate utopia of Libertalia. Many European sailors were shipwrecked along the coasts, among them the 17year-old Englishman Robert Drury, whose journal is one of the few written depictions of life in southern Madagascar during the early eighteenth century. The wealth generated by maritime trade spurred the rise of organized native kingdoms, some of which grew quite powerful. Unlike other African states, Madagascar had a relatively short period of colonization, lasting from the late nineteenth century until the early 1960s. The Mascarenes, by contrast, had already been settled by the Portuguese, Dutch, and French as early as the seventeenth century, in the process unleashing an unbelievably wanton destruction of animal life. In 1883 the French invaded Madagascar itself in what became known as the first Frano-Hova War. At its conclusion, the island ceded the northern port town of Diego Suarez (modern Antsiranana) to France. In 1890 the British accepted the full formal imposition of a French protectorate on the island, although this authority was not recognized by the native government. To force its capitulation, in 1894 the French bombarded and occupied the harbour of Tamatave (modern Toamasina) on the eastern coast, and the following year Majunga (modern Mahajanga) on the west coast. A French

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military expedition then marched to the capital Tana (modern Antananarivo), where the royal palace was bombed with heavy artillery leading Queen Ranavalona III to surrender and the royal family to go into exile. France formally annexed Madagascar in 1896 and declared the island a colony the following year. Slavery was abolished in 1896, and approximately 500,000 slaves were freed. Under colonial rule wide paved boulevards and buildings were constructed in Antananarivo, and the royal palace was turned into a museum. Schools were built throughout the island and a French-language education became mandatory. Taxation enabled the construction of a railway and roads linking key coastal cities to the capital. Plantations were established for the production of a variety of export crops, along with ambitious and often highly destructive development schemes. Even worse was the introduction of livestock and exotic species. Zebu cattle were brought in by the millions, causing enormous environmental damage. Many of the herds roamed in a semi-wild state, feeding, and reproducing freely but without being controlled by predators. Furthermore, between 1930 and 1940 red deer (Cervus elaphus) and fallow deer (Dama dama) were also introduced. They too were detrimental to the native vegetation, which had evolved without pressure from ungulate browsing. With independence and an ever-exploding human population, however, the destruction of the remaining forests accelerated, mainly due to slash-and-burn agriculture. It was only in relatively recent decades that an environmental awareness began to take hold

and a system of protected areas put in place, with the first national parks established by the late 1950s. By then it was almost too late. The occupation of Madagascar during World War II had tarnished the prestige of the colonial administration and galvanized a growing independence movement, and the Malagasy Republic was ultimately proclaimed in 1958 as an autonomous state within the French Community. Since then, Madagascar has transitioned through four republics with a rather neo-colonial arrangement, all noted for their corruption. Today, what little remains of the island’s ecosystems and unique wildlife continue to be under threat by the encroachment of humans. In recent historical time (i.e. since A . D . 1500), the Madagascan Realm has lost at least 49 species/3 subspecies of vertebrates. Among the extinct forms 2 species are mammals, 37 species/2 subspecies are birds, 7 species/2 subspecies are reptiles, and 3 species are freshwater fishes. One other species of mammal and 2 species of reptile are possibly extinct, and one species of fish is currently extinct in the wild. In addition, there are 636 species/20 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 143 species/5 subspecies are mammals, 65 species/6 subspecies are birds, 192 species/9 subspecies are reptiles, 163 species are amphibians, and 73 species are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

2 species

1 species

~ species

143 species

~ subspecies

~ subspecies

~ subspecies

5 subspecies

2 taxa

1 taxon

~ taxa

148 taxa

37 species

~ species

~ species

65 species

2 subspecies

~ subspecies

~ subspecies

6 subspecies

39 taxa

~ taxa

~ taxa

71 taxa

7 species

2 species

~ species

192 species

1 subspecies

~ subspecies

~ subspecies

9 subspecies

8 taxa

2 taxa

~ taxa

201 taxa

~ species

~ species

~ species

163 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

~ taxa

~ taxa

163 taxa

3 species

~ species

1 species

73 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

3 taxa

~ taxa

1 taxon

73 taxa

49 species

3 species

1 species

636 species

3 subspecies

~ subspecies

~ subspecies

20 subspecies

52 taxa

3 taxa

1 taxon

656 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

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Chapter

5

The Indo-Malaysian Realm

The Indo-Malaysian Zoogeographic Realm is comprised primarily of the tropical parts of South and South East Asia although extending some degrees north in places into the subtropical zone, particularly in northern India. It includes most of the Indian subcontinent south of the Himalayas together with most of Indochina, the Malay Peninsula, and the islands of the Sunda Archipelago. Biogeographically it is divided among the Oriental, Sundaic, and Wallacean zoogeographic regions. The majority of it was originally covered by both tropical moist and dry forests, with a few scattered areas of savanna and desert. Its evolution was greatly affected by iceage sea-level changes. At the height of the last glacial age, the oceans were hundreds of metres lower than at the present time. Lands like Indonesia and the Philippines, and perhaps New Guinea and Australia as well, were not as widely separated as today. At least Indonesia was certainly connected with the mainland and with the Philippines as late as about 18,000 years ago. These climatic oscillations, with long warm, interglacial stages between the cold periods, as well as the rise and fall of the oceans, had a tremendous impact on animal distribution patterns and migrations, and this is still going on. When sea levels dropped during the Pleistocene Bali was connected to both Java and Sumatra as well as to the Asian mainland, but the deep water of the Lombok Strait continued to keep the rest of the Sunda Islands isolated. The Lombok Strait lies to the immediate west of the island, marking the passage of the biogeographical division between the prolific fauna of the Indomalayan ecozone and the distinctly different, but similarly prolific, fauna of Australasia – this distinction is known as the ‘Wallace Line’ (or ‘Wallace’s Line’) and is named after Alfred Russel Wallace, the first to comment on the division between the two regions as well as on the abrupt boundary between them. Geologically speaking, the Indo-Malaysian Realm combines elements of the ancient supercontinents of Laurasia and Gondwana. Gondwanian elements were first introduced by what is now India, which detached from Gondwana approximately 90 million years ago, carrying its flora and fauna northward before colliding with Asia between 45 and 30 million years ago. Later, as what are now Australia and New Guinea drifted north, the collision of their plates pushed up the islands of the Wallacean Region, which were separated from one another by narrow straits, thereby allowing periodic

exchanges of plants and animals. All told, the IndoMalaysian Realm remains one of the richest areas for vertebrate diversity on Earth. The Asian or Asiatic elephant (Elephas maximus) is the world’s second largest land animal. At least three subspecies are found sporadically across South and South East Asia, which will be discussed in detail below. The Asiatic lesser one-horned rhinoceros (Rhinoceros sondaicus) is divided into three subspecies that historically occurred throughout South East Asia from eastern India and southern China to Indonesia. They will be discussed below. The Asiatic two-horned rhinoceros (Dicerorhinus sumatrensis) is similarly divided into three subspecies that historically occurred across South and South East Asia, including the islands of Sumatra, Borneo and Java. They too will be discussed below. The wild Asiatic buffalo or water buffalo (Bubalus arnee) was once widespread and common across southern Asia from Mesopotamia to Indochina and Indonesia. Habitat destruction, hunting, and diseases (particularly rinderpest) transmitted from domestic cattle have caused a considered decline, and the species is now found only in a few wet grassland, swamp and riverine areas of India, Nepal, Bhutan, Thailand, Cambodia, and possibly Myanmar. Unfortunately, many of these surviving populations are thought to have interbred with feral or domestic water buffalo (B. bubalus). The banteng (Bos javanicus) is a type of wild cattle that was historically widespread in the forests and grasslands of South East Asia, but which has declined in most parts of its range due to habitat destruction and heavy hunting pressure. Three subspecies will be discussed below. Carpenter’s white-handed gibbon (Hylobates lar carpenteri) is confined to northern and north-eastern Thailand, where it is threatened by hunting and habitat destruction. The macaques (Macaca) are a group of large, grounddwelling monkeys. Many are declining due to hunting and habitat destruction. The long-tailed macaque (M. fascicularis) is widespread across South East Asia where it is divided into at least 10 subspecies. The nominate form (M. f. fascicularis) occurs in Cambodia, Vietnam, the Malay Peninsula, the Philippines, Borneo, Sumatra, Java, Bali, Sumba, Timor and on a number of smaller islands. The northern pig-tailed macaque (M. leonina) ranges throughout much of mainland

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South East Asia, in a variety of habitats. The stump-tailed macaque (M. arctoides) is found in south-western China (Guangdong, Guangxi, Guizhou, and Yunnan), north-eastern India (Arunachal Pradesh, Assam, Manipur, Meghalaya, Mizoram, Nagaland, and Tripura), Laos, northern Myanmar, Thailand, Vietnam, and north-western peninsular Malaysia. It has been introduced to Hong Kong. The Bengal slow loris (Nycticebus bengalensis) is a small, nocturnal prosimian that remains widespread within the Indian subcontinent and Indochina, including the Malay Peninsula, but is everywhere threatened by habitat destruction, hunting for use in ‘traditional medicine’ and collection for the pet trade. The sun bear (Helarctos malayanus) is the world’s smallest bear species, and gets its name from the yellowish crescent patch on its chest. Two subspecies are found in the forests of South East Asia. The Malayan sun bear (H. m. malayanus) still ranges widely across Indo-China from north-eastern India to Sumatra, but has been extirpated from most of the areas it formerly inhabited. The main threats are loss of habitat and hunting for use in ‘traditional medicine’. The nominate form of the Asiatic black bear (Ursus thibetanus thibetanus) is still found widely from north-eastern India (Assam) and Nepal through Indochina to the Malay Peninsula, but is everywhere threatened by hunting for use in ‘traditional medicine’. The fishing cat (Prionailurus viverrinus) is a medium-sized wild cat found in widely scattered populations across South and South East Asia, where it favours wetlands. It has been extirpated over most of its historical range due to habitat destruction and human persecution. Pocock’s wild dog (Cuon alpinus fumosus) is found widely but patchily in Mongolia, southern China (Szechuan), Myanmar, Thailand, Laos, Cambodia, Vietnam, peninsular Malaysia and Java, but is everywhere threatened by loss of habitat, depletion of its prey base and human persecution. The greater hog badger (Arctonyx collaris) is found widely but patchily in South East Asia from southern China (Yunnan) to the Malay Peninsula, where it is threatened by hunting and trapping. The large-spotted civet (Viverra megaspila) is a rarely seen species found widely but patchily in southern China, Cambodia, Laos, Myanmar, Thailand, Vietnam, and peninsular Malaysia. The Sunda pangolin (Manis javanica) is an arboreal species still found throughout much of South East Asia from Myanmar and Vietnam to Java. It is everywhere seriously threatened, however, by overcollection for use as food and in Chinese ‘traditional medicine’. The Asian small-clawed otter (Aonyx cinereus) continues to have a very wide distribution extending from South Asia, through South East Asia to Sumatra, Borneo, and Palawan in the Philippines. The species has undergone considerable declines, however, due to hunting and loss of habitat, and has been extirpated from many areas. The escape of captive

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specimens in England in recent years has resulted in an introduced population there. The hairy-nosed otter (Lutra sumatrana) is the rarest and least-known of Asian otters. Once thought to be extinct due to overhunting, it has been rediscovered in a number of areas across South East Asia (Thailand, Vietnam, Cambodia, and Sumatra), with an additional record from northern Myanmar. There are also historical records from Borneo, peninsular Malaysia, Laos, and north-eastern India. The smooth-coated otter (Lutrogale perspicillata) is divided into three subspecies. The nominate form (L. p. perspicillata) remains widespread in South and South East Asia, but is everywhere threatened by loss of habitat, pollution, and hunting. Temminck’s flying squirrel (Petinomys setosus) is found patchily throughout much of South East Asia in northern Myanmar, Thailand, the Malay Peninsula, Sumatra, and northern Borneo. It is everywhere threatened by loss of habitat. The greater marmoset rat (Hapalomys longicaudatus) is known from a few scattered localities in southern Myanmar, western Thailand, and the Malay Peninsula. The lesser ranee mouse (Haeromys pusillus) is known only from a few specimens collected on Palawan and Calauit in the Philippines and from northern Borneo. The white-collared fruit bat (Megaerops wetmorei) is found widely but patchily in south-western peninsular Malaysia, central Sumatra, northern Borneo, and Mindanao in the Philippines. The tailless fruit bat (M. kusnotoi) is currently confined to two disjunct areas in eastern and western Java, although there is evidence to suggest that the species formerly occurred on Bali and Lombok as well. Both are threatened by loss of habitat. Andersen’s flying fox (Pteropus speciosus) is known for certain only from Mindanao and the Sulu Islands in the Philippines, as well as from the Talaud Islands and two small islands off the south-eastern coast of Borneo. The goldenmantled flying fox (P. pumilus) is widespread within the Philippines, occurring as well on Miangas (Palmas) Island off the north-eastern coast of Borneo. Both are threatened by loss of habitat, hunting, and human persecution. The Javan slit-faced bat (Nycteris javanica) is a forest and cave-dependent species known sporadically from Java, Nusa Penida, and the Kangean Islands of Indonesia. Thomas’ woolly bat (Kerivoula flora) has a highly disjunct range in north-eastern Borneo, Bali, and in the Lesser Sunda Islands (Sumbawa, Sumba, and Flores). It is threatened by loss of habitat. The slender-billed vulture (Gyps tenuirostris) and the white-rumped vulture (G. bengalensis) were both historically widespread and common in mainland South and South East Asia. Populations began to decline during the latter half of the nineteenth century owing to loss of habitat, but in recent years have all but collapsed due to poisoning and human persecution. Both are now largely confined to parts of India, Myanmar, and Cambodia.

The Indo-Malaysian Realm

The red-headed vulture (Sarcogyps calvus) was once similarly widespread and abundant in South and South East Asia from the Himalayas to southern peninsular Malaysia, but has undergone a massive and rapid decline mainly due to poisoning, habitat destruction and human persecution. The spot-billed pelican (Pelecanus philippensis) nested in huge numbers throughout South and South East Asia in the late nineteenth century. One colony alone on the Sittaung River, in what was then southern Burma, was described in 1877 as covering 300 km2 and containing millions. Immense numbers still bred in the area in 1910, but by 1939 all the birds had completely disappeared due to deforestation and human disturbance. Similar declines were noted elsewhere, but the species is now well-protected and still numbers between 13,000 and 18,000 overall. The milky stork (Mycteria cinerea) is found widely but sporadically in Cambodia, Peninsular Malaysia and on the islands of Sumatra, Sulawesi, Buton, Java, Bali, and Sumbawa in Indonesia, where it primarily inhabits mangroves, tidal mudfields, rice fields, and swamps. It is seriously threatened by intense hunting pressure at its nesting colonies as well as by human disturbance and loss of its coastal habitat. Two subspecies of white-necked stork (Ciconia episcopus) occur patchily throughout much of Asia where they are everywhere threatened by hunting and habitat destruction. The nominate form (C. e. episcopus) is found from India through Indochina to the Philippines, the Malay Peninsula and northern Sumatra. The Sundaic white-necked stork (C. e. neglecta) is found in southern Sumatra, Java, Sulawesi and the Lesser Sunda Islands. The lesser adjutant (Leptoptilos javanicus) is a type of stork found widely in wetland areas across South and South East Asia, but is everywhere threatened by hunting and habitat destruction. The white-shouldered ibis (Pseudibis davisoni) historically occurred widely, if patchily, from south-western China to Borneo, but is now restricted to northern and eastern Cambodia, extreme southern Laos, and a small area of Borneo (East Kalimantan, Indonesia). Based on its significant declines it has been described as the most seriously threatened large waterbird in South East Asia. The black-bellied tern (Sterna acuticauda) historically occurred throughout southern Asia from Pakistan to Vietnam but has been wholly or entirely extirpated in South East Asia due to loss of habitat and the overcollection of eggs. The surviving population, thought to be less than 10,000, is now confined to the vicinity of large rivers on the Indian subcontinent. The white-winged wood duck (Asarcornis scutulata) was historically widely distributed in the wetlands of South East Asia, but has undergone an astonishing decline over the last century. Today a total of perhaps 1000 birds survive in isolated pockets scattered from India to Sumatra. The masked finfoot (Heliopais personatus) is an aquatic, grebe-like bird found very patchily from north-eastern India

and Bangladesh through South East Asia to Sumatra and perhaps Java. In 2016 the total population was estimated at between 1000 and 2500. The green peafowl (Pavo muticus) was historically widespread and common throughout much of South East Asia but has been extirpated from most of its former range by loss of habitat and hunting for its meat and feathers. Populations are now confined to southern China (Yunnan), parts of Myanmar, Thailand, Laos, Cambodia and Vietnam, and the island of Java. The yellow-crested cockatoo (Cacatua sulphurea) historically ranged throughout a relatively wide area of Indonesia, with an additional, introduced population on Hong Kong. It is now seriously threatened by trapping for the international cagebird trade, with over 100,000 having been illegally exported from Indonesia between 1980 and 1992 alone. Several subspecies will be discussed in more detail below. The sunset lorikeet (Trichoglossus forsteni) was, as a species, historically found in the western Lesser Sunda Islands and smaller islands to the north, where it has suffered serious declines due to overcollection for the cage-bird trade. Mitchell’s sunset lorikeet (T. f. forsteni) is known from a few areas on Bali and Lombok. The grey imperial pigeon (Ducula pickeringii) is confined to a scattering of small islands in the southern Philippines and off the northern coasts of Borneo and Sulawesi, where it is threatened by habitat destruction and hunting. The straw-headed bulbul (Pycnonotus zeylanicus) was, up until the late twentieth century, common throughout the Malay Peninsula and Indonesia but has suffered massive declines primarily due to trapping for the cage-bird trade. It is now most likely extirpated from Thailand, Myanmar, Java and a few smaller islands, surviving only in peninsular Malaysia, Singapore, and parts of Borneo and Sumatra. The great slaty woodpecker (Mulleripicus pulverulentus) is the largest species of woodpecker that is certain to exist today. While still found across a large part of the Indian subcontinent and South East Asia it is everywhere threatened by the destruction of its preferred habitat (old-growth forests), and is hunted in some areas. The Siamese crocodile (Crocodylus siamensis) is among the most threatened of all crocodilians. Historically found over much of mainland South East Asia as well as in Indonesia, it was heavily hunted for its hide and by the 1990s virtually extinct in the wild. The situation has not changed much since, with only small wild populations scattered in remote areas of Vietnam, Cambodia, Laos, Thailand and Indonesia (Borneo and Java). The species is very common in captivity, however, particularly in crocodile farms. The Asian giant tortoise (Manouria emys) is divided into two subspecies found patchily in South East Asia. The southern Asian giant tortoise (M. e. emys) occurs in the Malay Peninsula and parts of Sumatra and Borneo, where it has suffered considerable declines due to loss of habitat and collection for use as food.

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The elongated tortoise (Indotestudo elongata) remains widespread in India (Tripura, Jalpaiguri, East Bengal, and Bihar), Nepal, Bangladesh, Myanmar, Laos, Thailand, Cambodia, Vietnam, peninsular Malaysia, and southern China, but is everywhere declining due to mass harvesting for food and overcollection by the international pet trade. The frog-faced softshell turtle (Pelochelys cantorii), one of the world’s largest extant freshwater turtles, historically occurred across much of southern Asia from India and Indochina to Sumatra, Borneo, Java and the Philippines. The species has disappeared from most of this range due to hunting and habitat destruction. The black-rayed softshell turtle (Amyda cartilaginea) remains fairly widespread from north-eastern India through South East Asia to Java and Borneo, but is everywhere threatened by overcollection for use as food. The Malayan flat-shelled turtle (Notochelys platynota) remains widespread in South East Asia but has suffered serious declines due to overharvesting for food. The South East Asian box turtle (Cuora amboinensis) is divided into at least four subspecies. The Burmese box turtle (C. a. lineata) is confined to parts of Myanmar, where it is seriously threatened by overcollection for use as food and in ‘traditional medicine’. The sunburst turtle (Heosemys spinosa) is found widely but patchily in Myanmar, the Malay Peninsula (including Singapore), Borneo and the Philippines, where it has been seriously reduced in number by overcollection for Asian food markets. The black marsh turtle (Siebenrockiella crassicollis) is found patchily in South East Asia, including the islands of Sumatra, Java and Borneo, where it is threatened by overharvesting. Two species of roofed turtle (Batagur) are among the most seriously endangered chelonians in the world. The northern roofed turtle (B. baska) continues to survive in India, Bangladesh, Cambodia, and parts of Indonesia and Malaysia, having been extirpated from Myanmar, Thailand, Vietnam, and Singapore. The southern roofed turtle (B. affinis) occurs in Cambodia, Sumatra, and the Malay Peninsula. The yellow-headed temple turtle (Heosemys annandalii) is found in Cambodia, Laos, the Malay Peninsula, Vietnam and possibly Myanmar, where it has been greatly reduced in number due to overcollection for Asian food markets. The Burmese python (Python bivittatus) is found in northeastern India (where it is known only from two small, disjunct areas), through Nepal and southern China (including the island of Hainan) and throughout most of mainland South East Asia. It is absent from the Malay Peninsula, Borneo and Sumatra, but occurs on Java, Bali, Sumbawa and possibly Lombok as well as in southern Sulawesi. Although widely distributed it is everywhere declining due to overcollection for its meat and leather, as well as by loss of habitat. The species has been introduced and established in southern Florida, USA via the pet trade, where it has become a serious threat to the native fauna.

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The king cobra (Ophiophagus hannah) remains relatively common across South and South East Asia, but is threatened by loss of habitat. As the world’s largest venomous snake, it is also highly persecuted by humans. Boulenger’s cross frog (Oreophryne monticola) is known only from the mountains of Bali and Lombok in Indonesia. Volcanic activity is a particular threat on the latter island. The Tasan frog (Alcalus tasanae) is known only from a few disjunct areas of western and southern Thailand but may occur as well in southern Myanmar. The giant freshwater whipray (Urogymnus polylepis), one of the world’s largest freshwater fish, was historically found in large rivers and estuaries across southern and South East Asia, including the island of Borneo and possibly western Java. Overfishing and habitat destruction have since reduced it to a few disjunct localities. The longnose marbled whipray (Fluvitrygon oxyrhyncha) is a very rare species known only from a few specimens collected from Cambodia, Thailand, southern Vietnam and Borneo. The white-edge whipray (F. signifer) is found widely but patchily in Thailand, peninsular Malaysia, Sumatra and Borneo. The Asian bony-tongues (Scleropages) are long-bodied fish that inhabit blackwater rivers, forested swamps and wetlands throughout much of South East Asia, although in very low densities. The Sunda bony-tongue (S. formosus) occurs in the Mekong drainage of Vietnam and Cambodia, as well as in south-eastern Thailand, the Malay Peninsula, Borneo, and Sumatra, with an additional introduced population on Singapore. It is threatened by overcollection for the international aquarium trade and habitat destruction. The Burmese bony-tongue (S. inscriptus) is known only from captive individuals that may have been either wild-caught or farm-bred. The species has been reported from the Great Tenasserim River drainage of peninsular Myanmar and from Pedu Lake, western Malaysia, with further anecdotal reports from rivers on the Andaman coast of Thailand. Jullien’s golden carp (Probarbus jullieni) historically occurred in the Mekong, Chao Phraya and Mae Klong river drainages of Thailand, Vietnam, Cambodia and Laos, as well as the Pahang, Terengganu and Perak rivers of peninsular Malaysia. The species has become rare or extirpated in many areas due to loss of habitat and damming, although it is also being bred commercially in Thailand and Laos, where it is considered a delicacy. The giant sharkminnow (Osteochilus schlegelii) is found patchily in Indochina, peninsular Malaysia, Sumatra, and Borneo.

The Oriental Region The Oriental Region includes all of the mainland areas of the Indo-Malaysian Realm along with the islands of Sri Lanka, Hainan, and numerous smaller islands. The Indian elephant (Elephas maximus indicus) historically ranged from West Asia along the Iranian coast into the Indian

The Oriental Region Figure 5.1 Northern two-horned rhinoceros in the London Zoo, 1904. (Credit: London Zoo.)

subcontinent and Indochina as far north and east as the Yangtze River. They have long been extinct in Iran, Pakistan and most of China, and in serious retreat everywhere else for centuries due to hunting for their ivory and conflict with humans. The Indian or greater one-horned rhinoceros (Rhinoceros unicornis) once ranged across the entire Indo-Gangetic Plain from northern Pakistan to at least the India/Myanmar border and including parts of Nepal, Bhutan, and Bangladesh. It may also have existed in Myanmar, southern China and Indochina, although this is uncertain. The species was common in northwestern India and Pakistan until around 1600, but disappeared from this region shortly thereafter. It declined sharply in the remainder of its range between 1600 and 1900 due to excessive hunting and agricultural development, until at last it found itself on the brink of extinction at the beginning of the twentieth century, with less than 200 remaining. Currently it survives only in a few small subpopulations in northern India (West Bengal, Uttar Pradesh and Assam) and southern Nepal, mainly in remote riverine swamps with tall, dense ‘elephant grass’. While not the rhinos’ preferred habitat (they seem to favour dry forest with open stretches of grassland), these areas have served as a last refuge for the species and are now mostly protected. Numbers have fluctuated and occasionally suffered dramatic declines, mainly due to the illegal settlement of squatters in protected areas, as well as poaching. In 2007 the total was estimated at 2575, with the majority living in Kaziranga National Park (Assam). An attempt to reintroduce a pair into Lal Sohanra National Park, Pakistan in 1983 was unsuccessful. The Asiatic lesser one-horned rhinoceros (R. sondaicus), previously discussed, is divided into three

subspecies historically found across South East Asia. The Indochinese lesser one-horned rhinoceros (R. s. annamiticus) was found in Vietnam, Laos, Cambodia and eastern Thailand. Heavily hunted, it was thought to be extinct until a small population was unexpectedly rediscovered in Cat Tien National Park, Vietnam in the 1990s. Sadly, the location of these animals was unwisely made widely known in the press and the subspecies was quickly hunted into certain extinction by poachers. The Indian lesser one-horned rhinoceros (R. s. inermis) was found in north-eastern India (West Bengal), Bangladesh, and Myanmar. It was almost certainly driven to extinction before 1925, although it is just possible that a small number may survive in Myanmar. The northern two-horned rhinoceros (Dicerorhinus sumatrensis lasiotis) once occurred in north-eastern India, Bangladesh and Myanmar, but has been declared extinct in these countries. Unconfirmed reports suggest a small population may still survive in northern Myanmar, but the political situation in that country has prevented verification. The kouprey (Bos sauveli) is (or was) a large, little-known forest ox historically found in Cambodia, southern Laos, south-eastern Thailand, and western Vietnam. Only discovered by science as late as 1937, it was intensively hunted and by the mid-1950s had disappeared from Thailand and Laos but was still found in Vietnam and Cambodia. At the end of the 1960s it was restricted to two areas of open deciduous forest and clearings on either side of the Mekong River in north-eastern Cambodia. Efforts to save the species were long discussed but little came of it, and it appears to have been a victim of the war in Indochina. There have been no confirmed

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Figure 5.2 A young male kouprey in the Vincennes Zoo, Paris, in 1937. (Credit: Vincennes Zoo, Paris.)

sightings since 1970, but the animals are nocturnal and notably skittish and a few may survive in remote areas. The gaur (B. gaurus), also known as the Indian bison, is the largest living cattle species. Two subspecies are generally recognized. The Indian gaur (B. g. gaurus) historically occurred throughout the Indian subcontinent but is now confined to scattered areas of south-western and centraleastern India, Bhutan and Nepal, having been extirpated in Bangladesh and on the island of Sri Lanka. The Indochinese gaur (B. g. laosiensis) is found patchily in southern China, Myanmar, Laos, Vietnam, Cambodia, Thailand and peninsular Malaysia. Both have been greatly reduced by ruthless slaughter, commercialized hunting and loss of habitat. In the 1930s large herds still grazed by night in open clearings along forest edges, with forest itself serving as their refuge. During World War II great numbers were killed, and populations have never really recovered. Diseases transmitted by domestic cattle, particularly rinderpest and foot-and-mouth disease, are also a serious threat and have occasionally come close to wiping out entire populations of gaur. A third type of wild cattle, the Burmese banteng (B. javanicus birmanicus), can still be found in Myanmar, Thailand, Cambodia, Laos, and Vietnam, and perhaps in north-eastern India (Manipur) as well. The largest single subpopulation, about 4600, lives in eastern Cambodia. The khting-vor (Pseudonovibos spiralis) is a mysterious bovid reputed to exist in Cambodia and Vietnam, and known only from a few curiously twisted horns that may be nothing more than artificially crafted cattle horns for use as talismans. However, there is a fairly reliable report of British hunters from the early twentieth century having observed the living creatures, and even shooting two of them for use as tiger bait. It is included here on a provisional basis, but its existence must be considered highly questionable. Two subspecies of tiger (Panthera tigris) historically inhabited the Oriental Region. The Indochinese tiger

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(P. t. corbetti) lives in mountainous and hilly areas of Laos, Myanmar, Thailand and Vietnam. It formerly occurred in Cambodia and south-western China, but is believed to be extinct in those countries. In 2010 the total world population was estimated at about 350 individuals. The Bengal tiger (P. t. tigris) is the most common subspecies of tiger and still lives in a wide range of habitats, from grasslands, rainforests, scrub and dry forests, and mangroves. At the end of the nineteenth century it ranged across South Asia from Pakistan to China, and it was estimated that probably about 40,000 of them roamed the Indo-Pakistan part of that range. That number had dropped to 2000–4000 owing to excessive shooting, poisoning, habitat destruction, and reduction of prey 50–60 years later. Today they are confined to a few suitable areas of Bangladesh, Bhutan, India and Nepal. There are a tremendous number in captivity. The Asiatic lion, long considered to be a distinct subspecies, is now generally thought to be a relict population of African lion (Panthera leo leo). Historically, outside of subSaharan Africa lions ranged across northern Africa and southern and eastern Europe throughout almost all of south-western Asia, from the Arabian Peninsula and Asia Minor, as far as India, where they were widespread across the northern half of the subcontinent. Herodotus reported almost 2500 years ago that there were many in Thrace, and declared that during the march of the Persian king Xerxes through Macedonia (480 B . C .) some of the baggage camels were killed by lions. Later, Aristotle (384–322 B . C .) assigned the same range to the lion, but referred to it as rare. By A . D . 80–100 the lion was considered to be entirely extirpated in Europe. The reason for the disappearance must be traced to the unceasing warfare man has carried on against this animal. Neither the climate nor the food supply could have been important factors in the elimination of the lion at that time. Until the nineteenth century lions still occurred in eastern Turkey, Iran, Mesopotamia, and from east of the Indus River to Bengal and the Narmada River in central India. As late as 1866 they were described as ‘not rare’ in Asia Minor, but hunting, loss of habitat and persecution led to their extermination through most of these areas within the last century and a half. In Iran they seem to have been wiped out during the 1930s – the last occurrence outside of India apart from an isolated population in the High Atlas Mountains of North Africa, where they perhaps survived up to the 1940s. Meanwhile, in India human occupation of the land and a tremendous slaughter by the military exterminated lions everywhere except in the Gir Forest, on the Kathiawar Peninsula of Gujarat, north-western India. Since 1884 they have been entirely confined to this small teak forest and to nearby open thorn scrub and scattered acacias among light stands of grass. The forest was officially protected to ensure the survival of the Asiatic lion, but continued to be utilized by thousands of squatters and tens of thousands of domestic animals. Nevertheless, the lion population remained stable despite the killing of around 100 annually. In 1936 there were thought to be 287 lions, in 1955, 290, and in 1963, 255. In 1968, however,

The Oriental Region

only 175 still survived, and in 1970 less than 150. Three Asiatic lions had also been introduced in the Chandraprabha Sanctuary in Uttar Pradesh in 1957, which by 1966 had increased to 11; by 1969 all of them had been killed. Today, this last vestige remains entirely confined to what is now Gir Forest National Park, where numbers have increased in recent years to around 650 as of 2017. As a single population they remain vulnerable to threats such as epidemics and forest fires, and some poaching continues. Reintroductions to other areas of India and Iran have long been discussed. The Indochinese leopard (Panthera pardus delacouri) is found in mainland South East Asia and southern China. The Indian leopard (P. p. fusca) is confined to the Indian subcontinent, Myanmar and southern Tibet. Both are threatened by loss of habitat and prey base and by human persecution. The Burmese wild dog (Cuon alpinus adjustus) is confined to north-eastern India and northern Myanmar, where it is threatened by loss of habitat, depletion of its prey base and human persecution. The sloth bear (Melursus ursinus) is a lanky, shaggy, mainly insectivorous species native to South Asia. The Indian sloth bear (M. u. ursinus) formerly occurred in a wide variety of habitats across India, Bangladesh and the southern lowlands of Nepal and presumably Bhutan, where it is now mostly confined to dry, sparsely forested areas. Habitat destruction is the chief reason for the decline, although human–bear conflicts often result in persecution. Several species and subspecies of crested gibbon (Nomascus) are seriously threatened by loss of habitat, hunting for food and ‘traditional medicine’ and live capture for use as pets. The northern white-cheeked crested gibbon (N. leucogenys) is known only from a few localities in extreme southwestern China (Yunnan), northern Laos and northern Vietnam. The southern white-cheeked crested gibbon (N. siki) is found patchily in central Laos and central Vietnam. The northern yellow-cheeked crested gibbon (N. annamensis) is found patchily in southern Laos, south-central Vietnam and north-eastern Cambodia. The southern yellow-cheeked crested gibbon (N. gabriellae) is found patchily in southern Vietnam and eastern Cambodia. The Laotian black crested gibbon (N. concolor lu) is confined to a small area in northwestern Laos (Luang Namtha and Bokeo provinces). Several species and subspecies of hoolock gibbon (Hoolock) are threatened by loss of habitat and hunting. The western hoolock gibbon (H. hoolock) is divided into two subspecies. The nominate form (H. h. hoolock) is found over a relatively wide area of north-eastern India (Arunachal Pradesh, Assam, Manipur, Meghalaya, Mizoram, Nagaland and Tripura), eastern Bangladesh, north-western Myanmar and possibly southwestern China (Tibet). The Mishmi Hills hoolock gibbon (H. h. mishmiensis) is confined to a small area of north-eastern India (Arunachal Pradesh and Assam). The eastern hoolock gibbon (H. leuconedys) is found in northern Myanmar between the Chindwin and Irrawaddy rivers. The tianxing hoolock gibbon (H. tianxing) occurs over a wide area of

Myanmar between the Irrawaddy and Salween rivers, and in extreme south-western China (Yunnan). The pileated gibbon (Hylobates pileatus) is found in southeastern Thailand, western Cambodia and parts of southwestern Laos, where it is threatened by hunting and habitat destruction. The golden long-tailed macaque (Macaca fascicularis aureus) is found in southern Bangladesh, Laos, Myanmar (including the Mergui Archipelago) and west-central Thailand, but is everywhere threatened by hunting and habitat destruction. Three species of douc (Pygathrix), brightly coloured leafeating monkeys, have long been persecuted by humans. During the Vietnam War their lowland and montane rainforest habitat was heavily bombed and sprayed with defoliants such as Agent Orange, and it is said that soldiers also occasionally used the animals for target practice. Since then, deforestation and hunting for food and ‘traditional medicine’ are the primary threats. The red-shanked douc (P. nemaeus) is found patchily in eastern Laos, central and north-central Vietnam and north-eastern Cambodia. The grey-shanked douc (P. cinerea) is largely confined to the Central Highlands of south-central Vietnam, extending slightly into north-eastern Cambodia. The total population is thought to be less than 2000. The black-shanked douc (P. nigripes) occurs in eastern Cambodia and southern Vietnam. Several leaf-eating monkeys of the genus Trachypithecus are threatened by hunting and habitat destruction. The Tenasserim langur (T. barbei) is known only from a few localities in south-western Thailand and adjacent northeastern Myanmar. Phayre’s langur (T. phayrei) is divided into two subspecies. The western Phayre’s langur (T. p. phayrei) is found in eastern Bangladesh, north-eastern India (Assam, Mizoram and Tripura) and western Myanmar. The Shan State Phayre’s langur (T. p. shanicus) occurs in south-western China (Yunnan) and in northern and eastern Myanmar. The Indochinese grey langur (T. crepusculus) occurs in south-western China (Yunnan), Laos, Myanmar, Thailand and northern Vietnam. The Laotian black langur (T. laotum) is confined to a small area of central Laos (Bolikhamxai and Khammouane provinces). The Ha Tinh langur (T. hatinhensis) is found in the limestone forests of north-central Vietnam (Ha Tinh, Quang Binh, and Quang Tri provinces) and east-central Laos (Khammouane and Savannakhet provinces). Germain’s langur (T. germaini) is still found throughout a fairly wide area of mainland South East Asia, but is everywhere declining. The orange-bellied capped langur (T. pileatus durga) from north-eastern India and Bangladesh, and the blond-bellied capped langur (T. p. pileatus) from north-eastern India (Arunachal Pradesh, Assam, Meghalaya, and Nagaland) and north-western Myanmar have both suffered population declines. The pygmy slow loris (Nycticebus pygmaeus) is a small, nocturnal prosimian found east of the Mekong River in Vietnam, Laos, eastern Cambodia and southern China (Yunnan). Its forest habitat was greatly reduced during the

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Vietnam War due to extensive burning, clearing and defoliation. Today, continuing habitat destruction and hunting for use in ‘traditional medicine’ are the main threats. The grey slender loris (Loris lydekkerianus) remains widespread within the forests of India and Sri Lanka, where it is divided into a number of subspecies. The Mysore grey slender loris (L. l. lydekkerianus) is found in southern and eastern India (Andhra Pradesh, Karnataka and Tamil Nadu). The Malabar grey slender loris (L. l. malabaricus) occurs in south-western India (Karnataka, Kerala and Tamil Nadu). Both are threatened by loss of habitat, hunting for ‘traditional medicine’ and live collection for use as pets. Two subspecies of brow-antlered deer (Rucervus eldii) are found in the dry forests, scrubs and grasslands of southern Asia (a third lives in wetland areas). The Burmese browantlered deer (R. e. thamin) was historically abundant on the central plains of Myanmar as well as in westernmost Thailand but was so depleted by hunting and habitat destruction that, by the 1940s, it was near extinction. The creation of sanctuaries saved it and by the mid-1980s it was once again widespread. Unfortunately, the population has since declined once more due to hunting for trophies as well as ‘traditional medicine’, and today it is once again seriously threatened. The Thai browantlered deer (R. e. siamensis) was historically found in the lowlands of Thailand, Cambodia, Laos, and Vietnam, with outlying populations in southern China and on the island of Hainan. By 1970 only a few small herds survived in northern Thailand, all other populations having been wiped out by hunters. In 2004 the wild population was estimated ‘in the low tens’. Fortunately, a longstanding captive breeding programme initially begun by the Vincennes Zoo, Paris, will at least continue to serve as safeguard against total extinction. The barasingha (Rucervus duvaucelii) is a type of large deer historically found throughout the Indian subcontinent from north-eastern Pakistan to Bangladesh, but which has been extirpated from most of its range due to unregulated hunting and habitat destruction. The hard-ground barasingha (R. d. branderi), the most seriously threatened of the three subspecies, survives only in Kanha National Park in central India (Madhya Pradesh), where in 2016 the total population was around 750. Two subspecies of sambar deer (Rusa unicolor), previously discussed, are threatened by habitat destruction and hunting. The Indian sambar deer (R. u. unicolor) is found in India, Bangladesh and Sri Lanka. The Indochinese sambar deer (R. u. cambojensis) is confined to parts of mainland South East Asia. The blackbuck (Antilope cervicapra) was historically one of the most common and characteristic of ungulates on the dry scrublands of India and Pakistan, with herds of 10,000 said to have occurred in the Punjab. Sadly, within a few decades, merciless hunting had brought it to the verge of extinction. Today only small, scattered herds are to be seen, which are largely confined to protected areas. However, large numbers have been introduced on hunting ranches worldwide,

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particularly in the United States, and for that reason the species is considered to be out of danger. The four-horned antelope or chousingha (Tetracerus quadricornis) is a small species found widely but patchily in India and, at least formerly, in the lowlands of Nepal. It is threatened by loss of habitat. The hog deer (Axis porcinus) is a type of small deer historically found over much of mainland South and South East Asia, with introduced populations in parts of Australia and the United States (Texas, Florida and Hawaii) and another in Sri Lanka that is either native or introduced. The two recognized subspecies have been much reduced by hunting and habitat destruction. The Indian hog deer (A. p. porcinus) is now confined to a few isolated populations in eastern Pakistan, northern India, Nepal, Bhutan, Bangladesh, Myanmar, and possibly south-western China (Yunnan). The Indochinese hog deer (A. p. annamiticus) has been almost entirely extirpated throughout its former range, although small numbers still survive in Cambodia and another population of unknown taxonomic origin has been reintroduced into north-central Thailand. The silver-backed chevrotain (Tragulus versicolor) is a little-known type of ‘mouse-deer’ that was first described in 1910 on the basis of four specimens collected from what is now central Vietnam. It was not recorded again until 1990, but was afterwards feared extinct as a result of hunting and habitat destruction. In 2019, however, camera traps confirmed its survival in southern coastal Vietnam. Williamson’s chevrotain (T. williamsoni) is known only from a single specimen collected in north-western Thailand. It is thought that the species may be found more widely in the highlands of South East Asia, in particular southern China and northern Vietnam. The leaf muntjac (Muntiacus putaoensis), standing as it does only about 60 cm in height, is considered the smallest true deer in the world. Only just discovered in 1997 in northern Myanmar (Kachin state), it is now known to occur as well in north-eastern India (Arunachal Pradesh and Nagaland). The species is believed to be decreasing in number due to hunting by local people. The giant muntjac (Megamuntiacus vuquangensis) is found throughout the Annamite Mountains and associated hill ranges in central and south-central Vietnam, eastern Laos and (marginally) eastern Cambodia. It is threatened by intense hunting pressure. Heude’s warty pig (Sus bucculentus) is (or was) a type of wild pig known only from two skulls collected from southern Vietnam in 1892. It was feared extinct until 1995, when the skull of a freshly killed specimen was discovered in the Annamite Mountains of central Laos. The Bhutan binturong (Arctictis binturong albifrons) is a type of large viverrid that was first described in 1822 based on a drawing by Alfred Duvaucel. In recent years it has been camera-trapped in Royal Manas National Park, and is thought to be present in other parts of southern Bhutan where habitat remains largely intact. The Thai binturong (A. b. gairdneri)

The Oriental Region

was first described in 1916 from a skull collected in southwestern Thailand. Owston’s palm civet (Chrotogale owstoni) is found widely in Laos, Vietnam and a small area of southernmost China (Yunnan), but is everywhere threatened by hunting and loss of habitat. The Indian pangolin (Manis crassicaudata) remains widespread in South Asia from northern and south-eastern Pakistan through much of India south of the Himalayas, southern Nepal, Sri Lanka, and parts of Bangladesh. Like all pangolins it is everywhere threatened by collection for use in Chinese ‘traditional medicine’, as well as local consumption of its meat. The long-eared gymnure (Hylomys megalotis) is a hedgehog-like mammal known only from limestone karst forests in central Laos (Khammouane province). The lesser marmoset rat (Hapalomys delacouri) has a patchy distribution in South East Asia, where very few specimens have ever been collected. Delacour’s lesser marmoset rat (H. d. delacouri) is known from northern Laos, Vietnam and southern China (southern Yunnan and Guangxi). Ranjini’s field rat (Rattus ranjiniae) is known only from three widely separated localities in south-western India (Kerala). The Myanmar flying fox (Pteropus intermedius) is known only from two roosting colonies in northern peninsular Myanmar. Lyle’s flying fox (P. lylei) is found in southern and central Cambodia, southern Thailand, and southern Vietnam, with a further outlying population in southern China (Yunnan). It is threatened by the loss of large roosting trees and by hunting. Wroughton’s mastiff bat (Otomops wroughtoni) was, until recently, known only from a single cave in the Western Ghats of south-western India (Karnataka). It has since been recorded from four localities in north-eastern India (Meghalaya), as well as from northern Cambodia, suggesting a much larger range than previously thought. Joffre’s pipistrelle bat (Hypsugo joffrei) is known only from a few specimens collected over a wide area in northern and north-western Myanmar, India, Nepal, and Vietnam. The mitred horseshoe bat (Rhinolophus mitratus) is known only from a single specimen collected in 1844 from north-eastern India (Jharkhand). The sarus crane (Antigone antigone), the world’s tallest flying bird, is a non-migratory species found disjunctly in South and South East Asia and in northern Australia. The Indian sarus crane (A. a. antigone) inhabits northern and central India, Nepal and, at least historically, parts of eastern Pakistan. Vagrants are occasionally seen in Bangladesh. The South East Asian sarus crane (A. a. sharpii) was also formerly widespread but is now confined to a few areas of Cambodia, southern Laos, southern Vietnam and southern Myanmar. Both subspecies have suffered significant declines due to loss of habitat, hunting, egg and chick collection, and pesticides. The Indian vulture (Gyps indicus) was historically very common in south-eastern Pakistan and peninsular India south

of the Gangetic plain, but since the mid-1990s has suffered a catastrophic decline throughout its range due to feeding on animal carcasses treated with the veterinary drug diclofenac. Attempts at banning the drug have thus far been unsuccessful. The Gingee Egyptian vulture (Neophron percnopterus ginginianus) is confined to the Indian subcontinent where it has suffered a rapid population decline mainly resulting from secondary poisoning (i.e. consuming the carcasses of livestock that have been treated with a veterinary medicine fatal to vultures). The Indian spotted eagle (Clanga hastata) is found widely but patchily across southern Asia and Indochina, but is everywhere rare and declining due to loss of habitat. The forest owlet (Heteroglaux blewitti) is a type of small owl confined to a few localities in west-central India. First described in 1873, it was long known only from a handful of specimens and not seen after 1884 until finally rediscovered in 1997. Other populations have been found since but the species remains seriously threatened, with a total population estimated in 2015 at less than 250. The Indian skimmer (Rynchops albicollis) is a tern-like bird that was historically widely distributed and common across the Indian subcontinent as well as along the major rivers of Myanmar and the Mekong in Indochina. The species has suffered a massive decline due primarily to habitat destruction and is now confined to a few sporadic colonies. The rufous-necked hornbill (Aceros nipalensis) was historically widespread in South and South East Asia but has declined dramatically due to hunting and loss of habitat. It is now very rare across much of its range and is thought to have been extirpated from Nepal and many areas in Thailand, and is close to disappearing from Vietnam. The Manipur bush-quail (Perdicula manipurensis) is a rare and elusive species known primarily from historical accounts originating in north-eastern India (West Bengal, Assam, Manipur, Nagaland and Meghalaya) and possibly the Chittagong Hill tracts of eastern Bangladesh. There was a gap in records after 1932 broken only by a single unconfirmed report in 1998, until its rediscovery within Manas National Park, Assam in 2006. Given the lack of more recent records and the small amount of remaining grassland habitat available, any remaining populations must be very small and fragmented. Two ill-defined subspecies (P. m. manipurensis and P. m. inglisi) have been described, separated by the Brahmaputra River. The Nilgiri wood pigeon (Columba elphinstonii) is found widely but patchily throughout southern and south-western India (Maharashtra, Goa, Karnataka, Kerala and Tamil Nadu), but is everywhere rare and declining. The pale-capped pigeon (C. punicea) was also historically widespread in South and South East Asia but is now confined to a few scattered pockets in north-eastern India, Bangladesh, Myanmar, Thailand, Laos, Cambodia and Vietnam. Both have declined due to heavy hunting pressure and loss of habitat. The dark-rumped swift (Apus acuticauda) is known only from a few breeding colonies in the Himalayan foothills of

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Bhutan and in the hills of north-eastern India (Meghalaya, Nagaland and Mizoram), from where it migrates to southern China (Yunnan) and western Thailand. In 2018 the total population was thought to be around 1000. The white-eyed river martin (Eurochelidon sirintarae) is known only from a single wintering ground (the reedbeds around Lake Boraphet in central Thailand). Not reported with certainty since 1978 despite targeted surveys, it may be extinct, although it is possible that it might still survive in isolated areas of Myanmar. The silver oriole (Oriolus mellianus) breeds in southern China (Sichuan, Guizhou, Guangxi and Guangdong), from where it winters in Thailand and western Cambodia. The total population is very small and continues to decline due to loss of habitat. The green avadavat (Amandava formosa) is a type of finch found in central India, where it is rapidly declining due to trapping for the cage-bird trade, compounded by habitat loss and degradation. The grey-crowned prinia (Prinia cinereocapilla) is a type of passerine bird found patchily in north-eastern India (Uttar Pradesh, West Bengal and Assam), Nepal and Bhutan. The yellow-throated bulbul (Pycnonotus xantholaemus) is a type of passerine bird found in southern peninsular India (Andhra Pradesh, Karnataka, Kerala and Tamil Nadu). Populations are small and severely fragmented. The white-naped tit (Machlolophus nuchalis) is a rare species found disjunctly in scattered pockets within northwestern and south-eastern India. It continues to decline due to loss of habitat. The broad-tailed grassbird (Schoenicola platyurus) is a type of warbler that was formerly common in the hills and mountains of south-western India, with a few additional unconfirmed records from Sri Lanka (presumably vagrants). The species has undergone a significant decline due to the mismanagement and destruction of native shola grasslands. The Burmese star tortoise (Geochelone platynota) was historically widespread in the dry forests and shrublands of central Myanmar, but is thought to have become extinct in the wild by the beginning of the twenty-first century owing to overcollection for local subsistence and export to Chinese food markets. Captive-bred specimens have since been reintroduced into two closely guarded protected areas (Shwesettaw and Minsontaung wildlife sanctuaries). The Indian star tortoise (G. elegans) is found patchily in dry areas of south-eastern Pakistan, north-western, eastern and southern India, and Sri Lanka. It is threatened by loss of habitat and collection for the international pet trade. Phayre’s Asian giant tortoise (Manouria emys phayrei) occurs from peninsular Thailand north through Myanmar to the north-eastern Indian and eastern Bangladeshi hill tracts. It has suffered considerable declines due to loss of habitat and collection for use as food. Bourret’s box turtle (Cuora bourreti) is a rare, primarily terrestrial species found in central Vietnam and adjoining

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south-eastern Laos (Savannakhet province). The Indochinese box turtle (C. galbinifrons) occurs in southern China (Guangxi and the island of Hainan), northern Vietnam and north-eastern Laos. The keeled box turtle (C. mouhotii) is divided into two subspecies found widely, if disjunctly, over much of South and South East Asia. The nominate form (C. m. mouhotii) is known from northern Laos, northern Vietnam and southern China (Guangdong, Guangxi, Hunan, Yunnan and the island of Hainan). Populations inhabiting south-eastern Bangladesh, north-eastern India (Arunachal Pradesh, Assam, Meghalaya and Mizoram), northern Myanmar and Bhutan may represent either this form or perhaps a separate taxon. Obst’s keeled box turtle (C. m. obsti) is known only from central Vietnam and possibly adjoining central Laos. All are seriously threatened by overcollection for use as food and pets. Annandale’s bent-toed gecko (Cyrtodactylus annandalei) is known only from two localities in western Myanmar. The black and white spitting cobra (Naja siamensis) is found widely in Thailand, Cambodia, Laos, southern Vietnam and eastern Myanmar, but is everywhere threatened by overcollection for use in Chinese ‘traditional medicine’. The Mandalay spitting cobra (N. mandalayensis) is known only from a few individuals collected from the dry central zone of Myanmar. The black-barred keelback (Rhabdophis callichroma) is a type of snake known only from northern Vietnam and the island of Hainan, where it has not been collected in nearly a century. The scarce bridal snake (Dryocalamus gracilis) is known only from a small number of specimens collected over a wide area of southern India, Sri Lanka, and Myanmar. The graceful racer (Platyceps gracilis) is found patchily in the dry forests and shrublands of western and north-western India (Gujarat, Madhya Pradesh, Maharashtra and Rajasthan). Phipson’s earth snake (Uropeltis phipsonii) is known only from a few localities in western coastal India (Maharastra and Goa). The Malabar toad (Duttaphrynus hololius) is known only from a few scattered localities in south-eastern India (Andhra Pradesh). The Charpa tree frog (Polypedates occidentalis) is known only from the Charpa Forest in south-western India (Kerala). The Dudhwa foam-nest tree frog (Chirixalus dudhwaensis) is known only from Dudhwa National Park in northern India (Uttar Pradesh). Quyet’s foam-nest tree frog (Gracixalus quyeti) is known from a small area of lowland and montane forest in northcentral Vietnam (Quang Binh and Ha Tinh provinces). Jerdon’s shrub frog (Philautus jerdonii) is known only from its original collection during the nineteenth century from north-eastern India (West Bengal). The Khao Yai odorous frog (Odorrana indeprensa) is known only from two small areas of south-central Thailand (Nakhon Ratchasima and Nakhon Nayok provinces), centered on Khao Yai National Park.

The Oriental Region

Boulenger’s golden-backed frog (Indosylvirana aurantiaca) is known from the Western Ghats of south-western India and from Sri Lanka. The Chengalam caecilian (Uraeotyphlus interruptus) is a fossorial amphibian known only from a few specimens collected from a rubber plantation in south-western India (Kerala). The Kuttal caecilian (Chikila fulleri) is known only from a single specimen collected at the beginning of the twentieth century in north-western India (Assam). The Mae Wang Valley caecilian (Ichthyophis acuminatus) is known only from the Mae Wang Valley in north-western Thailand (Chiang Mai province). The moustached caecilian (I. moustakius) is known only from a small area of northeastern India (Manipur). The Sendenyu striped caecilian (I. sendenyu) is known only from a few specimens collected within a banana plantation in north-eastern India (Nagaland). The Khumhzi striped caecilian (I. khumhzi) is known only from two closely situated localities in northeastern India (Manipur).

Mountains and Highlands There are high mountains in south-western India and throughout South East Asia. Myanmar (formerly Burma) in particular has range after range of rugged mountains, some of which have been denuded of forests and lie bare, while others remain cloaked in forests. The latter lose their leaves with a marked seasonality and therefore differ from true rainforests, with their distribution in mainland and insular Asia corresponding to the pattern of seasonal monsoon winds (bringing rainfall and moisture) and high summer temperatures. The monsoon forests of tropical Asia are among the most important habitats left for wildlife. Unfortunately, they are disappearing rapidly. Owing to heavy population pressure, cultivation, and grazing as well as improper use of land, many monsoon montane forests have been destroyed and have given way to the results of erosion. Serious landslides are common on devastated mountain slopes, violent floods are an annual occurrence, and the water table in valleys and plains is sinking. The deforestation of these mountains thus releases formidable challenges not only to agriculturists and conservationists but also to whole populations. The red serow (Capricornis rubidus) is a type of goatantelope known only from northern (and possibly western) Myanmar, where it is threatened by overhunting. Roosevelts’ muntjac (Muntiacus rooseveltorum) was long known only by a single specimen collected from northern Laos (Phongsali province) in 1929. In recent years, however, skulls along with a few sightings originating within the mountains of northern Laos and north-western Vietnam perhaps pertain to the species. Hume’s rat (Hadromys humei) is known only from three isolated localities in north-eastern India (Assam and Manipur).

The highland pygmy white-toothed shrew (Suncus montanus) is found widely but patchily in the Western Ghats of south-western India and in the highlands of Sri Lanka. Kelaart’s long-clawed shrew (Feroculus feroculus) is found widely but patchily in the Western Ghats of south-western India and in the highlands of Sri Lanka. The Surat serotine bat (Eptesicus dimissus) is known only from a single specimen collected in southern Thailand (Surat Thani province) and a few others from southern Nepal. The chestnut-eared laughingthrush (Ianthocincla konkakinhensis) is known from Kon Ka Kinh National Park in the Central Highlands of south-central Vietnam (Kon Tum province), and possibly adjacent areas of south-eastern Laos. The impressed tortoise (Manouria impressa) is widespread within the mountains of mainland South and South East Asia, but is everywhere threatened by hunting for food. The three-keeled hill turtle (Melanochelys tricarinata) is a little-known species from north-eastern India, Nepal, and Bangladesh. Fea’s bent-toed gecko (Cyrtodactylus feae) appears to be confined to the mountains of south-eastern Myanmar and possibly adjacent north-western Thailand. Boulenger’s ground skink (Kaestlea laterimaculata) is found widely but patchily in the both the Western and Eastern Ghats of southern India. Beddome’s coral snake (Calliophis beddomei) is known only from a few specimens collected from three mid-elevation localities in southern India (Karnataka and Tamil Nadu). Spencer’s mountain stream snake (Opisthotropis spenceri) is known only from a single specimen collected during the early twentieth century in north-western Thailand (Lampang province). Boomsong’s stream snake (Parahelicops boonsongi) is known only from three specimens collected in north-eastern Thailand (Loei province). Hampton’s green snake (Cyclophiops hamptoni) is known only from a single specimen collected in central Myanmar in the early twentieth century. The Rapp’s stripe-necked snake (Liopeltis rappi) is known only from a small area of north-eastern India and possibly neighbouring Nepal. Delacour’s mountain snake (Plagiopholis delacouri) is known only from disjunct historical records originating in northern Vietnam (Lao Cai and Vinh Phuc provinces) and northern Laos (Xiangkhouang province). The Garo Hills shrub frog (Philautus garo) is known from the Garo Hills on the Shillong Plateau of north-eastern India (Assam and Meghalaya) and from the Purvanchal Range (Nagaland). The gold-flanked odorous frog (Odorrana aureola) is known only from Phu Kradueng National Park and the Phu Luang Wildlife Sanctuary in north-central Thailand (Loei province). The musical slender litter frog (Leptolalax melicus) is known only from Virachey National Park in north-eastern

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Cambodia (Ratanakiri province), but may also occur in adjacent areas of southern Laos and southern Vietnam. It is threatened by logging and dam construction. The Laotian warty newt (Laotriton laoensis) is confined to the mountains of northern Laos, where it is threatened by overcollection by the international pet trade as well as use in ‘traditional medicine’.

The Western Ghats The Western Ghats are an old, isolated, low-level mountain range running parallel to the coast of south-western India (Maharashtra, Karnataka, Kerala and Tamil Nadu). Once covered in moist deciduous and montane rainforests, they still contain a large proportion of the subcontinent’s wildlife species and are among the most important biological hotspots on Earth. Sadly, the forests of the coastal lowlands are now completely gone, and those of the higher elevations (known as sholas) are highly fragmented. The lion-tailed macaque (Macaca silenus), one of the most threatened monkeys in Asia, is confined to the moist forests of the Western Ghats from the Kalakkadu Hills north to Ashi Ghat in the states of Karnataka, Kerala, and Tamil Nadu. During the twentieth century its range became increasingly isolated and fragmented by loss of habitat and numbers dropped to alarming levels, in part due to hunting as well. Thanks to decades of protection within a series of national parks and wildlife sanctuaries the population has grown once more, although it is still thought to be less than 4000. The Nilgiri langur (Semnopithecus johnii) is found, rather unevenly, in the hill country of the Western Ghats from the Aramboli Pass near the southern tip of India north to Srimangala. Hunting and habitat destruction remain a threat, but the population is thought to be relatively stable. The Nilgiri tahr (Nilgiritragus hylocrius) is confined to small pockets of high-elevation grassy slopes and cliffs in the Western Ghats of Kerala and Tamil Nadu, where the total population is currently less than 2500. Presumably the species had a much wider range in the past, because it has relatives in the Himalayas and the Arabian Peninsula. Excessive hunting is believed to be the cause of the decline. The Malabar civet (Viverra civettina) is another species thought to be endemic to the Western Ghats, although the few historical records lack precise locality information. Already considered rare or possibly even extinct by 1970, there were only two unconfirmed sightings until the late 1980s, when skins of recently killed civets (since lost) were obtained from hunters. The absence of any reports since the advent of widespread camera-trapping suggests that few, if any, remain. The Nilgiri marten (Martes gwatkinsii) is a rare species found widely but patchily within the Western Ghats. The Nilgiri striped palm squirrel (Funambulus sublineatus) is found widely but patchily within the Western Ghats (Kerala and Tamil Nadu).

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The Sahyadris forest rat (Rattus satarae) is confined to three small, widely separated forest areas of the Western Ghats (Maharashtra, Tamil Nadu and Karnataka). Bonhote’s mouse (Mus famulus) is known only from four localities within the Western Ghats of Kerala and Tamil Nadu. The Malabar spiny tree mouse (Platacanthomys lasiurus) is found widely but patchily within the Western Ghats (Karnataka, Kerala and Tamil Nadu). The Nilgiri long-tailed tree mouse (Vandeleuria nilagirica) is confined to two small areas of the Western Ghats (Karnataka and Tamil Nadu). Day’s pygmy white-toothed shrew (Suncus dayi) is known only from four localities within the Western Ghats of Kerala and Tamil Nadu. Salim Ali’s fruit bat (Latidens salimalii) is known only from a few localities within the Western Ghats. The Nilgiri laughingthrush (Montecincla cachinnans) was historically common within the Western Ghats but is now largely confined to the Nilgiri Hills, with a smaller disjunct population in the Palakkad-Siruvani Hills of Kerala. The Banasura laughingthrush (M. jerdoni) is confined to a few high-elevation shola localities in Karnataka and Kerala. Both are threatened by loss of habitat. The Nilgiri blue robin (Sholicola major) is confined to the Nilgiri Hills and South Wayanad Hills of Kerala, and to three peaks in south-western Karnataka. The white-bellied blue robin (S. albiventris) is found in scattered shola forest patches on the higher hills of the Western Ghats (Kerala and Tamil Nadu). Both are threatened by loss of habitat. The Nilgiri pipit (Anthus nilghiriensis) is a type of passerine bird found widely but patchily within the Western Ghats of Kerala, Tamil Nadu and Karnataka, where it is threatened by loss of its upland grassland habitat. The Travancore tortoise (Indotestudo travancorica) is a large species restricted to the forests of the Western Ghats (Kerala, Karnataka and Tamil Nadu). The Cochin Forest cane turtle (Vijayachelys silvatica) was first described from two specimens collected in 1912, and not seen again for 70 years. It was rediscovered in the Anamalai Hills in 1982, and has since been found elsewhere within the Western Ghats. The Indian kangaroo lizard (Otocryptis beddomii) is known from a few localities within the Western Ghats (Kerala and Tamil Nadu). Several species of day gecko (Cnemaspis) endemic to the Western Ghats are threatened by loss of habitat. The Wynad day gecko (C. wynadensis) is known only from two localities in Kerala. Jerdon’s day gecko (C. jerdonii) is known only from a few localities in Tamil Nadu. Indraneil Das’ day gecko (C. indraneildasii) is known from a few localities in Maharashtra, Karnataka, Tamil Nadu and Kerala. Beddome’s day gecko (C. beddomei) is known from a few localities in Tamil Nadu and Kerala. The coastal day gecko (C. littoralis) and the Nilgiri dwarf day gecko (C. indica) are both known from a few localities in Karnataka, Kerala and Tamil Nadu.

The Oriental Region

The Satara house gecko (Hemidactylus sataraensis) is confined to two localities within the Western Ghats (Maharashtra). Boulenger’s tree skink (Dasia subcaerulea) is known only from two specimens collected in the High Wavy Mountains and several others collected elsewhere in Tamil Nadu. There have been no records in recent years. Gans’ supple skink (Eutropis gansi) is known only from the Kalakkad-Mundunthurai Tiger Reserve in Tamil Nadu. Inger’s supple skink (E. clivicola) is known from two widely separated localities within the Western Ghats (Kerala). Rurk’s ristella (Ristella rurkii) is a type of skink known only from nineteenth-century historical records originating in the Anaimalai, Palni, and Travancore Hills (Tamil Nadu and Kerala). The Travancore ristella (R. travancorica) is confined to Pon Mudi Hill in Tamil Nadu and the Agasthyamalai Hills in Kerala. The Karnataka bronzeback tree snake (Dendrelaphis chairecacos) is known only from a few specimens collected from two widely separated localities within the Western Ghats (Karnataka and Kerala). The short-tailed kukri snake (Oligodon brevicauda) is known only a few specimens collected over a relatively wide area of Tamil Nadu and Kerala. The Travancore cat snake (Boiga dightoni) is known from a few disjunct areas of the Western Ghats in Kerala. The three-lined thorntail snake (Platyplectrurus trilineatus) is known only from three historical records originating in the Anamalai and Palani Hills (Kerala and Tamil Nadu). It was last reported in 1972. The Madura thorntail snake (P. madurensis) is known only from a few historical records originating from the Palani Hills in Tamil Nadu and the Cardamom Hills in Kerala. The Karnataka burrowing snake (Plectrurus canaricus) is known only from specimens collected during the late nineteenth century from Kudremukh Hill in Karnataka. The Kerala burrowing snake (P. aureus) is known only from a few specimens collected during the nineteenth century from the Western Ghats (Kerala). The spotted earth snake (Uropeltis maculata) is known only from two historical records from the southern Western Ghats (Anaimalai and Travancore Hills). The barred earth snake (U. myhendrae) is known from two localities in Tamil Nadu and Kerala. Günther’s earth snake (U. liura) is known only from the Madura and Agasthyamalai Hills of Tamil Nadu. Brougham’s earth snake (U. broughami) is known only from two specimens collected in 1878 from the Sirumalai Hills and Palani Hills of Tamil Nadu. The Travancore earth snake (Rhinophis travancoricus) is known from a few scattered localities in the Western Ghats (Kerala and Tamil Nadu). The two-lined black earth snake (Melanophidium bilineatum) is known from four localities within the Western Ghats (Kerala and Tamil Nadu). Thurston’s worm snake (Gerrhopilus thurstoni) is known only from three historical collections during the nineteenth

century within the Western Ghats (Kerala and Tamil Nadu). Beddome’s worm snake (G. beddomii) is known only from three nineteenth-century records originating from south of the Palghat Gap. The Belgaum worm snake (Indotyphlops exiguus) is known only from its original collection during the nineteenth century within the Western Ghats (Karnataka). The Malabar tree toad (Pedostibes tuberculosus) is known only from a few widely spaced localities within the Western Ghats, where it is threatened by loss of habitat. The Kempholey toad (Duttaphrynus brevirostris) is known only from its original collection in 1937 from Karnataka. The Silent Valley toad (D. silentvalleyensis) is known only from a single specimen collected in 1981 in or near Silent Valley National Park in Kerala. The small-eared toad (D. microtympanum) is known only from a few localities within the southern Western Ghats (Kerala and Tamil Nadu). Beddome’s toad (D. beddomii) is known from a few widely spaced localities in Karnataka, Kerala and Tamil Nadu. The Humbali toad (Xanthophryne koynayensis) is known only from two localities in the Western Ghats (Maharashtra). The Amboli toad (X. tigerina) is known only from Amboli, in Maharashtra. The red torrent toad (Ghatophryne rubigina) is known only from Silent Valley and Wynaad regions of Kerala. The ornate torrent toad (G. ornata) is known with certainty only from the Brahmagiri Hills and adjoining areas of Karnataka. The small gliding tree frog (Rhacophorus lateralis) is known only from two small areas in Karnataka and Kerala. The Kalakad gliding tree frog (R. calcadensis) is known only from the Agasthyamala Hills in Kerala and two national parks in Tamil Nadu. Both are threatened by loss of habitat. The ghat shrub frog (Ghatixalus asterops) is known only from the Palani and Anaimalai Hills (Kerala and Tamil Nadu). The Amboli shrub frog (Pseudophilautus amboli) is known only from a few localities in the vicinity of the Amboli Forest in southern Maharashtra. The dark-eared shrub frog (P. wynaadensis) is known only from a few widely spaced localities in Karnataka and Kerala. A great many species of bush frog (Raorchestes) are endemic to small areas of the Western Ghats, where they are highly threatened by habitat destruction. Sushil’s bush frog (R. sushili) is confined to Andiparai shola in Tamil Nadu. The Koadaikanal bush frog (R. dubois) is known only from a single locality in Tamil Nadu. The Kudremukh bush frog (R. tuberohumerus) is confined to two localities in Karnataka. Seshachar’s bush frog (R. charius) is known from three localities in Karnataka. The Kalpetta bush frog (R. nerostagona) is known from a few localities in Kerala. The Travancore bush frog (R. travancoricus) was long known only from a single specimen collected in the late nineteenth century, and thought to be extinct. Rediscovered in 2004, it is now known from two localities in Tamil Nadu. The Coorg yellow bush frog (R. luteolus) is known from three localities in Karnataka. The Ochlandra bush frog (R. ochlandrae) is known from a few

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localities in Kerala. The confusing green bush frog (R. chromasynchysi) is confined to a few localities in Karnataka and Kerala. The resplendent bush frog (R. resplendens) is confined to the summit of Mount Anamudi in Kerala, where it is protected within Eravikulam National Park. The Bombay bush frog (R. bombayensis) is found patchily in Maharashtra and Karnataka. The Munnar bush frog (R. munnarensis), greeneyed bush frog (R. chlorosomma) and Griet’s bush frog (R. griet) are all confined to the area of Munnar in Kerala. Jerdon’s bush frog (R. glandulosus) is found patchily throughout the Western Ghats. The yellow-bellied bush frog (R. flaviventris) was long known only from a single specimen collected from an undefined locality in 1882. It has since been rediscovered in the Anaimalai Hills of Tamil Nadu and Kerala, and may also occur in the Palani Hills. The Pon Mudi bush frog (R. ponmudi) was originally known only from Pon Mudi Hill in the southern Western Ghats, but has since been found in a few other localities in Kerala and Tamil Nadu. Several species of dancing frog (Micrixalus) are endemic to the Western Ghats, where they are threatened by habitat destruction. The elegant dancing frog (M. elegans) is known only from a single locality in Karnataka, while the Kottigehar dancing frog (M. kottigeharensis) is known only from two localities in Karnataka. The Malabar dancing frog (M. saxicola), Jerdon’s dancing frog (M. phyllophilus), Pillai’s dancing frog (M. nudis), Gadgil’s dancing frog (M. gadgili), and Boulenger’s dancing frog (M. silvaticus) are all found patchily throughout the Western Ghats. Günther’s leaping frog (Indirana brachytarsus), Leith’s leaping frog (I. leithii), Boulenger’s leaping frog (I. leptodactyla) and the spotted leaping frog (I. diplosticta) are all found sporadically throughout the Western Ghats, where they are threatened by loss of habitat. Several species of night frog (Nyctibatrachus) are endemic to the Western Ghats, where they are threatened by habitat destruction and pollution. The Kempholey night frog (N. kempholeyensis), meowing night frog (N. poocha), forest night frog (N. sylvaticus) and Coorg night frog (N. sanctipalustris) were each known only from a single specimen collected in the early twentieth century from Karnataka, and feared extinct. They were all rediscovered in 2011. The Dattatreya night frog (N. dattatreyaensis) is known only from the vicinity of Manikyadhara Falls within the Bhadra Wildlife Sanctuary, Karnataka. Humayun’s night frog (N. humayuni) is known from a few localities in Maharashtra. The Kudremukh night frog (N. karnatakaensis) is known only from Kudremukh National Park in Karnataka. The miniature night frog (N. minimus) is known only from the Wayanad Plateau in Kerala. The Shola night frog (N. sholai) is known only from Eravikulam National Park in Kerala. The Deccan night frog (N. deccanensis), Beddome’s night frog (N. beddomii), Alicia’s night frog (N. aliciae), Malabar night frog (N. major) and the small night frog (N. minor) are all found patchily throughout the Western Ghats. The Sholiga rice frog (Microhyla sholigari) is known from the southern Western Ghats (Karnataka and Kerala).

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The purple frog (Nasikabatrachus sahyadrensis) is known only from a few localities in the southern Western Ghats. The Sahyadris frog (Minervarya sahyadris) is known from two localities in the southern Western Ghats (Karnataka and Kerala). Four species of wart frog (Zakerana) endemic to the Western Ghats are threatened by loss of habitat. The Mysore wart frog (Z. mysorensis) is known only from its original collection during the 1920s in Karnataka. Murthy’s wart frog (Z. murthii) is known only from a few specimens collected from a single locality in Tamil Nadu. Rao’s wart frog (Z. sauriceps) is known only from its original collection during the 1930s from Karnataka. The short-webbed wart frog (Z. brevipalmata) is found widely but patchily within the Western Ghats. The Malabar narrow-mouthed frog (Melanobatrachus indicus) is known only from three localities in Tamil Nadu and Kerala. The Wattakole burrowing frog (Sphaerotheca leucorhynchus) is known only from a single (now lost) specimen collected from Karnataka in the 1930s. The triangular balloon frog (Uperodon triangularis) and the marbled balloon frog (U. mormorata) are both found patchily throughout the Western Ghats. The Kodagu striped caecilian (Ichthyophis kodaguensis) is known for certain only from the Venkidds Valley Estate in southern Karnataka. The Mudur caecilian (Gegeneophis madhavai) and the Mhadei caecilian (G. mhadeiensis) are each known only from a small area of Karnataka. The pink blind caecilian (G. carnosus) is known from a few localities in Karnataka and Kerala. The Humbarli caecilian (Indotyphlus maharashtraensis) is known only from a single locality in Maharashtra. Oommen’s caecilian (Uraeotyphlus oommeni), Menon’s caecilian (U. menoni) and the Kannan caecilian (U. narayani) are all known from the Western Ghats in Kerala. The Nilgiri Hills The Nilgiri Hills (or Nilgiris) are located in Karnataka, Kerala and Tamil Nadu. They span around 90 km in a north–south direction and nearly 80 km from east to west, and are clearly distinguished from the surrounding lowlands in the west, south, and east. The Nilgiri day gecko (Cnemaspis nilagirica) is known only from a single specimen collected in the Nilgiri Hills. Perrotet’s vine snake (Ahaetulla perroteti) is confined to grassland patches in shola forests of the Nilgiri Hills. Günther’s burrowing snake (Plectrurus guentheri) is known only from two specimens collected during the late nineteenth-century in the Nilgiri Hills. Tindall’s worm snake (Gerrhopilus tindalli) is known only from two specimens collected in the 1940s from Kerala. The variable shrub frog (Ghatixalus variabilis) is confined to a few localities within the Nilgiri Hills.

The Oriental Region

The spotted bush frog (Raorchestes tinniens) and the cross-marked bush frog (R. signatus) are both confined to the Nilgiri Hills. Thampi’s dancing frog (Micrixalus thampii) is known only from Silent Valley National Park in Kerala. The Nilgiri wart frog (Zakerana nilagirica) is known only from two localities in Kerala and Tamil Nadu. The long-headed caecilian (Ichthyophis longicephalus) is known only from Silent Valley National Park in Kerala. The Malabar caecilian (Uraeotyphlus malabaricus) is known with certainty only from a single locality in Tamil Nadu. The Anaimalai Hills Spanning the border between Kerala and Tamil Nadu, the Anaimalai Hills are a range of mountains that form the southern part of the Western Ghats. They are separated from the latter to the north by the Palakkad Gap. Günther’s mountain snake (Xylophis stenorhynchus) is known for certain only from the Valparai Plateau, Anaimalai Hills (Kerala and Tamil Nadu). Beddome’s earth snake (Uropeltis beddomii) is known only from a single specimen collected in 1862 in the Anaimalai Hills (Tamil Nadu). The Ponachi earth snake (U. macrorhyncha) is known only from a single specimen collected in 1877 in the Anaimalai Hills (Tamil Nadu). The Cochin earth snake (U. nitida) is known only from a few specimens collected in 1878 in the Anaimalai Hills (Kerala). Peters’ earth snake (U. petersi) is known only from its original collection in 1878 from an imprecise locality in Tamil Nadu. The Anamalai gliding tree frog (Rhacophorus pseudomalabaricus) is known only from an area centered on Indira Gandhi National Park in Tamil Nadu. The Anamalai leaping frog (Indirana phrynoderma) is known only from a single locality in the Anaimalai Hills (Tamil Nadu). The Anaikatti Hills are a subrange of the Anaimalai Hills located near the city of Coimbatore in Tamil Nadu. The Anaikatti Hills day gecko (Cnemaspis anaikattiensis) is known only from the Anaikatti Hills. The Palani Hills The Palani Hills (also known as the Palni Hills) are an eastward extension of the Western Ghats located in Kerala and Tamil Nadu. They adjoin the high Anaimalai Hills to the west. The Palani ground skink (Kaestlea palnica) is known only from its original collection at two localities within the Palani Hills in the late nineteenth century. Nikhil’s kukri snake (Oligodon nikhili) is known only from the Palani Hills in Kerala. The Sungam Hills The Sungam Hills are located between the Anaimalai and Nelliyampathi Hills in Kerala. The Sungam dot frog (Ramanella anamalaiensis) was long known only from a (now lost) type specimen collected in the

Anaimalai Hills in 1937. A population was rediscovered in the Parambikulam Wildlife Sanctuary in 2010. The Parambikulam frog (Fejervarya parambikulamana) is known only from a now-lost holotype collected from what is now the Parambikulam Wildlife Sanctuary. The Nelliyampathi Hills The Nelliyampathi Hills are located in Kerala. Kaikatt’s bush frog (Raorchestes kaikatti) and Mark Wilkinson’s bush frog (R. marki) are both confined to the Nelliyampathi Hills. The Cardamom Hills The Cardamom Hills are located in south-eastern Kerala and south-western Tamil Nadu. They conjoin the Anaimalai Hills to the north-west, the Palani Hills to the north-east and the Agasthyamalai Hills to the south. The Cardamom Hills earth snake (Rhinophis fergusonianus) is only known from a single specimen collected near Trivandrum in Kerala. The white-spotted bush frog (Raorchestes chalazodes) is known only from the south-western slopes of the Cardamom Hills, where it was rediscovered in 2011 after not having been reported for 125 years. The Agastyamalai Hills The Agastyamalai Hills (formerly known as the Ashambu Hills) are located in the southern Western Ghats (Kerala and Tamil Nadu). The Ashambu laughingthrush (Montecincla meridionalis) is confined to the Agastyamalai Hills (Kerala and Tamil Nadu), where it is threatened by habitat destruction. The orange-lipped forest lizard (Microauris aurantolabium) is known only from the Kalakkad-Mundunthurai Tiger Reserve and Peppara Wildlife Sanctuary (Tamil Nadu and Kerala). Vasanthi’s night frog (Nyctibatrachus vasanthi) is known only from the Kalakkad-Mundanthurai Tiger Reserve in Tamil Nadu. Gans’ caecilian (Uraeotyphlus gansi) is known only from the Kalakkad-Mundanthurai Tiger Reserve in Tamil Nadu. Pon Mudi Hill is located in the south-western Agastyamalai Hills (Kerala). The Pon Mudi grassland bush frog (Raorchestes graminirupes) and Bob Inger’s bush frog (R. bobingeri) are both confined to Pon Mudi Hill.

The Eastern Ghats The Eastern Ghats are a discontinuous range of mountains running along India’s eastern coast (Odisha, Andhra Pradesh, Madhya Pradesh, Tamil Nadu, Karnataka and Telangana). They are dry, heavily eroded and cut through by four major rivers. The large rock rat (Cremnomys elvira) is known only from a single locality within the Eastern Ghats (Tamil Nadu). The

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species went unrecorded from the 1940s until its rediscovery in the Shervaroy Hills in 2013. Jerdon’s courser (Rhinoptilus bitorquatus) is a long-legged, nocturnal ground bird confined to a few areas of the Eastern Ghats of Andhra Pradesh and extreme southern Madhya Pradesh. Long known only from a few historical records and thought to have gone extinct around 1900, when the last individual was collected, it was unexpectedly rediscovered in 1986. It is currently known for certain only from the Sri Lankamalleswara Wildlife Sanctuary, where the total population is believed to number between 50 and 250. The Visakhapatnam bush frog (Raorchestes terebrans) is known only from a single locality in Andhra Pradesh. The Similipal shrub frog (Philautus similipalensis) is known only from the region of Similipal National Park in Odisha. The Jeypore Hills The Jeypore Hills are located in central-eastern India (Andhra Pradesh and Odisha). The Jeypore bent-toed gecko (Cyrtodactylus jeyporensis) was long known only from a single specimen collected in the Jeypore Hills in 1877. It was rediscovered in 2010. The Chitteri Hills The Chitteri Hills (also known as the Sitheri Hills) are located in south-eastern India (northern Tamil Nadu). Pruth’s supple skink (Lygosoma pruthi) is known only from a single specimen collected from the Chitteri Hills in 1929. The Sirumalai Hills The Sirumalai Hills are located in southern India (southern Tamil Nadu). The Sirumalai Hills earth snake (Uropeltis dindigalensis) is known only from the Sirumalai hills, where it was last recorded in the 1970s.

The Shillong Plateau The Shillong Plateau is located in north-eastern India (Assam and Meghalaya). Its southern, northern and western ridges form the Garo, Khasi, and Jaintia hills, respectively. The tawny-breasted wren-babbler (Spelaeornis longicaudatus) now appears to be confined to the Khasi Hills of Meghalaya due to loss of habitat. The Garo Hills The Garo Hills are located in north-eastern India (Assam and Meghalaya). Kemp’s toad (Bufoides kempi) is known only from two specimens collected before 1919 from the Garo Hills. The Tura shrub frog (Philautus kempiae) is known only from a single specimen collected in the early twentieth century. The Siju Cave stone loach (Schistura sijuensis) is known only from Siju Cave in the Garo Hills.

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The Khasi Hills The Khasi Hills are located in north-eastern India (Meghalaya). The Khasi rock toad (Bufoides meghalayanus) is known only from three montane forest localities in the Khasi Hills, where it is threatened by loss of habitat. The Shillong bush frog (Raorchestes shillongensis) is confined to a small area of forest within the Khasi Hills. The Khasi wart frog (Limnonectes khasianus) is known only from a now-lost type specimen collected in the Khasi Hills. The Jaintia Hills The Jaintia Hills are located to the east of the Khasi Hills (Meghalaya). The Krem Synrang stone loach (Schistura papulifera) is confined to the Krem Synrang cave system in the Jaintia Hills.

The Shan Hills The Shan Hills are a vast mountainous region in centraleastern Myanmar (Shan State) and north-western Thailand. They are comprised of numerous mountain ranges separated mainly by steep, narrow river valleys and a few plateaus. The Celestial pearl danio (Danio margaritatus) is a type of fish known from a few heavily vegetated pools on the Shan Plateau of Myanmar and Thailand. It has been heavily exploited for the international aquarium trade. The Hwe-gna-sang stone loach (Schistura acuticephala) is known only from the Hwe-gna-sang River, on the Shan Plateau of northern Myanmar (Shan State). The Daen Lao Range The Daen Lao Range is located in the southern and southeastern area of the Shan Hills. It separates the Salween watershed from that of the Mekong. Doi Suthep Mountain is located west of Chiang Mai in north-western Thailand. The Doi Suthep caecilian (Ichthyophis youngorum) is known only from a handful of adult and larval specimens collected in 1957 from a small forested valley. The Thanon Thong Chai Range The southernmost part of the Shan Hills, the Thanon Thong Chai Range is located in north-western Thailand and centraleastern Myanmar. The green odorous frog (Odorrana livida) is known only from the Dawna Hills in central-eastern Myanmar.

The Arakan Mountains The Arakan Mountains (Rakhine Yoma in Burmese) are located in western Myanmar (Rakhine and Chin states) and north-eastern India (Manipur). The Rakhine Yoma bent-toed gecko (Cyrtodactylus wakeorum) is known only from the Rakhine Yoma Elephant Range in southern Rakhine state. The Irrawaddy bent-toed

The Oriental Region

gecko (C. ayeyarwadyensis) is known only from two localities within the Arakan Mountains. The spotted foam-nest tree frog (Chiromantis punctatus) is known only from the south-western foothills of the Arakan Mountains. The Arakan large-eyed litter frog (Leptobrachium rakhinensis) is known only from two localities within the Arakan Mountains. The Chin Hills The Chin Hills are located in western Myanmar (Chin State), extending northward into north-eastern India (Manipur). Mount Victoria (Khonu Msung in Burmese) is located in the southern Chin Hills. It the highest peak in the Arakan Mountains. The white-browed nuthatch (Sitta victoriae) is known only from the Mount Victoria area. It is threatened by loss of its montane forest habitat.

The Pegu Range The Pegu Range (Pegu Yoma in Burmese) is located in southcentral Myanmar. It is essentially an area of low mountains, uplands and hills between the Irrawaddy and Sittaung rivers. The Popa langur (Trachypithecus popa) is confined to the Pegu Range, where the total population is thought to be less than 260. The Toungoo frog (Humerana oatesii) is only known from specimens collected in the Pegu Mountains in the 1890s. The Pegu longtail catfish (Olyra burmanica) is known only from a few old museum specimens collected from the Pegu Range. The Pegu stone loach (Neonoemacheilus peguensis) is known only from the Pegu Range. Mount Popa is an extinct volcano located in central Myanmar. The Popa bent-toed gecko (Cyrtodactylus brevidactylus) is confined to Mount Popa.

The Cardamom Mountains The Cardamom Mountains (Chuor Phnom Kravanh in Khmer/Thio Khao Banthat in Thai) are located largely in south-western Cambodia, extending marginally into eastern Thailand. Dense tropical rainforest prevails on the wet westward slopes, while the eastern slopes are drier. The Cardamom wolf snake (Lycodon cardamomensis) is known only from a few specimens collected in the Cardamom Mountains. The Cardamom horned frog (Megophrys auralensis) is known only from the Phnom Aural Wildlife Sanctuary in Kampong Speu province, and the Central Cardamoms Protected Forest in Pursat province. The Cardamom spiny frog (Quasipaa fasciculispina) is confined to the Cardamom Mountains, where it is threatened by habitat destruction and overcollection for human consumption.

Mount Samkos Mount Samkos (Phnom Samkos in Khmer) is located in western Cambodia (Pursat province). It is the country’s second highest peak. The Samkos foam-nest tree frog (Chiromantis samkosensis) is currently known only from Samkos Mountain, but may also be found in neighbouring areas. The Cardamom shrub frog (Philautus cardamonus) is known only from the Phnom Sankos Wildlife Sanctuary, but likely ranges more widely. The Damrei Mountains The Damrei Mountains (Chuor Phnum Damrei in Khmer – literally ‘Elephant Mountains’) are a southern offshoot of the Cardamom Mountains located in south-western Cambodia (Koh Kong and Kampot provinces). The Damrei horned frog (Megophrys damrei) is known only from the Bokor Plateau within the Damrei Mountains. Long known only from a single specimen collected in 1914, it was rediscovered in 2011.

The Annamite Mountains The Annamite Mountains (Day Truong Son in Vietnamese/ Xai Phou Luang in Laotian/Chaîne annamitique in French) extend approximately 1100 km through central and southern Vietnam, south-eastern Laos, and a small area of northeastern Cambodia. Its montane rainforests are among the last relatively intact examples in Indochina. Known for their globally outstanding biodiversity, in recent years a number of spectacular large mammal discoveries have been made here. The saola or spindle-horned antelope (Pseudoryx nghetinhensis) is one of the world’s rarest large mammals, being known only from the Annamite Mountains of Laos and Vietnam. The species was only just discovered in 1992 after the discovery of remains in the Vu Quang Nature Reserve, and is still rarely seen in the wild. It has been kept in captivity on a few occasions, although only for short periods. Hunting and trapping are thought to be a threat to the small population. The Annamite muntjac (Muntiacus truongsonensis) is a tiny species first discovered in the Annamite Mountains of central Vietnam (Quang Nam province) in 1997. Taxonomic confusion with related species makes it impossible to be more precise about its distributional limits or status. The Annamite striped rabbit (Nesolagus timminsi) occurs at low and medium altitudes in the northern and central Annamite Mountains along the Vietnam–Laotian border. It is threatened by hunting, trapping and loss of habitat. The Annamite mouse-eared bat (Myotis annamiticus) is known for certain only from a single valley in central Vietnam (Quang Binh province), although its range likely extends further and perhaps into neighbouring Laos. The golden-winged laughingthrush (Trochalopteron ngoclinhense) is known only from Mount Ngoc Linh and Mount Ngoc Boc in south-central Vietnam (Kon Tum and Quang

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Nam provinces), where it is threatened by habitat destruction and trapping. The grey-crowned crocias (Laniellus langbianis) is a rare passerine bird was long known only from five specimens collected on the Lang Bian Plateau of south-central Vietnam (Lam Dong province) in the late 1930s, it was rediscovered there in 1994 and has since also been recorded from Kon Tum province. It is highly threatened by habitat destruction. Smith’s blind skink (Dibamus smithi) is known only from two localities in southern Vietnam (Lam Dong and Khanh Hoa provinces). The Annam keelback (Parahelicops annamensis) is a type of snake known only from a few specimens collected in central Vietnam (Da Nang and Kon Tum provinces) and southeastern Laos (Xe Kong province). Thanh’s reed snake (Calamaria thanhi) is known only from a single specimen collected near Phong Na-Ke Bang National Park in central Vietnam (Quang Binh province). The concolor reed snake (C. concolor) is known only from a single specimen collected in Bach Ma National Park in central Vietnam (Thua Thien-Hue province). Bourret’s cat snake (Boiga bourreti) is known only from a few sporadic localities in central Vietnam (Quang Binh and Kom Tum provinces). The Truong Son pit viper (Viridovipera truongsonensis) is a rare species known with certainty only from the vicinity of Phong Nha-Ke Bang National Park, in the Annamite Mountains of central Vietnam (Quang Binh province). It may be found more widely. The Ananjeva tree frog (Kurixalus ananjevae) is confined to central Vietnam (Ha Tinh province). Quang’s foam-nest tree frog (Gracixalus quangi) is known only from three isolated areas of montane forest in northern Vietnam (Nghe An, Thanh Hoa, and Son La provinces). The Khammouane wart frog (Limnonectes khammonensis) is known only from the Khammouane Plateau in central Laos, near the border with Vietnam. The hidden shrub frog (Philautus abditus) is known only from a small area of south-central Vietnam (Gia Lai and Kon Tum provinces) and extreme north-eastern Cambodia (Ratanakiri province). The Dak Cheung large-eyed litter frog (Leptobrachium xanthops) is known only from the Dak Cheung Plateau in south-eastern Laos (Sekong province). Firth’s dwarf litter frog (Leptobrachella firthi) is known only from two localities in south-central Vietnam (Kon Tum and Quang Nam), although it likely occurs at least a little more widely. The Annamite sticky frog (Kalophrynus cryptophonus) is known only from a small area of the southern Annamite Mountains in southern Vietnam (Lam Dong province). The cricket bush frog (Raorchestes gryllus) is known from a few montane localities in south-central Vietnam. The Laotian knobby newt (Tylototriton notialis) is known only from the Nakai-Nam Theun National Protected Area in

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central Laos (Khammouane province) and the Pu Hoat Proposed Nature Reserve in northern Vietnam (Nghe An province). The Central Highlands of Vietnam The Central Highlands (Cao nguyen Trung phan in Vietnamese) are a series of contiguous plateaus in southcentral Vietnam (Dak Lak, Dak Nong, Gia Lai, Kon Tum and Lam Dong provinces). They are surrounded by higher mountains of the southern Annamite Range. The wrinkly skink (Vietnascincus rugosus) is known only from a single locality on the border of Gia Lai and Kon Tum provinces. The Gia Lai reed snake (Calamaria gialaiensis) is known only from a single specimen collected from Kon Ka Kinh National Park in Gia Lai province. Sang’s reed snake (C. sangi) is known only from two specimens collected from Kon Tum province. The marble-backed gliding tree frog (Rhacophorus marmoridorsum) is known only from a single locality in Gia Lai province. The vampire gliding tree frog (R. vampyrus) is known only from two localities in Lam Dong and Dak Nong provinces. Both are threatened by loss of habitat. The Buon Luoi horned frog (Megophrys hansi) is known only from a single locality in south-central Vietnam (Gia Lai province). Appleby’s slender litter frog (Leptolalax applebyi) is known only from two localities in Quang Nam and Kon Tum provinces. The pale-eyed large-eyed litter frog (Leptobrachium leucops) is known only from a few localities in Lang Dong, Khanh Hoa, Ninh Thuan, and Dak Nong provinces. The Tay Nguyen Plateau is located in Gia Lai province. The Tay Nguyen large-eyed litter frog (Leptobrachium xanthospilum) is known only from the Tay Nguyen Plateau. The dusty-speckled rice frog (Microhyla pulverata) is known only from the Tay Nguyen Plateau. The Kon Tum Plateau is located in Kon Tum province. Cuc’s mountain stream snake (Opisthotropis cucae) is known only from the Kon Tum Plateau. Ryabov’s bug-eyed tree frog (Theloderma ryabovi) is known only from a single locality on the Kon Tum Plateau. The Mang Canh odorous frog (Odorrana gigatympana) is known only from a small area of the Kon Tum Plateau. The Lang Bian Plateau (also known as the Da Lat Plateau) is located in south-central Vietnam (Lam Dong province). It has been designated an Endemic Bird Area by Birdlife International. The collared laughingthrush (Trochalopteron yersini) is an uncommon species known only from a handful of localities on the Lang Bian Plateau including Mount Lang Bian, Mount Bi Doup and Chu Yang Sin National Park. The southern Vietnam box turtle (Cuora picturata) is confined to the eastern slopes of the Lang Bian Plateau, where it is seriously threatened by overcollection for use as food.

The Oriental Region

The Lang Bain bug-eyed tree frog (Theloderma palliatum) is known only from Bidoup Nui Ba National Park, but may range more widely. The Lang Bain golden-backed frog (Hylarana montivaga) is known for certain only from a few areas of the Lang Bian Plateau, where it is threatened by agricultural expansion and overcollection for food. The Bidoup dwarf litter frog (Leptobrachella bidoupensis) is known only from Bidoup Nui Ba National Park. The Lang Bian rice frog (Microhyla fusca) is confined to the Lang Bian Plateau. Mount Ba Na Mount Ba Na is located in coastal central Vietnam (Thua Thien-Hue province), where it is isolated from the rest of the Annamite Mountain chain by lowlands. The Ba Na tree frog (Kurixalus banaensis) is confined to the region of Mount Ba Na, where it was reported to be very common in the 1930s but has not been reported since. Mount Ngoc Linh Mount Ngoc Linh is an isolated mountain in south-central Vietnam (Quang Nam and Kon Tum provinces). Much of its fauna is believed to be endemic. Sokolov’s shrew (Crocidura sokolovi) is known only from Mount Ngoc Linh. Abramov’s reed snake (Calamaria abramovi) is known only from two specimens collected from Mount Ngoc Linh. The misty bug-eyed tree frog (Theloderma nebulosum) is known only from a single adult specimen and some larvae collected on Mount Ngoc Linh. The thorny foam-nest tree frog (Gracixalus lumarius) is known only from Mount Ngoc Linh. The Ngoc Linh large-eyed litter frog (Leptobrachium ngoclinhense) is known only from Mount Ngoc Linh. The thumbless rice frog (Microhyla nanapollexa) is known only from a single specimen collected on Mount Ngoc Linh.

Miscellaneous Mountains and Highlands The Satpura Range is located in north-central India (Madhya Pradesh, Chhattisgharh, and Odisha). The sacred grove bush frog (Raorchestes sanctisilvaticus) is known only from two localities within the Maikal Hills in north-central India (Madhya Pradesh and Odisha). The Singharh Plateau is located in western coastal India (Maharashtra). The Kondana rat (Millardia kondana) is confined to a few localities on the Singharh Plateau. The Chittagong Hills are located in south-eastern Bangladesh. Cox’s Bazar danio (Devario anomalus) is known only from a single small stream within the southern Chittagong Hills. The Kakhien Hills are located in north-eastern Myanmar. The Kakhien Hills tiny frog (Nanorana feae) is known only from the Kakhien Hills. The Bolaven Plateau is located in southern Laos (Champasak, Salavan, Sekong, and Attapeu provinces). The

Bolaven odorous frog (Odorrana bolavensis) and Buchard’s large-eyed litter frog (Leptobrachium buchardi) are each known only from a small area of the Bolaven Plateau.

Lowland Tropical Moist Forests and Swamp Forests Tropical rainforest once covered much of South and South East Asia, although today only scattered fragments remain, chiefly in mountainous and limestone karst areas. Within the Oriental Region pockets of lowland rainforest are mainly to be found in Laos, Thailand and Cambodia. The hinpoon limestone rat (Niviventer hinpoon) is confined to limestone karst forests in central Thailand (Saraburi, Lopburi, and Nakhon Sawan provinces). Paulina’s limestone rat (Saxatilomys paulinae) is known only from specimens collected within the Khammouane Limestone NBCA in central Laos and from north-central Vietnam (Quang Binh province). Edwards’ pheasant (Lophura edwardsi) is (or was) confined to the few remaining areas of lowland rainforest in central Vietnam, where it may already be extinct in the wild. Fortunately, there is a large captive population. A form previously known as the Hat Tinh pheasant (L. hatinhensis) is now known to be a colour variant, while the so-called ‘imperial pheasant’ is thought to be a naturally occurring hybrid between this species and the Annam silver pheasant (L. nycthemera annamensis), itself a threatened form confined to southern Vietnam. Another subspecies, Engelbach’s silver pheasant (L. n. engelbachi), is confined to a small area of southern Laos. The rufous limestone babbler (Turdinus calcicola) is a type of passerine bird found only in the limestone hills of central Thailand (Saraburi province). The Arakan forest turtle (Heosemys depressa) is confined to a small area of western Myanmar and south-eastern Bangladesh, where it is threatened by hunting for food and overcollection for the international pet trade. The Goan day gecko (Cnemaspis goaensis) is known only from three lowland forest localities in western coastal India (Goa and Karnataka). The ruby-eyed green pit viper (Trimeresurus rubeus) is known only from a few localities in southern Vietnam and eastern Cambodia (Mondolkiri province). The Kanburi pit viper (Cryptelytrops kanburiensis) is known only from western Thailand (Kanchanaburi province). The three-horned scaled pit viper (Protobothrops sieversorum) is known only from two localities within the limestone karst forests of eastern Laos and north-central Vietnam. The Red River krait (Bungarus slowinskii) is a type of venomous snake known only from two localities in northern and central Vietnam. Andrea’s keelback (Hebius andreae) is a type of nonvenomous snake known only from a single locality adjacent to Phong Nha-Ke Bang National Park in north-central Vietnam (Quang Binh province).

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Smith’s rough water snake (Fimbrios smithi) is known only from a single specimen collected from lowland karst forest in the Annamite Mountains of central Vietnam. Angel’s kukri snake (Oligodon macrurus) is confined to a small area of southern coastal Vietnam (Khanh Hoa and Ninh Thuan provinces). Morice’s kukri snake (O. moricei) is known only from a single locality in southern coastal Vietnam (Kranh Hoa province). Hampton’s kukri snake (O. hamptoni) is known only from two specimens collected in northern Myanmar during the early twentieth century. The red-striped kukri snake (O. erythrorhachis) is known only from two localities in north-eastern India (Arunachal Pradesh). The Annam kukri snake (O. annamensis) is known only from two specimens collected during the mid-twentieth century from southern Vietnam (Lam Dang province). Kundu’s wolf snake (Lycodon kundui) is known only from a single specimen collected in the 1940s from south-central Myanmar. Repeated surveys of the locality have failed to find it since. Rendahl’s wolf snake (L. paucifasciatus) is known only from a small area of central Vietnam (Thua Thien-Hue and Quang Binh provinces), where it is threatened by loss of habitat. Nguyenvansang’s snake (Colubroelaps nguyenvansangi) is known only from two specimens collected in southern Vietnam (Binh Phuoc and Lam Dong provinces). Flower’s worm snake (Gerrhopilus floweri) is known only from fruit plantations near the city of Bangkok, central Thailand. The Rakhine Yoma toad (Duttaphrynus crocus) is known only from two, closely situated localities in southern Myanmar. Stuart’s toad (D. stuarti) is known only from a few localities within a small area on northern Myanmar (Kachin state) and north-eastern India (Arunachal Pradesh and possibly Assam). Helen’s gliding tree frog (Rhacophorus helenae) is known only from two widely separated forest fragments in southern Vietnam (Binh Thuan and Dong Nai provinces). The bamboo bug-eyed tree frog (Theloderma bambusicolum) is known only from a small area of southern Vietnam (Binh Phuoc, Dak Nong, Lam Dong, and Dong Nai provinces). The slender bug-eyed tree frog (T. petilum) is known only from a small area of north-western Vietnam (Dien Bien province) and northern Laos (Phongsaly province). The Seima horned frog (Megophrys synoria) is known only from the Seima Biodiversity Conservation Area in Cambodia (Mondolkiri province). The Gundia leaping frog (Indirana gundia) and the Kempholey leaping frog (I. longicrus) are both known only from a single locality each in south-western India (Karnataka). Annandale’s oriental frog (Ingerana borealis) remains relatively widespread in the lowland forests of north-eastern India, Bhutan and possibly Bangladesh, but is everywhere under threat by water pollution and loss of habitat. The Chilapata frog (Minervarya chilapata) is known only from the Chilapata Forest Reserve in north-eastern India (West Bengal).

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The Goa caecilian (Gegeneophis goaensis) is known only from three specimens collected in south-western India (Goa). The Krishna Farms caecilian (G. krishni) is known only from a single locality in south-western coastal India (Karnataka). Nadkarni’s caecilian (G. nadkarnii) is known only from the Bondla Wildlife Sanctuary in south-western India (Goa).

Lowland Tropical Dry Forests and Shrublands Areas of lowland tropical dry forest have largely disappeared from South and South-East Asia, although there are still scattered patches remaining in parts of India and Indo-China. The Vellore day gecko (Cnemaspis otai) is known only from scrub jungle areas in the town of Vellore, south-eastern India (Tamil Nadu). The Kolhapur Hills day gecko (C. kolhapurensis) is known only from the Kolhapur Hills in centralwestern India (Maharashtra). The Vagheshwari Mata house gecko (Hemidactylus gujaratensis) is known only from a single temple in western coastal India (Gujarat). Malcolm Smith’s house gecko (H. malcolmsmithi) is known only from a small area of northern India (Punjab). The Poona skink (Eurylepis poonaensis) is known only from a few specimens collected in central-western India (Maharashtra). Blanford’s supple skink (Eutropis innotata) is known only from a few, mostly historical records in east-central India (Maharashtra, Madhya Pradesh and Andhra Pradesh). Ashwamedh’s supple skink (E. ashwamedhi) is confined to a small area of eastern coastal India (Andhra Pradesh). The pygmy writhing skink (Subdoluseps frontoparietale) is known only from two specimens collected some 40 years appear within a small area of central Thailand (Lopburi province). Roule’s snake skink (Isopachys roulei) is known only from two disjunct localities in south-central Thailand (Chonburi and Nakhon Ratchasima provinces). The Myanmar short-tailed python (Python kyaiktiyo) is confined to secondary scrub in southern Myanmar (Mon State). Ozaki’s blind snake (Indotyphlops ozakiae) is known only from a few specimens collected in the Sakaerat Biosphere Reserve in central Thailand (Nakhon Ratchasima province). Seshachari’s caecilian (Gegeneophis seshachari) and the Amboli caecilian (G. danieli) are each known only from a single locality in western coastal India (Maharashtra).

Tropical Savannas and Grasslands Grasslands and savannas within the Oriental Region once covered a large part of the Indo-Gangetic Plain, extending across India from the Arabian Sea to the Bay of Bengal, with smaller regions found patchily in Indochina. Sadly, much of these areas have been changed into deserts through overgrazing by domestic cattle. The Bengal florican (Houbaropsis bengalensis) is a type of bustard found in north-eastern India from Uttar Pradesh to

The Oriental Region

Assam, through Nepal and, at least historically, into Bangladesh. There is another disjunct population in central Cambodia and possibly adjacent southern Vietnam. The species has disappeared from most of this range due to loss of habitat and only survives now in scattered pockets. In 2017 the total population was estimated at less than 1000. The swamp grass-babbler (Laticilla cinerascens) is a notably rare species known from north-eastern India (Assam and Bihar) and parts of northern Bangladesh, although there have been records from the latter country since the early 1970s. In India it is scattered into small populations that are under threat from loss of habitat. The marsh babbler (Pellorneum palustre) is found in north-eastern India (Assam, Arunachal Pradesh, and Meghalaya) and eastern Bangladesh, where it is threatened by loss of habitat. The bristled grassbird (Chaetornis striata) inhabits tall, grass-covered marshlands in the Indian subcontinent, where it is threatened by habitat destruction.

Seasonally Flooded and Riverine Grasslands Remnant areas of seasonally flooded forest are to be found in India and Indo-China. Schomburgk’s deer (Rucervus schomburgki) was a graceful species similar in appearance to the barasingha (R. duvaucelii) previously discussed. It lived in the swampy plains of central Thailand, in particular the Chao Phraya River valley near Bangkok, where in the late nineteenth century it was apparently ‘not uncommon’. During the rainy season flooding would force herds together onto higher ground that often turned into islands, where they became easy targets for hunters. However, it was the commercial production of rice for export which began in the late nineteenth century that appears to have doomed the animals, leading as it did to the loss of nearly all the grassland and swamp areas on which they depended. The species appears to have undergone a very rapid decline after 1910. The last-known wild individual was shot in 1932, and the last captive died in 1938. Only one mounted specimen remains, although there are still hundreds of skulls and antlers that had been taken as trophies, and which still show up occasionally in the Chinese ‘traditional medicine’ trade. The swamp barasingha (Rucervus duvaucelii duvaucelii) is a large deer adapted to the tall, flooded grasslands of the IndoGangetic plain north of the Ganges. It has declined considerably owing to the cultivation of the marshy grasslands. Only a few thousand still survive in India (West Bengal and Uttar Pradesh) and in the Sukla Phanta Wildlife Reserve in southwestern Nepal. It has been extirpated from Pakistan. Another subspecies, the eastern barasingha (R. d. ranjitsinhi) historically occurred across the Brahmaputra plain in India and Bangladesh, but is now confined to a small area of Assam. The vast majority, about 1150, live in Kaziranga National Park. The Manipur brow-antlered deer (R. eldii eldii) has long been confined to the floating marshy grasslands on the shores of

Figure 5.3 Schomburgk’s deer in the Berlin Zoo, 1911. (Credit: Lothar Schlawe.)

Loktak Lake in eastern India (Manipur), where it is protected within Keibul Lamjao National Park. In the 1950s it had been reduced to just a few herds, although today the population is more or less stable at around 250. The black-breasted parrotbill (Paradoxornis flavirostris) was historically found in Bangladesh, eastern Nepal and northern India but now appears to be confined to a few localities within the Brahmaputra Valley in the latter country (Arunachal Pradesh and Assam). The Myanmar Jerdon’s babbler (Chrysomma altirostre altirostre) is a type of passerine bird that was thought to have become extinct in 1941, but was rediscovered in 2015 in southcentral Myanmar. Terai-Duar Savannas and Grasslands The Terai-Duar Savannas and grasslands are a narrow lowland region, about 25 km wide, located at the base of the Himalayas in Bhutan, Bangladesh, Nepal and India (West Bengal and Assam). It consists of a mosaic of tall riverside grasslands, savannas, and forests. The pygmy hog (Porcula salvania) is the smallest and rarest of the world’s suids. Nocturnal and the size of a hare, it is difficult to detect and known only from a few localities in northern West Bengal and north-western Assam, although most likely it ranged at one time throughout the tall, swampy grasslands of the Terai-Duar in India, Bangladesh and Nepal. Loss of habitat due to cultivation and grazing resulted in a marked decline during the mid-twentieth century. Today it is now largely confined to small, isolated pockets in and around Manas National Park in north-western Assam, where the total wild population is currently thought to number less than 150. A further 16 captive-bred animals were released in the Sonai

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Rupai Wildlife Sanctuary in 2008, and similar reintroductions have been planned for other protected areas in Assam. The hispid hare (Caprolagus hispidus) formerly occurred in areas of elephant grass in the Himalayan foothills from Uttar Pradesh through southern Nepal, northern West Bengal, Assam, and into Bangladesh. Like the pygmy hog it has suffered considerably from habitat destruction and degradation, and indeed by the 1960s was thought by many to be extinct. In 1971, however, a live specimen was captured in the Barnadi Wildlife Sanctuary in north-western Assam. It is currently thought to persist throughout much of its former range, although sporadically. The swamp francolin (Francolinus gularis) is a type of partridge historically found in the wet grasslands of the Ganges and Brahmaputra river drainages of north-eastern India, Nepal, and Bangladesh. It has been extirpated from much of its range due mainly to loss of habitat. The slender-billed babbler (Chatarrhaea longirostris) was fairly widespread and common during the nineteenth century but is today confined to three disjunct areas (one in central Nepal and two others in north-eastern India), where it mainly occurs in national parks. The Terai Jerdon’s babbler (Chrysomma altirostre griseigularis) is a type of passerine bird confined to the Terai of northeastern India and western Nepal. Finn’s weaver (Ploceus megarhynchus) is a type of passerine bird endemic to the Terai of the northern Indian subcontinent, where it is known from three disjunct populations.

Dry Grasslands The dry grasslands of the Oriental Region are of two main types: steppe-like plains partly covered by shrubs, and somewhat richer grasslands, bushy country or grassy plains with scattered thin woods. The great Indian bustard (Ardeotis nigriceps) is a large, ostrich-like species that formerly ranged widely over the arid plains of the Indian subcontinent. Heavy hunting and habitat loss due to cultivation and overgrazing have reduced it to a few pockets of suitable habitat, mainly in central and western India. In 1969 the total population was estimated at 1260, but had declined to around 250 by 2011 and to as few as 150 in 2018. The lesser florican (Sypheotides indicus), the smallest of the bustards, was historically widespread and common across the Indian subcontinent but has been steadily declining since the 1870s due to hunting and habitat destruction. Today it is largely confined to areas of central and western India.

Deserts and Xeric Shrublands Important deserts and xeric shrubland areas within the Oriental Region include the Thar Desert (a.k.a. the Great Indian Desert) and the Rann of Kutch, a large area of salt marshes combined with arid grasslands and saline deserts located mostly in north-western India (Gujarat), as well as in

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the southern tip of Sindh province, Pakistan. The latter is divided into two main areas: the Great Rann of Kutch, and the Little Rann of Kutch. Together they serve as an important refuge for many species. The Indian wild ass or khur (Equus hemionus khur) once ranged from south-eastern Iran through southern Afghanistan and Pakistan (Sindh and Balochistan provinces) into western India. Heavily hunted at one time, its last refuge became the Little Rann of Kutch where, in 1947, the population was estimated at between 2000 and 5000. From 1958 to 1960, however, an infectious disease (South African horse sickness) had caused it to crash suddenly, and in 1969 an aerial census assessed the total number at just 400. The Indian Wild Ass Sanctuary was established in 1972 and helped considerably with the recovery. Today some 4800 live within the sanctuary and, increasingly, areas beyond. Sadly, the international boundary with Pakistan remains fenced, thus preventing any possibility of the animals reclaiming areas west. The white-browed bushchat (Saxicola macrorhynchus) is endemic to the arid areas of the north-western India and eastern Pakistan. It has disappeared from much of its historic range due to loss of habitat. The white-striped house gecko (Hemidactylus albofasciatus) is confined to a few localities in coastal west-central India (Maharashtra). Russell’s legless skink (Barkudia melanosticta) is known only from a single locality in central-eastern coastal India (Andra Pradesh). Sharma’s racer (Coluber bholanathi) is a type of snake known only from a few rocky hillocks and outcrops in southcentral India (Andhra Pradesh, Karnataka, Tamil Nadu, and Telangana).

Isolated Caves, Springs and Wells The Oriental Region is relatively rich in subterranean habitats, particularly limestone karst caves and underground rivers. The Kolar leaf-nosed bat (Hipposideros hypophyllus) is now confined to a single roosting cave in south-central India (Karnataka). Durga Das’ leaf-nosed bat (H. durgadasi) is confined to a small area of north-central India (Madhya Pradesh). The shield-nosed leaf-nosed bat (H. scutinares) is known only from a few karst localities in central Laos and central Vietnam. The Phou Khao Khouay leaf-nosed bat (H. khaokhouayensis) is known from a small area of central Laos (Vientiane province) and from the Cat Ba Biosphere Reserve in north-eastern Vietnam. The Thai leaf-nosed bat (H. halophyllus) is found patchily throughout western and central Thailand. All are threatened by disturbance of their roosting caves, habitat destruction, and (in some area) hunting for food. Kitti’s hog-nosed bat (Craseonycteris thonglongyai) is the smallest of the bats and arguably the world’s smallest mammal, being roughly the size of a bumblebee. The species is confined to limestone caves on the Thailand–Myanmar border, where it

The Oriental Region

is threatened by collectors who catch large numbers to make taxidermy mounts that are sold to tourists. Pyadalin Cave bent-toed gecko (Cyrtodactylus chrysopylos) is confined to a single cave within Pyadalin Cave Wildlife Sanctuary in north-central Myanmar (Shan State). The cave gliding tree frog (Rhacophorus spelaeus) is known only from limestone karst caves in and around the Khammouane Limestone National Biodiversity Conservation Area in central Laos (Khammouane province). The cave brook carp (Neolissochilus subterraneus) is known only from subterranean streams within Tham Phra Wangdaeng Cave, west-central Thailand (Pitsanulok province). The blind cave barb (Poropuntius speleops) is confined to an underground stream system in Phatewada Cave, within the Phu Khiew Wildlife Sanctuary, northern Thailand (Chaiyaphum province). Musae’s bangana (Speolabeo musaei) is known only from a few specimens collected from the underground course of the Xe Bang Fai River in central Laos. The Khammouane cavefish (Troglocyclocheilus khammouanensis) is known only from a single specimen collected in 1998 from central Laos (Khammouane province). Alikunhi’s blind catfish (Horaglanis alikunhii) is known only from a single specimen collected from subterranean wells in the village of Parappukara, in Kerala, south-western India. The Western Ghats blind cave catfish (H. krishnai) is confined to wells, where it migrates from one to another through underground channels. Several stone loaches of the genus Schistura are threatened by declining water levels and/or overcollection for the international pet trade. Dean Smart’s stone loach (S. deansmarti) is known only from subterranean streams in Tham Phra Sai Ngam Cave, in the Nernmaprang karst formation within Thung Salaeng Luang National Park, northern Thailand (Phitsanulok province). Spies’ stone loach (S. spiesi) is known only from Tham Prawangdaeng Cave in northern Thailand (Phitsanulok province). The Laotian stone loach (S. kaysonei) is known only from specimens collected from subterreanean steams within a cave system in Laos (Khammouane province). Spekul’s stone loach (S. spekuli) is confined to a single limestone cave in the centre of Chin Chu Chai village, northern Vietnam (Lai Chau province). Jarutanin’s stone loach (S. jarutanini) is known only from specimens collected from an underground stream in Lam Klong Ngu Cave, western Thailand (Kanchanaburi province). The Sai Yok Noi stone loach (Nemacheilus troglocataractus) is an eyeless species known only from a single subterranean stream within Sai Yok Noi Cave, in the Mae Klong River drainage of west-central Thailand (Kanchanaburi province). The Calicut swamp eel (Rakthamichthys digressus), Rosen’s swamp eel (R. roseni) and Eapen’s swamp eel (R. eapeni) are all known only from subterranean wells in southwestern coastal India (Kerala).

The Pang Mapha Karst Formation The Pang Mapha karst formation is located in north-western Thailand (Mae Hongson province). It has a large system of underground caves, some of which are connected. The waterfall-climbing cave fish (Cryptotora thamicola) is known only from eight subterranean sites within the Pang Mapha karst formation. The total population is thought to be less than 2000. The Pang Mapha stone loach (Schistura oedipus) is also confined to streams within the Pang Mapha cave system.

Lakes, Rivers, and Marshes Mainland Asia features a number of large river systems, but in general is much poorer in lakes than is Eurasia. There are, however, many swamps along the watercourses and in the deltas. Because of the disappearance of wetlands, populations of aquatic vertebrates, mostly birds, are becoming increasingly threatened. The vast rice fields in South East Asia have become an important habitat for many species. The Ganges River dolphin (Platanista gangetica gangetica) was formerly found in large groups in both the Ganges and Brahmaputra rivers as well as their tributaries, but is now seriously threatened by pollution and dam construction. The Sindh smooth-coated otter (Lutrogale perspicillata sindica) is confined to the wetlands of north-eastern Pakistan (Khyber Pakhtunkhwa, Punjab and Sindh provinces). The giant ibis (Thaumatibis gigantea) is a very rare wading bird nowadays mostly confined to northern and eastern Cambodia, with a few birds from the same population occasionally observed in southern Laos and southern Vietnam. The species formerly occurred in south-eastern and peninsular Thailand but has disappeared from there. In 2015 the total population was estimated at less than 300. The greater adjutant (Leptoptilos dubius) is a type of large scavenging stork that was historically widespread and common across much of South and continental South East Asia, but which declined dramatically during the first half of the twentieth century due to habitat destruction, hunting and egg collection, and improved human sanitation. Today only three breeding populations are known, two in north-eastern India and another in Cambodia. The species disperses widely at other times. The white-bellied heron (Ardea insignis) was historically known from the eastern Himalayan foothills of north-eastern India, Nepal and Bhutan through Bangladesh, much of Myanmar and south-western China (Yunnan and possibly Tibet). Highly sensitive to human disturbance, the species has been extirpated in many areas due to habitat destruction. The pink-headed duck (Rhodonessa caryophyllacea) historically had an extensive but discontinuous range in the wetlands of north-eastern India, adjacent areas of Bangladesh, north-western Myanmar and perhaps Nepal, where it seems never to have been common. Intensively hunted, the last known wild specimens (a flock of six birds,

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The Indo-Malaysian Realm Figure 5.4 Pink-headed duck: mounted specimen (Credit: unknown)

of which one was shot) were observed at Lake Manroona in 1947, with a few others surviving around the same time in captivity. While there have been occasional reports of its continued existence in recent years these appear to be based on misidentifications, and the species is most likely extinct. The marsh crocodile or mugger (Crocodylus palustris) is a medium-sized species found from southern Iran and Pakistan to the Indian subcontinent and Sri Lanka. It has become locally extinct over large parts of its range, with viable populations now only occurring in protected areas. The gharial (Gavialis gangeticus), a fish-eating crocodile from the slow-moving rivers of southern Asia, was long exploited for its skin. Historically it ranged from the Indus River in present-day Pakistan to, perhaps, the Irrawaddy River in Myanmar. It has now been extirpated from the latter country and perhaps Pakistan, Bangladesh and Bhutan as well owing to habitat alteration and other causes, and is now confined to three widely separated subpopulations in northern India and Nepal. The peacock softshell turtles (Nilssonia) are endemic to the rivers of South and South East Asia, where they are threatened by overcollection for East Asian food markets. Gray’s peacock softshell turtle (N. hurum) remains fairly widespread in India, Pakistan, Nepal and Bangladesh. Cuvier’s peacock softshell turtle (N. gangetica) is found in India, Pakistan and Bangladesh. Leith’s peacock softshell turtle (N. leithii) is found in peninsular India (Andhra Pradesh, Karnataka, Kerala, Madhya Pradesh, Maharashtra, Tamil Nadu and

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Odisha). The eastern peacock softshell turtle (N. formosa) is found in Myanmar and possibly Thailand. The Indian narrow-headed softshell turtle (Chitra indica) remains widespread in Pakistan, India, Nepal and Bangladesh but in very low densities, even in protected areas. It is threatened by hunting and habitat destruction. The Burmese flapshell turtle (Lissemys scutata) is a littleknown species from the Irrawaddy and Salween river drainages of Myanmar, north-eastern Thailand, and possibly southwestern China (Yunnan). It is traded in large numbers in East Asian food markets. Roofed turtles of the genus Batagur are a group of large river turtles from South and South East Asia. Historically very common, they have consequently been regarded as a useful food resource and have been driven to the point of extinction by intensive slaughter and overcollection of their eggs. The Burmese roofed turtle (B. trivittata) was long thought to be extinct, not having been reliably reported since 1935 despite mass trade of riverine turtles between Myanmar and China. It was rediscovered in small numbers in 2002, however, and although functionally extinct in the wild survives in captivity. The three-striped roofed turtle (B. dhongoka) from India and Bangladesh has disappeared from much of its former range. The red-crowned roofed turtle (B. kachuga) from India, Nepal and Bangladesh is also highly threatened. The Assamese roofed turtle (Pangshura sylhetensis) is a rare species known only from a few specimens collected in

The Oriental Region

parts of Bangladesh, north-eastern India (Assam) and possibly Bhutan. The crowned river turtle (Hardella thurjii) occurs in the Ganges, Brahmaputra and Indus river drainages of Pakistan, northern India and Bangladesh, where it is considered threatened by various factors. Spengler’s black-breasted leaf turtle (Geoemyda spengleri), while still relatively widespread in mainland South East Asia, is heavily threatened by overcollection for food, the international pet trade, and for use in ‘traditional medicine’. The Burmese eyed turtle (Morenia ocellata) is found in Myanmar and possibly southern China (Yunnan), where it was so heavily depleted by overharvesting for food that it actually disappeared from food markets by the end of the twentieth century. The Indian eyed turtle (M. petersi) is found in northeastern India and Bangladesh. The four-eyed turtle (Sacalia quadriocellata) is found in southern China, Hainan, and parts of Laos and Vietnam. The species is threatened by overcollection for its shells and bones, which are used in ‘traditional medicine’. Two South East Asian subspecies of big-headed turtle (Platysternon megacephalum), previously discussed, are threatened by overcollection for use as food. The Pegu bigheaded turtle (P. m. peguense) is found in Myanmar and Thailand, while Shiu’s big-headed turtle (P. m. shiui) occurs in Cambodia, Laos and Vietnam. The giant Asian pond turtle (Heosemys grandis) is found in Cambodia, Vietnam and parts of Laos, where it has been greatly reduced in number due to overcollection for Asian food markets. The black pond turtle (Geoclemys hamiltonii) occurs in the Indus, Ganges, and Brahmaputra river drainages of Pakistan, northern India, Bangladesh, and Nepal. Widely collected for local consumption up to the 1980s, it is now threatened primarily by harvesting for East Asian food markets as well as for use in the international pet trade. The Annam pond turtle (Mauremys annamensis) was historically endemic to a small area of central Vietnam, where it was reportedly abundant in the 1930s. Heavily harvested, all field surveys after 1941 failed to locate it although the species continued to be seen occasionally in food markets. In 2006 a small wild population was discovered in Quang Nam province) and a few are being kept and bred in captivity. Jagor’s water snake (Enhydris jagorii) is known with certainty only from a small, human-modified water system in north-central Thailand. The Mada paddy frog (Micryletta erythropoda) is known only from wetlands within the Ma Da Forest in southern Vietnam (Dong Nai province). Thin Thin’s stream toad (Ansonia thinthinae) is known only from the Tanintharyi Nature Reserve in peninsular Myanmar, but may occur in adjacent areas of Thailand. Swan’s burrowing frog (Sphaerotheca swani) is known only from a small area of sandy marshland in south-eastern Nepal.

The roughback whipray (Fluvitrygon kittipongi) is confined to the Mae Klong and Chao Phraya rivers of southwest-central Thailand. The ornate-finned snakehead (Channa ornatipinnis) is known only from a single locality in south-central Myanmar (Rakhine state). The Malabar snakehead (C. diplogramma) is known only from a few localities within the Western Ghats of south-western India (Kerala and Tamil Nadu states). Both are threatened by overfishing and habitat destruction. The Arakan snakehead (Channa pulchra) is a type of fish confined to streams in the Arakan Mountains of south-central Myanmar. The giant carp (Catlocarpio siamensis) is another enormous species that can reach up to three metres in length. Historically found in the larger rivers and floodplain areas in the Mae Klong, Mekong and Chao Phraya drainages of Thailand, Cambodia, Laos and Vietnam, it has been severely depleted by overfishing. Wild populations no longer occur in the Chao Phraya River, and the species is very rare in the Thai and Laos Mekong and its associated tributaries. The Sandkhol carp (Thynnichthys sandkhol) occurs in a number of river systems and reservoirs in peninsular India, but is everywhere threatened by dams, pollution, and introduced species. The Nukta carp (Schismatorhynchos nukta) occurs in the Krishna and Kaveri river drainages of south-western India, where it is threatened by habitat destruction and degradation. The finely striped shuttle-like carp (Luciocyprinus striolatus) is a large, predatory species known from a few river drainages in Laos and Vietnam, and at least formerly from southern China as well. It is threatened primarily by illegal dynamite fishing within the deep pools that it favours. The small-scaled mud carp (Cirrhinus microlepis) was historically known from the Chao Phraya and Mekong river drainages of Thailand, Laos, Cambodia and Vietnam. The Chao Phraya population was extirpated decades ago due to pollution, damming and overfishing, and the species is still commercially targeted in the Mekong. The Mrigal mud carp (C. cirrhosus) was historically found throughout the Godavari, Krishna and Kaveri river drainages of south-western India, but suffered major decline due to competition with introduced fish species. Today the only part of its native range where it still survives is in the Kaveri River, although it has also been introduced to many countries outside of India. The Mandalay mud carp (Henicorhynchus inornatus) is known only from a specimen collected from a market in Myanmar in the 1990s, of uncertain provenance. The Chittagong perch (Badis chittagongis) is known only from a few hillstreams in south-eastern Bangladesh. The Siamese tiger perch (Datnioides pulcher) was historically widespread in Indochina within the Chao Phraya, Mae Klong and Mekong river drainages, but has suffered considerable declines due to overcollection for the international pet trade. The species has been extirpated from Thailand and in a number of other areas.

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The Canara pearlspot (Etroplus canarensis) is a rare type of cichlid confined to the Kumaradhara and Nethravati rivers in south-western India (Karnataka). It is much sought-after for use in the international aquarium trade. Pointon’s fish (Oxygaster pointoni) occurs in the Chao Phraya and Mekong river drainages of Thailand, Cambodia and Laos, where it is threatened by pollution and damming. Chaudhuri’s kingfish (Semiplotus cirrhosus) is known only from its original collection in 1919 from northern Myanmar. Three barbs of the genus Dawkinsia are endemic to the Western Ghats of south-western India. The mascara barb (D. assimilis) and the Aruli barb (D. arulius) are both confined to a few rivers in Kerala, Karnataka, and Tamil Nadu. Rohan’s barb (D. rohani) is known only from a few hillstreams in Tamil Nadu. Several barbs of the genus Puntius from the Western Ghats of southern and south-western India are seriously threatened by habitat destruction and degradation. The Deccan barb (P. deccanensis) is known only from a single locality in Maharastra. Recent surveys have failed to locate the species, and it may possibly be extinct. The Mahecola barb (P. mahecola) is a rare species confined to a few rivers in Kerala. The Arenatus barb (P. arenatus) is confined to the southern Western Ghats. The Mudumalai barb (P. mudumalaiensis) is confined to a few localities in Tamil Nadu. The crescent barb (P. crescentus) is known only from a few localities in Karnataka and Kerala. The Boloven barb (Poropuntius bolovenensis) is confined to the Boloven Plateau in southern Laos (Champasak province), where it is threatened by overfishing and dam construction. The yellow-tail brook barb (P. deauratus) is found patchily in the coastal river drainages of southern Vietnam. The Channa barb (Eechathalakenda ophicephalus) is confined to hillstreams within the Western Ghats of south-western India (Kerala and Tamil Nadu), where it is threatened by habitat destruction and introduced fish species. Suvatti’s barb (Hypsibarbus suvattii) is known for certain only from the Mae Klong River of Thailand, where it is known to be declining. Sporadic and rather unreliable records from Vietnam, Cambodia and southern Thailand may represent remnants of a once-continuous distribution pre-dating the last glacial period. Several barbs of the genus Hypselobarbus are endemic to India, where they are threatened by overfishing, habitat destruction and pollution. The beautiful little barb (H. pulchellus) is known only from its original collection in 1870 from south-western India (Karnataka). Surveys of this locality have failed to record the species, and it is most likely extinct. The Canara barb (H. lithopidos) is known only from museum specimens collected from south-western India (Karnataka) and was last recorded in the 1940s. It is possibly extinct. The Curmuca barb (H. curmuca) is confined to highland rivers and streams of the Western Ghats in south-western India (Kerala). The red Canarese barb (H. thomassi) is

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confined to the Western Ghats of south-western India (Karnataka and Kerala). Dobson’s barb (H. dobsoni) and the Kolus barb (H. kolus) are both found widely but patchily in southern India (Kerala, Karnataka, Tamil Nadu and Maharashtra). The humpback barb (H. mussullah) is known only from a few isolated localities within the Krishna River and possibly the Godavari River in south-western India (Karnataka, Maharashtra and Kerala). Three species of glass barb (Laubuka) are threatened by loss of habitat. The striped glass barb (L. fasciata) is confined to a few rivers in south-western India (Kerala). The Siamese glass barb (L. siamensis) is confined to peninsular Thailand. The leaping glass barb (L. caeruleostigmata) is found in the Mae Klong, Mekong and Chao Phraya drainages of Thailand, southern Laos and Cambodia. Several species of sawfin barb (Pethia) are threatened by habitat destruction and degradation. The Chennai sawfin barb (P. sharmai) is known only from two freshwater ponds in south-western India (Tamil Nadu). The ornate sawfin barb (P. ornata) is known only from the Lokchou River in northeastern India (Manipur) and from another locality in Myanmar. The indigo sawfin barb (P. setnai) is known only from two river drainages within the Western Ghats of southwestern India (Goa and Karnataka). The long-lined sawfin barb (P. macrogramma) and the Nan Kywe Chaung sawfin barb (P. nankyweensis) are both known only from a few streams near Myitkyina in northern Myanmar (Kachin state). The Shalyni sawfin barb (P. shalynius) is confined to hillstreams in north-eastern India (Meghalaya). The red line torpedo barb (Sahyadria denisonii) and the Chalakkudy barb (S. chalakkudiensis) are both confined to a few isolated streams within the Western Ghats of southwestern India (Kerala and Karnataka), where they are threatened by habitat destruction and overcollection for the international aquarium trade. Evezard’s baril (Barilius evezardi) is known only from a few historical records over a wide area of southern India. Jerdon’s baril (B. canarensis) is confined to a few localities in the Western Ghats of south-western India (Kerala and Karnataka), where it is threatened by habitat destruction and overcollection for the international aquarium trade. The Putitor mahseer (Tor putitora) is naturally distributed throughout the rivers and associated reservoirs of the south Himalayan drainage, from Pakistan (and possibly Afghanistan) to Myanmar. Populations have also been introduced into some Himalayan lakes, parts of peninsular India, and even to New Guinea. However, the species is everywhere under threat by overfishing and loss of habitat. The Malabar mahseer (T. malabaricus) is confined to a few rivers in southern India. The Wayanad mahseer (Barbodes wynaadensis) is confined to upland streams and rivers within a small area of the Western Ghats in south-western India (Karnataka and Kerala), where it is seriously threatened by overfishing and habitat degradation.

The Oriental Region

The Mekong fighting betta (Betta smaragdina) is found over much of the Mekong River drainage in central and eastern Thailand, Laos and northern Cambodia. The Siamese fighting betta (B. splendens), a popular aquarium species the world over, is endemic to the Mae Klong and Chao Phraya drainages of west-central Thailand, including the Isthmus of Kra. The black paradise fish (Macropodus spechti) is a type of gourami known only from the Thu Bon and Houng river drainages of central Vietnam. Day’s paradise fish (Pseudosphromenus dayi) occurs in the Western Ghats and the coastal areas of south-western India, where it is threatened by loss of habitat. The Jaintia danio (Danio jaintianensis) is confined to streams within the Jaintia Hills of north-eastern India (Meghalaya). The scarlet chameleonfish (Dario dario) is known only from tributary streams in the Brahmaputra drainage of northern India (West Bengal and Assam), although it may occur as well in Bhutan. It is likely threatened by habitat degradation and overcollection for the international aquarium trade. Several species of danio (Devario) are threatened by habitat destruction and overcollection for the international aquarium trade. McLelland’s danio (D. ostreographus) is known only from nineteenth-century historical records from north-eastern India (Assam and West Bengal). The Nilgiri danio (D. neilgherriensis) is known only from a few lakes and streams in south-western India (Tamil Nadu and Kerala). Hora’s danio (D. acuticephala) is known only from two localities in northeastern India (Manipur and Nagaland). The Assamese danio (D. assamensis) is known only from a few localities in northeastern India (Assam). The Manipur danio (D. manipurensis) is known only from single locality in north-eastern India (Manipur). Myer’s danio (D. strigillifer) is known only from two localities in central and northern Myanmar. The Cardamom garra (Garra hughi), Periyar garra (G. surendranathanii), Kalakad garra (G. kalakadensis) and Silent Valley garra (G. menoni) are all confined to montane streams within the Western Ghats of south-western India. The panda garra (G. flavatra) is known only from three localities within the Arakan Mountains of south-western Myanmar. The Karen garra (G. imberbis) is confined to hillstreams within the Karen Hills of eastern Myanmar (Kayah state). The Hai Van Pass garra (G. apogon) is known only from creeks within the Hai Van Pass of central Vietnam. Two species of freshwater pufferfish, the Malabar pufferfish (Carinotetraodon travancoricus) and the dwarf Malabar puffer (C. imitator), are both confined to a few rivers in coastal south-western India (Kerala and southern Karnataka), where they are threatened by loss of habitat and overcollection for the international aquarium trade. The Deccan labeo (Labeo potail) remains fairly widespread within the Western Ghats of south-western India but has undergone considerable declines due to habitat destruction and degradation, overfishing, and introduced species.

The lipped algae-eater (Horalabiosa joshuai) and the Palani algae-eater (H. palaniensis) are both known only from fast-flowing streams in south-western India. Krempf’s sharpbelly (Hemiculter krempfi) is known only from the Cai and Da Rang rivers of central Vietnam (Khanh Hoa and Phu Yen provinces). The South Asian river shark (Glyphis gangeticus) is found within a narrow band of coastal rivers and estuaries of eastern and western India and western Bangladesh, where it is seriously threatened by overfishing and pollution. The redtail shark (Epalzeorhynchos bicolor) is a popular and common aquarium fish that was historically found in the lower Mae Klong, Chao Phraya and Bang Pakong drainages of Thailand. At one time thought to be extinct in the wild, most likely due to overcollection for the aquarium trade, it is now believed to survive within a small area of the Chao Phraya. The redfin shark (E. munense) is known from the Mekong, Chao Phraya and Mae Klong drainages of Thailand, Cambodia and Laos, where it is threatened mainly by pollution. The species is rare within the aquarium trade and is not known to be bred in commercial farms. The Siamese sharkminnow (Balantiocheilos ambusticauda) was historically known from the Mae Klong and Chao Phraya drainages of north-central Thailand. The species was severely impacted by overfishing, collection for the international aquarium trade and habitat destruction, and has not been recorded either in the wild or in captivity since 1974. It may be extinct. The Laotian sharkminnow (Osteochilus striatus) is found in the Xe Bangfai and Nam Mang drainages of northern and central Laos, where it is threatened by habitat degradation and dam construction. The Ratnagiri minnow (Parapsilorhynchus discophorus) and the elongated minnow (P. elongatus) are both confined to streams within the northern Western Ghats of southwestern India, where they are threatened by loss of habitat. Rahman’s torrent minnow (Psilorhynchus rahmani) is known only from a single small hillstream located north of Chittagong in south-eastern Bangladesh. The Melissa torrent minnow (P. melissa) and the Ann Chaung torrent minnow (P. pavimentatus) are both known only from the headwaters of the Ann Chaung drainage, on the western slopes of the Arakan Mountains in western Myanmar. Gokky’s torrent minnow (P. gokkyi) and the Man Chaung torrent minnow (P. piperatus) are each known only from a single locality on the eastern slopes of the Arakan Mountains, western Myanmar. Pierre’s labeo (Labeo pierrei) is a migratory species that is widespread in the Mekong and Chao Praya river drainages, but has declined extensively due to overfishing. Miller’s goby (Rhinogobius milleri) is known only from small creeks within the Nam Mat and Nam Mo river drainages of north-eastern Laos. The Huong Son goby (R. vinhensis) is known only from creeks and streams within the Annamite Mountains of north-central Vietnam (Ha Tinh province).

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The Cuc Phong sheatfish (Pterocryptis cucphuongensis) is known only from the Luong River drainage in northern Vietnam (Than Hoa province). The Malabar sheatfish (P. wynaadensis) is found widely but patchily in the Western Ghats of south-western India, where it is threatened by destructive fishing practices and pollution. Four shark catfish of the genus Pangasius are seriously threatened by overfishing, pollution and dam construction. Sanitwong’s shark catfish (P. sanitwongsei) is confined to the Chao Phraya and Mekong drainages in Cambodia, southwestern China (Yunnan), Laos, Thailand and Vietnam, where it has been depleted to the point of extinction. The Myanmar shark catfish (P. myanmar) is known from the Irrawaddy and Salween drainages in Myanmar. The elongated shark catfish (P. elongatus) is found widely but patchily in Thailand, Laos, Cambodia, and Vietnam, where it is everywhere uncommon. Krempf’s shark catfish (P. krempfi) is a freshwater and marine species that feeds in coastal waters of the South China Sea and migrates up the Mekong River to spawn. The Siamese flat-barbelled catfish (Platytropius siamensis) historically occurred in the Chao Phraya and Bang Pakong river drainages of south-central Thailand. Damming, canalization, pollution and the reclamation of wetlands around Bangkok all contributed to its decline, and the species has not been recorded since the late 1970s despite numerous surveys. It is most likely extinct. Children’s catfish (Silonia childreni) is confined to the Western Ghats of south-western India, where it has been recorded from the Krishna, Godavari and Kaveri rivers. It has undergone a significant decline due to overfishing and habitat destruction. The iridescent catfish (Pangasianodon hypophthalmus) is still found sporadically throughout the Chao Phraya, Mae Klong and mid to lower Mekong drainages of Indochina, but is everywhere threatened by overfishing. It is a very common aquaculture species, however, particularly in Vietnam and Thailand, and has been widely introduced outside of its native range (e.g. Myanmar and Indonesia). The sun catfish (Horabagrus brachysoma) is found patchily in lakes, rivers and backwaters in south-western India (Kerala and Karnataka), where it is threatened by overfishing and habitat degradation. The Quang Binh catfish (Hemibagrus centralus) is known only from hillstreams within the Annamite Mountains of central Vietnam (Quang Binh province). Mitchell’s catfish (Pseudeutropius mitchelli) is confined to a few rivers in the Western Ghats of south-western India (Kerala), where it is threatened mainly by habitat destruction and degradation. Bocourt’s catfish (Mystus bocourti) remains fairly widespread in the Chao Phraya and Mekong drainages of Cambodia, Laos, Thailand and Vietnam, but is everywhere rare and declining due to damming and pollution. Hamilton’s airbreathing catfish (Clarias magur) remains fairly widespread within the Ganges and Brahmaputra river

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drainages of northern India, Nepal, Bhutan and Bangladesh, but suffered considerable declines due to overfishing, loss of habitat and introduced species. The Thirumurthi airsac catfish (Heteropneustes longipectoralis) is known only from a single specimen collected in the 1990s in an irrigation reservoir of the Thirumurthi Dam, within the Anaimalai Hills of south-western India (Tamil Nadu). The Xiangkhouang bat-catfish (Oreoglanis macronemus) is known only from four specimens collected in the mid 1920s from the Xiangkhouang Plateau of northern Laos. The short-finned mountain catfish (Glyptothorax brevipinnis) is known only from four specimens collected in 1923 from an undefined locality in India. Sais’ mountain catfish (G. saisii) is known for certain only from the Chota Nagpur Plateau in northern India (Jharkand). The Cardamom mountain catfish (G. coracinus) is known only from the short coastal drainages on the western face of the Cardamom Mountains in south-western Cambodia. House’s mountain catfish (G. housei) and the Anamalai mountain catfish (G. anamalaiensis) are both confined to the Anamalai Hills of south-western India (Tamil Nadu and Kerala). The Madraspatana catfish (G. madraspatanus), Poona catfish (G. poonaensis) and Trewavas’ catfish (G. trewavasae) are all confined to montane streams within the Western Ghats of southwestern India. The Arakan stream catfish (Akysis vespertinus) is known only from the western face of the Arakan Mountains in southcentral Myanmar. The Stung Chhay stream catfish (A. clinatus) is known only from a single hillstream in south-western Cambodia. The Travancore naked catfish (Batasio travancoria) is confined to hillstreams within the Western Ghats of southwestern India, where it is threatened by overfishing and habitat destruction. The Sharavati naked catfish (B. sharavatiensis) is confined to the Joginmatha and Yennehole tributaries of the Sharavati River and to the Vimalanathi tributary of the Tunga River, in south-western India (Karnataka). It is threatened by intermittent water flow in the hillstreams where it lives due to dams. The elongated naked catfish (B. elongatus) is known only from hillstreams within the Arakan Mountains of southcentral Myanmar. Three loaches of the genus Sewellia are threatened by overfishing and habitat destruction. The small-lined loach (Sewellia lineolata) is confined to a few coastal drainages in south-central Vietnam. Roberts’ loach (S. pterolineata) is known only from the Ve and Tra Khuc rivers in south-central Vietnam. The kneecap loach (S. patella) is known only from the Ba (Da Rang) and Sesan basins of central Vietnam (Gia Lia and Kon Tum provinces). The splendid loach (Yasuhikotakia splendida) is known only from a small area of south-west-central Thailand, where it is threatened by dam construction. The black-lined loach (Ambastaia nigrolineata) is found widely but patchily in southern China (Yunnan), Laos,

The Oriental Region

Vietnam, and Thailand, where it is threatened by dam construction and pollution. The Santhampara loach (Ghatsa santhamparaiensis) is known only from two streams within the Western Ghats of south-western India (Kerala). The Travancore loach (Travancoria jonesi) is known only from two montane streams in the Western Ghats of southwestern India (Kerala), where it is threatened by habitat degradation and introduced species. The spearfin loach (Acanthocobitis pavonacea) is found in the Brahmaputra and Ganges river drainages, where it is threatened by habitat destruction and degradation. Rendahl’s weather loach (Misgurnus multimaculatus) is known only from a few specimens collected in the 1940s from central coastal Vietnam (Hué province). The Western Ghats striped loach (Botia striata) is confined to mountain streams within the Western Ghats of southwestern India (Karnataka and Maharastra). The ladder loach (B. rostrata) is confined to highland streams within the lower Ganges and Brahmaputra drainages of north-eastern India (Mizoram, West Bengal, Nagaland and Tripura) and Bangladesh. The slender stone loach (Balitora mysorensis) is confined to mountain streams within the Western Ghats of southwestern India (Kerala and Karnataka). The Khao Soi Dao stone loach (B. meridionalis) is known only from its original collection in 1978 from a small area of south-central Thailand (Chanthaburi province). The Banar stone loach (Nemacheilus banar) is known only from a few rivers in south-central Vietnam. The Shimoga stone loach (Nemachilichthys shimogensis) is confined to the Western Ghats of south-western India (Maharashtra and Karnataka), where it is threatened by dam construction. The Kerala stone loach (Indoreonectes keralensis) is known only from a few localities within the Western Ghats of southwestern India (Kerala), where it is threatened by introduced species and pollution. Day’s stone loach (Mesonoemacheilus pulchellus) is confined to a few rivers within the Western Ghats of southwestern India (Kerala). Herre’s stone loach (M. herrei) is confined to a very small area within the Indira Gandhi Wildlife Sanctuary in south-western India (Kerala and Tamil Nadu), where it is threatened by pesticide run-off from tea plantations. Petrubanarescu’s dwarf stone loach (M. petrubanarescui) is known only from the Nethravati and Kabani rivers within the Western Ghats of south-western India (Karnataka and Kerala). It is threatened by sand mining and pollution. A number of stone loaches of the genus Schistura are threatened by habitat degradation, pollution, overcollection for the international aquarium trade and dam construction. McClelland’s stone loach (S. zonata) is known only from its original collection in 1839 from an undefined locality in northeastern India (Assam). Pellegrin’s stone loach (S. nasifilis) is known only from its original collection in 1936 from the Ko

and Tan rivers of south-central coastal Vietnam. It has not been recorded since, and may be extinct. The Nam Xi stone loach (S. sertata) is known only from a single small stream in northern Laos. The Manipur stone loach (S. chindwinica) is known only from the Barak and Brahmaptura drainages of north-eastern India (Manipur). The Nam Mo stone loach (S. finis) is known only from small streams in the headwaters of the Mo River, on the border between northern Laos and northern Vietnam. The Pu Mat stone loach (S. pumatensis) is known only from creeks and small rivers within Pu Mat National Park in northern Vietnam (Nghe An province). Singh’s stone loach (S. singhi) is known only from a single locality in north-eastern India (Nagaland). Sokolov’s stone loach (S. sokolovi) is confined to a few rivers in south-central Vietnam. Yersin’s stone loach (S. yersini) is confined to a few mountain streams in south-central Vietnam. The parrot stone loach (S. psittacula) is confined to the Cam Lo and Ben Hai rivers in central Vietnam (Quang Tri province). The Huong stone loach (S. huongensis) is confined to the Huong and Cam Lo rivers in central Vietnam (Thua Thuen Hue and Quang Tri provinces). The Shan stone loach (Physoschistura rivulicola) is known only from Lake Inlé and part of the Salween River drainage in central Myanmar (Shan State). The Garo Hills stone loach (Aborichthys garoensis) is known from a few hillstreams in north-eastern India (Meghalaya and Arunachal Pradesh). The striated stone loach (Longischistura striatus) is known only from a few localities within the Western Ghats of southwestern India (Kerala, Karnataka and Tamil Nadu). The Nan Kwe eel-loach (Pangio lumbriciformis) is known only from Nan Kwe stream near Myitkina in northern Myanmar. The Bombay swamp eel (Monopterus indicus) and the Malabar swamp eel (M. fossorius) are both known only from a few localities within the Western Ghats of central and southwestern coastal India. The Hpa Lap spiny eel (Macrognathus obscurus) is known only from specimens collected from a single stream in northern Myanmar. The peacock spiny eel (M. pavo) is known only from a single stream within the Arakan Mountains of western Myanmar. The Kyeintali Chaung spineless eel (Pillaia indica) is known only from a single stream within the Arakan Mountains of western Myanmar. The Kachin spineless eel (P. kachinica) is known only from two streams in northern Myanmar (Kachin State).

Lake Inlé Lake Inlé and its tributaries lie in the Shan Hills of east-central Myanmar (Shan State). It is long, narrow and relatively shallow, but supports a remarkable number of endemic fishes. Unfortunately, the lake has been drastically modified by land-reclamation schemes and negatively affected by pollution and invasive species. The lake’s outlet disappears into the

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ground at some point, although it is thought that its waters ultimately reach a tributary of the Salween River. The Inlé snakehead (Channa harcourtbutleri) is known only from the Lake Inlé basin. The Inlé carp (Cyprinus intha) is confined to Lake Inlé, where it is threatened mainly by hybridization with introduced common carp (C. carpio). The Inlé mud carp (Gymnostomus horai) is confined to Lake Inlé. The Sawbwa barb (Sawbwa resplendens) is confined to Lake Inlé, where it is threatened by introduced fish species, overcollection for the international aquarium trade, and pollution. The Inlé barb (Systomus compressiformis) has not been reported since 1994, and is likely extinct. The red dwarf rasbora (Microrasbora rubescens) is confined to Lake Inlé. Two species of danio (Devario) are endemic to the Lake Inlé basin. The Inlé danio (D. auropurpureus) is confined to the lake itself, while Jayaram’s danio (D. jayarami) is known only from a few tributary streams. The emerald dwarf danio (Danio erythromicron) is confined to Lake Inlé. The Inlé catfish (Silurus burmanensis) has not been reported since its discovery in the 1960s. It may survive in some tributary streams. The Inlé stone loach (Yunnanilus brevis) is confined to Lake Inlé and its tributaries to the south. The Inlé spiny eel (Mastacembelus oatesii) is confined to Lake Inlé and surrounding streams.

Lake Indawgyi Lake Indawgyi is located in northern Myanmar (Kachin State). One of the largest lakes in South East Asia, it and the surrounding area have been set aside as the Indawgyi Lake Wildlife Sanctuary, which serves as a particularly important refuge for waterfowl. The Indawgyi perch (Badis corycaeus) appears to be confined to Lake Indawgyi and a few nearby streams. The Indawgyi glassfish (Pseudambassis roberti) is confined to Lake Indawgyi and a few nearby streams. The Indawgyi longtail catfish (Olyra horae) is known only from a few museum specimens originating from Lake Indawgyi. The Indawgyi spiny eel (Mastacembelus pantherinus) is known only from Lake Indawgyi and one of its tributaries.

The Kaveri River The Kaveri River (also known as the Cauvery or Ponni River) is located in south-central India. Originating in the foothills of the Western Ghats, it flows south and east through the states of Karnataka and Tamil Nadu before emptying into the Bay of Bengal. The Bhavani brook carp (Neolissochilus bovanicus) is known only from the Kaveri River and its tributary, the

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Bhavani River, at the base of the Nilgiri Hills. It suffered a massive population decline for unknown reasons during the late twentieth century and it is thought that only a single population, in the Mettur reservoir, may still survive, although it has not been recorded there since 1998. The Kaveri barb (Puntius cauveriensis) is confined to shallow pools within the Kaveri River drainage of the Western Ghats. The Nilgiri barb (Hypselobarbus dubius) is confined to four streams within the Kaveri River drainage, in the southern part of the Western Ghats. The Korhi barb (H. micropogon) is similarly restricted to the headwaters of the Kaveri River. Hora’s razorbelly minnow (Salmophasia horai) is confined to the headwaters of the Kaveri River in Karnataka. The Nilgiri catfish (Hemibagrus punctatus) is known with certainty only from the Kaveri River and its principal tributaries. Not seen since 1998, it may already be extinct. The Kaveri stone loach (Nemacheilus kodaguensis) is known only from a single stream within the Kaveri River drainage.

The Periyar River The Periyar River is located in south-western India (Kerala). It originates high in the Western Ghats and flows west before emptying into the Arabian Sea. The peninsular hilltrout (Lepidopygopsis typus) is known only from a few localities in the Periyar River drainage (including Lake Periyar), within the Periyar Tiger Reserve. The Periyar barb (Hypselobarbus periyarensis) is confined to the Periyar River drainage, including Lake Periyar. The Periyar garra (Garra periyarensis) is confined to the upper reaches of the Periyar River. The long-finned sharkminnow (Osteochilus longidorsalis) is confined to the Periyar River and its tributary, the Chalakudy River. The Periyar algae-eater (Crossocheilus periyarensis) is confined to the upper reaches of the Periyar River within the Periyar Tiger Reserve, where it is threatened by introduced fish species. The imperial catfish (Horabagrus nigricollaris) is confined to the Periyar River and its tributary, the Chalakudy River. The Periyar hillstream loach (Travancoria elongata) is confined to the Chalakudy and Pooyankutty tributaries of the Periyar River. The Periyar reticulated loach (Mesonoemacheilus periyarensis) and the Periyar blotched loach (M. menoni) are both confined to the upper reaches of the Periyar River within the Periyar Tiger Reserve

The Panniyar River The Panniyar River is located within the Western Ghats of south-western India. The Panniyar algae-eater (Horalabiosa arunachalami) is known only from a single locality within the Panniyar River, where it is seriously threatened by agro-based pollution originating from tea and cardamom plantations.

The Oriental Region

The Chalakudy River

The Godavari River

The Chalakudy River is located within the Western Ghats of south-western India. The Anaimalai loach (Homaloptera montana) is known only from streams passing through tea plantations within the Anaimalai Hills.

The Godavari River runs west to east through most of central India. It is the country’s second longest river after the Ganges. Day’s barb (Puntius ambassis) is known only from its original collection during the nineteenth century, most likely from the Godavari River. Fraser’s danio (Devario fraseri) appears to be confined to the upper reaches of the Godavari River in the Western Ghats (Maharashtra).

The Sharavati River The Sharavati River is located in south-western India. It originates in the central Western Ghats and flows west through Karnataka state before emptying into the Arabian Sea. The Nagodihole River The Nagodihole River is an upper tributary of the Sharavati River located in the Western Ghats (Karnataka). The Nagodihole stone loach (Schistura nagodiensis) is known only from a few rivulets within the Nagodihole River. The Kalkatte River The Kalkatte River is an upper tributary of the Sharavati River located in the Western Ghats (Karnataka). The Kalkatte stone loach (Schistura sharavathiensis) is confined to a short (c.100 m) stretch of the Kalkatte River above Dabbe Falls.

The Krishna River The Krishna River is located in south-western India (Maharashtra, Karnataka, Telangana and Andhra Pradesh). It originates in the Western Ghats and flows east through southcentral India before emptying into the Bay of Bengal. The Deccan catfish (Gagata itchkeea) is known for certain only from the Krishna River drainage. The Khavalchor catfish (Neotropius khavalchor) is known for certain only from the Krishna River drainage. The Gongwaree catfish (Eutropiichthys goongwaree) is known for certain only from the Krishna River drainage.

The Seetha River The Seetha River (also spelled Sita and Sitha) is located within the Western Ghats in south-western India (Karnataka). The Seetha danio (Betadevario ramachandrani) is confined to the upper Seetha River drainage. The Thuttinjet stone loach (Nemacheilus stigmofasciatus) is known only from the Seethanathi River, a small tributary of the Seetha River.

The Darna River The Darna River is a small upper tributary of the Godavari River located in the Western Ghats of west-central India (Maharashtra). General threats include pollution, habitat destruction and overfishing. The dwarf mahseer (Tor kulkarnii) is known only from four specimens collected from the Darna River in the early 1990s. The Darna barb (Puntius fraseri) is confined to the Darna River. The Deolali minnow (Parapsilorhynchus prateri) is known only from the Darna River. The species has not been reported since 2004 despite repeated surveys, and may be extinct. The Indravati River The Indravati River is located in east-central India (Chhattisgarh). The Indravati catfish (Clupisoma bastari) is known only from the Indravati River and the Ravishankar Sagar Reservoir.

The Namdapha River The Namdapha River is located in north-eastern India (Arunachal Pradesh). While nominally protected within Namdapha National Park, deforestation has been extensive, leading to sedimentation. The Namdapha danio (Devario horai) is known only from its original collection in 1983 from hillstreams within Namdapha National Park. It may be extinct. The Namdapha sheatfish (Pterocryptis indicus) is known only from a single specimen collected from the Namdapha River. The Namdapha loach (Lepidocephalichthys arunachalensis) is known only from hillstreams within Namdapha National Park.

The Lower Ganges River The Tunga River The Tunga River is located within the Western Ghats of southwestern India (Karnataka). The Tunga torrent minnow (Psilorhynchus tenura) is confined to two localities on the Tunga River. The Kudremukh mountain catfish (Glyptothorax kudremukhensis) is confined to the headwaters of the Tunga River within Kudremukh National Park, where it is seriously threatened by sedimentation.

The lower Ganges (Ganga in Hindustani) flows south and east through northern India into central Bangladesh, where it ultimately empties into the Bay of Bengal. Almora’s bangana (Bangana almorae) is known only from its original description in 1912 from north-western India (Uttar Pradesh). The torrent-loving catfish (Sisor rheophilus) and the muscular catfish (S. torosus) are each known only from a few specimens collected from the lower Ganges River drainage.

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The Ganges torrent catfish (Amblyceps tenuispinis) is known only from a single locality within the Ganges River drainage in northern India (Uttar Pradesh). The Sone River The Sone River is located in north-central India. It is the second largest of the Ganges’ southern tributaries after the Yamuna River. The Rihand catfish (Erethistoides pipri) is known only from its original collection in 1950 from the Rihand River, a tributary of the Sone River in Uttar Pradesh. An enormous dam has since been constructed at the type locality. The Bhagirathi River The Bhagirathi River is located in north-eastern India (West Bengal). The Bhagirathi catfish (Pseudolaguvia flavida) is known only from a single locality within the Bhagirathi River.

The Lower Brahmaputra River The lower Brahmaputra River, as here defined, flows through north-eastern India (Assam, Nagaland, Meghalaya and West Bengal) into central Bangladesh, where it merges with the Ganges River. The black peacock softshell turtle (Nilssonia nigricans) was historically found throughout the lower Brahmaputra River drainage of north-eastern India (Assam) and Bangladesh. The species was long thought to be extinct in the wild, surviving only in small numbers within a single artificial temple pond near Chittagong, in south-eastern Bangladesh. Recently, however, it has been rediscovered in northeastern India. The orange-spotted snakehead (Channa aurantimaculata) is known only from a single locality within the Brahmaputra drainage of north-eastern India (Assam). The Barca snakehead (C. barca) is known for certain only from the Brahmaputra drainage of north-eastern India (Assam and Nagaland) and Bangladesh. Both are threatened by overfishing and collection for the international aquarium trade. The Kanabos perch (Badis kanabos) and the Assam perch (B. assamensis) are both known only from a few localities in the lower Brahmaputra River drainage of north-eastern India (Assam). The spot-tailed catfish (Batasio spilurus) and the Meriani catfish (B. merianiensis) are both known only from the Brahmaputra River drainage of north-eastern India (Assam). The Jaintia stone loach (Schistura reticulofasciata) is confined to streams within the Jaintia Hills of north-eastern India (Meghalaya). The elongated stone loach (Physoschistura elongata) is known only from a single locality within the Brahmaputra drainage of north-eastern India (Meghalaya). The Jorai River The Jorai River is located in north-eastern India (Assam).

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The Jorai barb (Oreichthys crenuchoides) is known only from the Jorai River. The Priapus fish (Danionella priapus) is known only from the Jorai River. The Mansai River The Mansai River is located in north-eastern India (West Bengal). The Schutunga catfish (Erethistoides sicula) is known only from a few specimens collected from the Schutunga River, a tributary of the Mansai River. The Mahananda River The Mahananda River is located in north-eastern India (West Bengal). The Mahananda catfish (Conta conta) is known for certain only from the Mahananda River drainage.

The Surma–Meghna River Drainage The Surma–Meghna river drainage is one of the longest in Asia, and one of the three river complexes that form the Ganges–Brahmaputra delta. It rises in the Manipur Hills of north-eastern India as the Barak River and flows west, becoming the Surma River, before finally flowing south as the Meghna River to the Bay of Bengal by way of Bangladesh. The ornate catfish (Nangra ornata) is known only from the Surma–Meghna drainage in north-eastern Bangladesh (Sylhet province). The Barak River The Barak River (also known as the Ovurei River) is located in north-eastern India (Manipur, Nagaland, Mizoram, and Assam) and Bangladesh. Proposed dam construction poses a serious threat to a number of freshwater fish species. The Barak perch (Badis tuivaiei) is known only from the Tuivai and Irang rivers, tributaries of the Barak River. The Barak mahseer (Tor barakae) is confined to two streams within the Barak River drainage. The Manipur garra (Garra manipurensis) is confined to the Barak River drainage. The Barak sheatfish (Pterocryptis barakensis) is known only from a single locality within the Barak River. The Barak mountain catfish (Glyptothorax manipurensis) is confined to the Barak River. The Barak stone loach (Schistura tigrinum) is known only from a single locality within the Barak River drainage. The Baleshwar River is located in north-eastern India (Assam). The Baleshwar torrent minnow (Psilorhynchus amplicephalus) is confined to the Baleshwar River. The Jiri River is located in north-eastern India (Manipur). The Jiri catfish (Sisor barakensis) is known only from the Jiri River. The Iyei River is located in north-eastern India (Manipur). The Iyei catfish (Exostoma barakensis) is known only from the Iyei River.

The Oriental Region

The Lower Irrawaddy River The lower Irrawaddy, as here defined, begins at the confluence of the upper Irrawaddy and Chindwin rivers in central Myanmar. From there it runs directly south into the Irrawaddy delta, a low-lying expanse of land, streams and rivers located in coastal southern Myanmar, where it fans out across the Andaman Sea and Bay of Bengal. The Burmese keelback water snake (Xenochrophis bellula) is known only from two historical and once recent specimen collected from the lower Irrawaddy River. Blanford’s mud snake (Gyiophis maculosa) and Voris’ mud snake (G. vorisi) are both known only from a few specimens collected from the Irrawaddy delta in southern Myanmar. The ruby barb (Pethia padamya) is a popular aquarium fish that, in the wild state, is known only from the lower Irrawaddy drainage along with an artificial pond above the Anisakan Falls in central Myanmar. The Chindwin River The Chindwin is the largest tributary of the Irrawaddy River. It originates in northern Myanmar and north-eastern India (Nagaland and Manipur), and flows south before ultimately joining the lower Irrawaddy mainstream in central Myanmar. The Chindwin freshwater stingray (Makararaja chindwinensis) is known only from a single specimen collected in the Chindwin River. Dogar Singh’s baril (Barilius dogarsinghi) is known only from the Chindwin drainage in north-eastern India (Nagaland and Manipur). The Chindwin danio (Devario naganensis) is known only from its original description during the early twentieth century from the Chindwin River drainage. The Chindwin torrent catfish (Amblyceps tuberculatum) is confined to fast-flowing hillstreams of the Chindwin River drainage. Prashad’s stone loach (Schistura prashadi), Hora’s stone loach (S. kangjupkhulensis) and the reticulated stone loach (S. reticulata) are all confined to the Chindwin River drainage of north-eastern India (Manipur), where they are threatened by loss of habitat. The Uyu River (also known as the Yu or Laniye River) is located in north-eastern India (Manipur and Nagaland) and north-western Myanmar (Sagaing region). The Uyu barb (Pethia yuensis) is known only from the Lockchao and Maklang rivers, tributaries of the Uyu River in north-eastern India (Manipur). The Khujairok barb (Chela khujairokensis) is known only from Khujairok stream, a tributary of the Uyu River on the India/Myanmar border. The Chatrickong barbil (Barilius chatricensis) is known only from its original collection within the Chatrickong River, a tributary of the Uyu River in north-eastern India (Manipur). The ornate rasbora (Rasbora ornata) is confined to the Chatrickong and Lokchao rivers, tributaries of the Uyu River in north-eastern India (Manipur).

The Lokchao danio (Devario yuensis) is known from three localities within the Lokchao River, a tributary of the Uyu River in north-eastern India (Manipur) and northwestern Myanmar. The Uyu catfish (Myersglanis jayarami) is known only from the Uyu River, where it is threatened by habitat destruction and overfishing. The Uyu torrent catfish (Amblyceps torrentis) is confined to the Uyu River headwaters. The Tizu stone loach (Schistura nagaensis) is confined to hillstreams within the Tizu River, a tributary of the Uyu River in north-eastern India (Manipur and Nagaland). The Manipur River is located in north-eastern India (Manipur) and north-western Myanmar. Two species of barb (Pethia) are endemic to the Manipur River drainage. The Imphal barb (Pethia ater) is known only from the Imphal River, a tributary of the Manipur River in north-eastern India (Manipur). The Khuga barb (P. khugae) is confined to the Khuga River, a tributary of the Manipur River in north-eastern India (Manipur). The Sherou barbil (Barilius ngawa) is confined to the Sherou River, a tributary of the Manipur River in northeastern India (Manipur). The Khuga garra (Garra paralissorhynchus) is confined to the Khuga River, a tributary of the Manipur River in northeastern India (Manipur). The Chapki torrent minnow (Psilorhynchus microphthalmus) is confined to the Chapki stream, a tributary of the Manipur River in north-eastern India (Manipur). The Khuga stone loach (Schistura khugae) is confined to the Khuga River, a tributary of the Manipur River in northeastern India (Manipur). The Ukhrul River is located in north-eastern India (Manipur). The Wanze garra (Garra compressa) and the Litan garra (G. litanensis) are each known only from a single stream within the Ukhrul River drainage. The Ukhrul danio (Aspidoparia ukhrulensis) is known only from two localities within the Ukhrul River. The Ukhrul catfish (Pseudecheneis ukhrulensis) is confined to the Ukhrul River drainage. The Saman River is located in north-central Myanmar. The Ma Gawe torrent minnow (Psilorhynchus breviminor) is known only from the Ma Gawe River, a tributary of the Saman River. The Hwe-nga-sang River is located in east-central Myanmar. The Hwe-nga-sang stone loach (Schistura paucifasciata) is known only from its original description in 1929 from the Hwe-nga-sang River.

The Lower Salween River The lower Salween runs the length of eastern Myanmar, with its drainage basin extending into parts of north-western Thailand.

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Brown’s danio (Devario browni) is confined to the Salween River in central-eastern Myanmar, where it is threatened by overcollection for the ornamental fish trade. The flat-headed torrent catfish (Amblyceps platycephalus) is known only from a single specimen collected from the Salween River drainage in north-western Thailand. Rao’s stone loach (Physoschistura raoi) is known only from its original collection from the Salween River in central-eastern Myanmar (Shan State) in the 1920s. The Yuam River The Yuam River (Maenam Yuam in Thai) is located in northwestern Thailand (Mae Hong Son province). The Yuam bat-catfish (Oreoglanis heteropogon) is confined to the upper tributaries of the Yuam River drainage. The Yuam stone loach (Schistura alticrista) is known only from the Yuam River. The Mae Sa Nga River The Mae Sa Nga River (Maenam Mae Sa Nga in Thai) is located in north-western Thailand. The Mae Sa Nga bat-catfish (Oreoglanis laciniosus) is known only from two streams in the Mae Sa Nga drainage. The Ataran River The Ataran River (known as the Kasat River in Thailand) is located in south-eastern Myanmar (Karen state) and northwestern Thailand. The humphead glassy perchlet (Parambassis pulcinella) is confined to the Ataran River and its tributaries. It is threatened by overcollection for the international aquarium trade as well as by habitat degradation. The robust torrent minnow (Psilorhynchus robustus) is known only from a single stream at the headwaters of the Ataran River. The Ataran stream catfish (Akysis vespa) is known only from the Ataran River drainage. The Moei River The Moei River (Maenam Moei in Thai) is located on the south-western Thailand–south-eastern Myanmar border. The Moei catfish (Hemibagrus imbrifer) is known only from the Moei River. The Moei stone loach (Schistura moeiensis) is known only from four tributary streams within the Moei River drainage, but may be more widespread.

The Great Tenasserim River The Great Tenasserim River (also known as the Tanintharyi River) is located in south-eastern Myanmar. The Tenasserim mud carp (Cirrhinus rubirostris) is known only from the Great Tenasserim River, where it is potentially threatened by dam development. The emperor loach (Botia udomritthiruji) is confined to hillstreams within the upper Great Tenasserim River drainage.

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It was at one time a popular aquarium fish, but in recent years has become harder to obtain due to political instability.

The Chao Phraya River Drainage The Chao Phraya (Maenam Chao Phraya in Thai) is the major river drainage in north-western and central-western Thailand. It begins at the confluence of the Ping and Nan rivers and flows south for roughly 370 km to the Gulf of Thailand. Coleman’s barb (Discherodontus colemani) is an uncommon species from the upper Chao Phraya drainage in northwestern Thailand. The Chiang Mai goby (Rhinogobius chiengmaiensis) is confined to the upper Chao Phraya drainage, where it is threatened by dam construction and other water diversion projects. Hardman’s catfish (Pseudobagarius hardmani) is known only from its original collection from the Chao Phraya River in west-central Thailand. Buchanan’s mountain catfish (Glyptothorax buchanani) is known only from the upper Chao Phraya River drainage. The Doi Inthanon bat-catfish (Oreoglanis siamensis) is known only from the upper Chao Phraya drainage within Doi Inthanon National Park in northern Thailand (Chiang Mai province). Walton’s stone loach (Schistura waltoni) is confined to the Ping, Wang and Yom tributaries of the upper Chao Phraya drainage in north-western Thailand, where it is threatened by habitat destruction and degradation. The Ping River The Ping River (Maenam Ping in Thai) is located in northwestern Thailand (Chiang Mai and Mae Hong Son provinces). It is one of the two main tributaries of the Chao Phraya River. Nakasathian’s bat-catfish (Oreoglanis nakasathiani) is known only from the Ping River drainage within the Doi Chiang Dao Wildlife Sanctuary. The Ping stone loach (Schistura spilota) is confined to the upper Ping River drainage. The Taeng River (Nam Mae Taeng in Thai) is located in Chiang Mai province. Sinkler’s bangana (Bangana sinkleri) is known only from the upper Taeng River. The firebar danio (Devario maetaengensis) is confined to the Taeng River and tributary streams. The mini dragon stone loach (Schistura pridii) is confined to streams within the Doi Chiang Dao Wildlife Sanctuary. The Nai River (Mae Ping Nai in Thai) is located in Mae Hong Son province. Sudara’s bat-catfish (Oreoglanis sudarai) is known only from the Nai River, where it is threatened by loss of habitat and overfishing. The Nan River The Nan River (Maenam Nan in Thai) is located in northwestern Thailand (Nan province), and is the other of the two major sources of the Chao Phraya River.

The Oriental Region

Three species of bat-catfish (Oreoglanis) are confined to the Nan River drainage. The slender-tailed bat-catfish (O. tenuicauda) is confined to a small area of north-western Thailand. The Silaphet Waterfall bat-catfish (O. vicinus) is known only from Silaphet Waterfall. The Doi Bhuka bat-catfish (O. colurus) is known only from Doi Bhuka National Park, where it is threatened by dam construction. The Nan river loach (Hemimyzon nanensis) is confined to the Nan River drainage. The Nan stone loach (Schistura menanensis) is confined to the Nan River drainage. Smith’s stone loach (Sectoria atriceps) is confined to the Nan River drainage. The Yom River is located in west-central Thailand. The falcate chameleon catfish (Acrochordonichthys gyrinus) is confined to the Yom River. The dwarf loach (Yasuhikotakia sidthimunki) is confined to the Yom River. The Lower Chao Phraya River The lower Chao Phraya River is located in centralwestern Thailand. Chanard’s water snake (Enhydris chanardi) is known only from the Chao Phraya freshwater swamp forest in central Thailand, where it is threatened by expansion of the city of Bangkok. The Bangkok goby (Rhinogobius vexillifer) is known only from streams and tributary rivers within the lower Chao Phraya drainage, in what is now the city of Bangkok.

The Mae Klong River The Mae Klong River (Maenam Mae Klong in Thai, and also known as the Meklong River) is located in western central Thailand (Kanchanaburi, Ratchaburi, and Smut Songkhram provinces). It begins at the confluence of the Khwae Noi and Khwae Yai rivers and flows south to the Gulf of Thailand. The Mae Klong barb (Poropuntius melanogrammus) is confined to the upper Mae Klong River and adjacent areas, where it is threatened by dam construction and habitat degradation. Somphongs’ rasbora (Trigonostigma somphongsi) is confined to the lower Mae Klong drainage (Ratchaburi province). Long close to extinction in the wild, it is threatened by habitat destruction and degradation and harvesting for the international aquarium trade. The Mae Klong sheatfish (Pterocryptis buccata) is a blind species known from a few caves as well as surface streams within the Mae Klong drainage of Kanchanaburi province. The Khwae Noi River The Khwae Noi River (Maenam Khwae Noi in Thai) is located in Kanchanaburi province. The Khwae Noi naked catfish (Batasio tigrinus) and Khwae Noi torrent catfish (Amblyceps variegatum) are both known only from hillstreams within the Khwae Noi River drainage.

The Lower Mekong River Known as the ‘Amazon of South East Asia,’ the Mekong is one of the longest rivers in the world. Like the Yangtze, Ganges and Indus it starts among the glaciers of Tibet, where it is known as the Lancang River (see ‘Upper Mekong’). Fed by melting snow it begins its 4500-km journey through steep mountain gorges in China, gathering power from streams along the way and passing through the ‘Golden Triangle’ formed by the borders of Myanmar, Laos and Thailand, before heading into Cambodia and ultimately entering the South China Sea through a network of distributaries (the Mekong delta) in south-western Vietnam. The drainage is home to some 1300 fish species and a rich turtle fauna. The river is also one of the last strongholds of the Irrawaddy dolphin (Orcaella brevirostris). Wetlands in the basin, meanwhile, harbour rare species such as Siamese crocodile (Crocodylus siamensis) and sarus crane (Antigone antigone). General threats include overfishing, pollution and large-scale hydroelectric projects. The Mekong snail-eating turtle (Malayemys subtrijuga) historically occurred throughout the Mekong River drainage, but has been much reduced due to a variety of factors. Additional introduced populations in the Malay Peninsula, Sumatra and Java are small and, in some places, extirpated. The Tay Minh water snake (Enhydris innominata) is known only from a few specimens collected in the Mekong River drainage of southern Vietnam. The Mekong giant catfish (Pangasianodon gigas), one of the world’s largest freshwater fish, was abundant within the lower Mekong River drainage in the early 1900s, but began to disappear by the 1970s. Overfishing and loss of migratory routes owing to the construction of dams are to blame for the decline. While now protected, the species is very near extinction. The Mekong freshwater stingray (Hemitrygon laosensis) was historically endemic to the Mekong River drainage where it is known from Cambodia and northern Thailand. An additional introduced population lives in the Chao Phraya River in Thailand. It is threatened by heavy fishing and bycatch pressure, damming and habitat degradation. The Mekong shad (Tenualosa thibaudeaui) remains widespread in the Mekong River drainage but has suffered declines due to overfishing and dam construction. The Mekong tiger perch (Datnioides undecimradiatus) is confined to the lower Mekong drainage in Thailand, Laos, Cambodia and Vietnam, where it is threatened by overcollection both for food as well as the international aquarium trade. The bristly fish (Laocypris hispida) is known only from the Nam Leuk and Nam San (a small tributary of the Negum Reservoir) in northern Laos, where it is threatened by overfishing and loss of habitat. The Mekong giant salmon carp (Aaptosyax grypus) is confined to the Mekong River drainage of northern Cambodia, Laos, and Thailand, where it is threatened by overfishing and habitat degradation.

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Behr’s bangana (Bangana behri) remains widespread in the Mekong River drainage but has suffered declines due to overfishing and dam construction. Lagler’s barb (Hypsibarbus lagleri) is widespread within the middle Mekong River drainage of eastern Thailand, western Laos and north-eastern Cambodia, but is everywhere threatened by dam construction. The thin-tailed barb (Mystacoleucus lepturus) remains widespread in the Mekong River drainage but has suffered declines due to dam construction and pollution. The Mekong sharpmouth barb (Scaphognathops bandanensis) is a migratory species still found throughout much of the Mekong River drainage, but is everywhere threatened by overfishing and dam construction. The thick-lipped barb (Probarbus labeamajor) is found patchily throughout much the Mekong River drainage, where it is threatened by overfishing and habitat destruction. Deignan’s bitterling (Acheilognathus deignani) is confined to a few small tributaries in north-western Laos and northwestern Vietnam, where it is threatened by loss of habitat and overfishing. The elephant ear gourami (Osphronemus exodon) is confined to the Mekong River drainage, where it is threatened by dam construction and pollution. The Mekong betta (Betta stiktos) is known only from a small area of the Srepok River in north-eastern Cambodia, but is expected to occur as well in the lower Kong River in southern Laos and north-eastern Cambodia. The Mekong sharkminnow (Osteochilus brachynotopteroides) is known only from a small number of localities dispersed throughout the Mekong River drainage. Kottelat’s garra (Garra cyrano) is known from the Leuk, Theun and Kong tributaries in Laos, where it is restricted to the upper parts of the respective drainages in areas with strong current. The line-cheek goby (Glossogobius sparsipapillus) is known only from the tidal zone of the Mekong delta and from the lower parts of the Mekong River in southern Vietnam and southern Cambodia. The long-finned loach (Yasuhikotakia longidorsalis) is confined to the Mekong drainage of central Laos and northeastern Thailand, where it is threatened by dam construction. Baird’s stone loach (Schistura bairdi) is known only from the mainstream Mekong River of south-western Laos in the Khone Falls area, and from Kaoh Han Island in north-eastern Cambodia (Stung Treng province). Schultz’s stone loach (S. schultzi) is known from northern Thailand (Chiang Rai and Loei provinces) and is expected from adjacent areas of northern Laos (Bokeo province). Both are threatened by loss of habitat. The Mekong eel-loach (Pangio longimanus) is known only from specimens collected in 1997 from the confluence of the Leuk and Gnong rivers in north-central Laos. The belted serpent loach (Serpenticobitis cingulata) is confined to the Mekong River drainage of northern Laos and

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Thailand, where it is threatened by dam construction and pollution. The Khone Phapheng Falls are located within the mainstream Mekong River in southern Laos (Champasak province), near the border with Cambodia. It is the largest waterfall in South East Asia and the widest in the world. The Khone Falls river loach (Hemimyzon khonensis) is known only from a single specimen collected from Khone Falls. The Kok River The Kok River (Mae Nam Kok in Thai) is located in northwestern Thailand (Chiang Mai province) and Myanmar (Shan State). The Doi Tung bat-catfish (Oreoglanis suraswadii) is known only from two rocky forest streams near Doi Tung, northern Thailand. The beautiful stone loach (Schistura bella) and the shortheaded stone loach (S. breviceps) are both known only from the Kok River drainage. The Leuk River The Leuk River (Nam Leuk in Laotian) is located in northern Laos and north-eastern Thailand. The construction of the Nam Leuk Dam in the late 1990s, which diverted much of the water flow to the nearby Xan River, may have seriously threatened a number of endemic species. The Leuk danio (Devario acrostomus) is known only from forested streams within the Leuk River drainage. The bumblebee catfish (Pseudomystus bomboides) is known only from three specimens collected in 1997 from the Leuk River. The Tenura stone loach (Schistura tenura), Leuk stone loach (S. leukensis) and the bark stone loach (S. suber) are all known only from their original collection in the 1990s within the Leuk River catchment. Their known geographical range now occurs almost entirely downstream of the Nam Leuk Dam, where very little water now remains. The Leuk spineless eel (Chaudhuria fusipinnis) is known only from its original collection in 1997 from the Leuk River drainage. The Ngiap River The Ngiap River (Nam Ngiap in Laotian) is located in northern Laos. Delacour’s bat-catfish (Oreoglanis delacouri) is known only from the upper Ngiap River. The Ngiap stone loach (Schistura crabro) is known only from a single specimen collected in 1999. The Tha River The Tha River (Nam Tha in Lao) is located in north-western Laos (Luang Namtha province). Three species of loach (Schistura) are confined to hillstreams in the upper Tha River drainage, where they are

The Oriental Region

threatened by habitat destruction and degradation. The globeheaded stone loach (S. globiceps) and the nine-rayed stone loach (S. novemradiata) are each known only from a single stream. The russet stone loach (S. russa) is known only from two localities. The Khan River The Khan River (Nam Khan in Laotian) is located in northcentral Laos (Xiangkhouang province). The Khan bat-catfish (Oreoglanis frenata) is known only from a single locality within the Khan River drainage. The irregular stone loach (S. irregularis) is known only from two localities within the Khan River drainage. The Xhat River (Nam Xhat in Laotian) is located in northcentral Laos. The Xhat stone loach (Schistura xhatensis) is known only from its original collection in the upper Xhat River. The Banghiang River The Banghiang River (Se Banghiang in Laotian/Se Bang Hieng in Vietnamese) is located in southern Laos (Savannakhet province) and central Vietnam. It originates from the western side of the Annamite Mountains, and joins the Mekong near Savannakhet city. The Banghiang stone loach (Schistura latidens) is known only from a single locality in the Banghiang River. The Ou River The Ou River (Nam Ou in Laotian) is located in northern Laos (Phongsaly and Luang Prabang provinces) and north-western Vietnam (Dien Bien province). The pole stone loach (Schistura pertica) is known only from its original collection in 1997 from a small stream in northern Laos. The Nua River (Nam Nua in Laotian) is located on the Vietnam–Laos border. The Aramis stone loach (Schistura aramis) is known only from a few specimens collected from the Nua River. The Ngum River The Ngum River (Nam Ngum in Laotian) is a major tributary of the Mekong River located in northern Laos (Xiangkhoang and Vientiane provinces). The Houay Sala Yai stone loach (Schistura sigillata), sought stone loach (S. quaesita) and the freckled stone loach (S. ephelis) are all known only from the Ngum River. The Ngum river loach (Hemimyzon confluens) is known only from a single locality within the Ngum River, where it is threatened by mining activities and dams. The San River (Nam San in Laotian) is located in northern Laos (Vientiane province). General threats include pollution from mining activities, sedimentation from deforestation, and dam construction. The San goby (Rhinogobius albimaculatus) is known only from the San River.

The Quasimodo stone loach (Schistura quasimodo) and the San stone loach (S. coruscans) are both confined to the San River. The Theun River The Theun River (Nam Theun in Laotian, and also known as the Kading River) is located in central Laos (Khammouane and Bolikhamsai provinces). The Theun barb (Scaphognathops theunensis) is confined to the Theun and Gnouang catchments. It is seriously threatened by dam construction, which has flooded much of its habitat with reservoirs. The dark mahseer (Tor ater) is known only from two streams within the upper Theun River drainage. The Theun bitterling (Rhodeus laoensis) is known only from the Theun and Gnouang catchments. Formerly common, it suffered serious declines after the construction of numerous dams between 1999 and 2009, which now impact much of its range. The lined goby (Rhinogobius lineatus) is known only from the Theun River and its tributary, the Gnouang River. The Theun sheatfish (Pterocryptis inusitata) is confined to the Theun River drainage, where much of its original range was inundated by a reservoir while the remainder lies within a protected area. Kottelat’s stone loach (Schistura tubulinaris), the atra stone loach (S. atra), and the naked-backed stone loach (S. nudidorsum) are all confined to the Theun River drainage, where they are threatened by dam construction. The Phao River (Nam Phao in Laotian) is located in eastcentral Laos (Bolikamsai province). The Phao bat-catfish (Oreoglanis lepturus) is known only from a very short stretch (about 2 km) of a branch of the upper Phao River, where it is seriously threatened by habitat destruction and pollution. The Kong River The Kong River (Se Kong in Laotian; Song Se Kong in Vietnamese) originates in central Vietnam and flows 480 km through southern Laos and eastern Cambodia before finally joining the Mekong River. Kottelat’s barb (Poropuntius solitus), the confusing barb (P. consternans), the Aluoi barb (P. aluoiensis), and the lobocheiloides barb (P. lobocheiloides) are all confined to the Kong River drainage, where they are threatened by dam construction and overfishing. The Bolaven danio (Devario salmonatus) is confined to the Bolaven Plateau of south-eastern Laos, within the Kong River drainage. It is threatened by habitat destruction and degradation. The Kong mountain catfish (Glyptothorax filicatus) is confined to the Kong River drainage in central Vietnam and south-eastern Laos. The Kong stream catfish (Akysis bilustris) is known only from two localities within the Kong River drainage of southern Laos (Attapeu province).

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Diard’s loach (Sewellia diardi) is known only from the Kong River drainage in southern Laos. Tizard’s stone loach (Schistura tizardi), Rikik’s stone loach (S. rikiki), and the Boloven Plateau stone loach (S. bolavenensis) are all confined to the Kong River drainage, where they are threatened by dam construction. The Kong serpent loach (Serpenticobitis octozona) is an uncommon species confined to the Kong River drainage. The Bang Fai River (Xe Bang Fai in Laotian) is located in central Laos (Khammouane and Savannakhet provinces). It is notable for the Khoun Xe Cave, the largest river cave in the world. The Bang Fai sleeper goby (Terateleotris aspro) is confined to a small area of the Bang Fai River, downstream of the subterranean course and upstream of the Nam Theun 2 hydroelectric project. The Bang Fai stone loach (Schistura punctifasciata) is confined to the Bang Fai River downstream of the subterranean section. The Tonlé San River The Tonlé San River (also known as the Tonlé Se San River or Sesan River) flows through south-central Vietnam, southeastern Laos, and north-eastern Cambodia. The Tonlé San carp (Mekongina bibarba) is confined to the Tonlé San River. The Kon Tum stone loach (Schistura kontumensis) is known for certain only from a few specimens collected from the headwaters of the Tonlé San River in south-central Vietnam (Kon Tum province). The Pako River is located in south-central Vietnam (Kon Tum province). The butterfly loach (Sewellia breviventralis) is known only from a single locality within the Pako River. The Tonlé Sap River The Tonlé Sap River is located in central Cambodia (Siem Reap, Battambang, and Pursat provinces). Tsukawaki’s dragonet (Tonlesapia tsukawakii) is known only from a small area of the Tonlé Sap River. Tonlé Sap Lake is the largest freshwater lake in South East Asia. The Tonlé Sap water snake (Enhydris longicauda) is confined to the Tonlé Sap Lake floodplain. It is threatened by overcollection and loss of habitat.

The Thu Bon River The Thu Bon River (Song Thu Bon in Vietnamese) is located in central Vietnam (Quang Nam province). The Thu Bon barb (Spinibarbus brevicephalus) is confined to the Thu Bon River. The Thu Bon loach (Sewellia albisuera) is confined to the Thu Bon River, where it is seriously threatened by habitat destruction and overcollection for the pet trade.

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The Phong Nha River The Phong Nha River is located in central Vietnam (Quang Binh province). It is protected within Phong Nha-Ke Bang National Park. The Phong Nha-Ke Bang carp (Cyprinus exophthalmos) and Hien’s carp (C. hieni) are both confined to the Phong Nha River drainage. Trang’s danio (Devario trangi) and the Quang Binh danio (D. quangbinhensis) are both confined to hillstreams within the Phong Nha River drainage. The Yleng spined loach (Cobitis ylengensis), Ngô’s spined loach (C. squataeniatus), the longitaeniatus spined loach (C. longitaeniatus) and the Phong Nha spined loach (C. phongnhaensis) are all known only from limestone mountain streams within the Phong Nha–Ke Bang drainage.

The Lam River The Lam River (Song Lam or Song Ca in Vietnamese/Nam Khan in Laotian) originates in the Loi Mountains of Laos and crosses northern Vietnam before emptying into the Gulf of Tonkin. The Lam minnow (Opsariichthys bea) is known only from the Lam River in northern Vietnam (Nghe An province). The Lam stream goby (Rhinogobius nganfoensis) is confined to creeks and small rivers within the catchment area of the Lam River in northern Vietnam (Nghe An province). The Ha Tinh stone loach (Schistura antennata) is known from a few small headwater streams in northern Vietnam (Ha Tinh province).

The Ma River The Ma River (Song Ma in Vietnamese/Nam Ma in Laotian) is located in north-western Vietnam (Thanh Hoa and Son La provinces) and north-eastern Laos (Houaphan and Louangphrbang provinces). The Ma goby (Rhinogobius nammaensis) is known only from the Ma River drainage. The thick-tailed loach (Vanmanenia crassicauda) is known only from the Ma River drainage. The Chu River The Chu River (Song Chu in Vietnamese, and known as the Nam Xam River in Laos) is located in north-western Vietnam (Thanh Hoa province) and north-eastern Laos (Houaphan and Louangphrabang provinces). It is the largest tributary of the Ma River. The Chu barb (Acrossocheilus xamensis) is known only from two specimens collected from the Chu River in northeastern Laos (Louangphrabang province). The Nam Xam goby (Rhinogobius vermiculatus) is known only from small forest streams in north-eastern Laos (Houaphan province). The Chu sleeper goby (Sineleotris namxamensis) is known only from the Chu River drainage in Laos (Houaphan province).

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Miscellaneous Lakes, Rivers, and Marshes Lake Loktak is a large freshwater lake located in north-eastern India (Manipur). The Loktak barb (Pethia manipurensis) is confined to Lake Loktak, where it is threatened by habitat degradation. Lake Pookode is located in the Western Ghats of India (Kerala). The Pookode barb (Pethia pookodensis) is known for certain only from Lake Pookode. The Narmadha River is located in north-western India (Madhya Pradesh). Günther’s baril (Barilius radiolatus) is confined to the Narmadha River. The Nelligudda Reservoir is located in south-western India (Karnataka). The large razorbelly minnow (Salmophasia belachi) is known only from its original collection from the Nelligudda Reservoir in the 1990s. The Chaliyar River is located in south-western India (Kerala). The Chaliyar mountain catfish (Glyptothorax davissinghi) is known only from the Karimpuzha and Paanapuzha tributaries of the Chaliyar River, within the New Amarambalam Reserve Forest. The Kallada River is located in south-western India (Kerala). The exclamation barb (Dawkinsia exclamatio) is confined to the Kallada River drainage. The Pambar River is located in south-western India (Kerala). The Pambar banded loach (Mesonoemacheilus pambarensis) is known only from the Pambar River, within the Chinnar Wildlife Sanctuary. The Bharathapuzha River is located in south-western India (Kerala). The Kunthi catfish (Pseudolaguvia austrina) is confined to the Kunthi River, a tributary of the Bharathapuzha River in the southern Western Ghats. The Valapattanam River is located in south-western India (Kerala). The Urutty mountain catfish (Glyptothorax malabarensis) is known only from a single hillstream within the Valapattanam River drainage. The Tambraparniei River is located in south-western India (Tamil Nadu). The Tambraparniei barb (Dawkinsia tambraparniei) is confined to the upper and middle reaches of the Tambraparniei River in the Western Ghats. The Kaladan River (also known as the Chimtuipui River) is located in north-eastern India (eastern Mizoram) and western Myanmar (Rakhine State). It remains one of the largest rivers in the world to be completely unfragmented by dams. The Kaladan mountain catfish (Glyptothorax chimtuipuiensis) is known only from the Kaladan River. The Nambul River is located in north-eastern India (Manipur). The Nambul garra (Garra nambulica) is known only from one or two streams within the Nambul River drainage, where it is threatened by loss of habitat. The Iyei River is located in north-eastern India (Manipur). The Iyei stone loach (Schistura minutus) is known only from the Iyei River. The Ichamati River is located in north-eastern India (West Bengal) and western Bangladesh. Andrew’s leaf fish (Nandus andrewi) is known only from the Ichamati River.

The Feni River is located in north-eastern India (Tripura) and south-eastern Bangladesh. The Feni catfish (Pseudolaguvia inornata) is known only from the Koilla Khal creek, a tributary of the Feni River. The Ganges–Brahmaputra delta is located in southern Bangladesh. Formed by the large, sediment-laden flows of the Ganges, Brahmaputra and Surma–Meghna rivers, it is the world’s largest river delta. The Tangail catfish (Nangra bucculenta) is known only from a small floodplain area between two large tributaries of the Ganges in central Bangladesh. The Sittaung River is located in south-central Myanmar. Flowing from the Shan Hills to the Andaman Sea, the Pegu Range separates its basin from that of the Irrawaddy. The Sittaung stream catfish (Akysis portellus) is known only from hillstreams in the Sittaung River drainage. The Bang Pakong River (Maenam Bang Pakong in Thai) is located in south-central Thailand. The club-barbel sheatfish (Ceratoglanis pachynema) is known for certain only from the lower Bang Pakong River drainage. A specimen was recorded from a market in Savannakhet, Laos, but was not preserved. The Kampot River is located in southern coastal Cambodia. The Kampot sheatfish (Pterocryptis bokorensis) is known only from the Kampot River. The Dong Nai River is located in southern Vietnam (Lam Dong province). The Dong Nai stone loach (Schistura dalatensis) is known only from the Dong Nai River drainage. The Da Rang River (Song Da Rang in Vietnamese) is located in south-central coastal Vietnam (Kon Tum, Tuy Hoa, and Phu Yen provinces). The Da Rang bumblebee catfish (Pseudomystus sobrinus) is known only from the Da Rang River drainage. The Huong River (Song Huong in Vietnamese) is located in central coastal Vietnam (Thua Thien-Hue province). The Huong barb (Hypsibarbus annamensis) is known only from specimens collected from the Huong River in 1936. The Mong Mo River is located in coastal central Vietnam. The Mong Mo stone loach (Schistura susannae) is confined to the Mong Mo River. The Vu Gia River (Song Vu Gia in Vietnamese) is located in central Vietnam (Thua Thien-Hue, Da Nang, and Quang Nam provinces). The Vu Gia gudgeon (Parasqualidus maii) is known only from the Vu Gia River, where it is threatened by hydropower developments. The Ben Hai River (Song Ben Hai in Vietnamese) is located in central coastal Vietnam (Quang Tri and Quang Binh provinces). The Vietnamese cardinal minnow (Tanichthys micagemmae) is confined to the Ben Hai River. The Kien Giang River (Song Kien Giang in Vietnamese) is located in central coastal Vietnam (Quang Binh province). The Kien Giang carp (Cyprinus melanes) is confined to the Kien Giang River. The Vinh Thanh River (Song Vinh Thanh in Vietnamese) is located in central Vietnam (Binh Dinh province). The marbled loach (Sewellia marmorata) and the Vinh Thanh

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stone loach (Schistura thanho) are both known only from the Vinh Thanh River drainage. The Thua Luu River (Song Thua Luu in Vietnamese) is located in central Vietnam. The Bach Ma stone loach (Schistura bachmaensis) is known only from a single small stream within the Thua Luu River drainage. Valenciennes’ stone loach (S. spiloptera) is known only from the Thua Luu River drainage, where it too now appears to be confined to a single stream. Both are seriously threatened by loss of habitat. The Thuy Loan River (Song Thuy Loan in Vietnamese) is located on the eastern face of the Annamite Mountains in central Vietnam (Da Nang province). The Thuy Loan catfish (Pseudecheneis maurus) is known only from the Thuy Loan River.

Coasts and Satellite Islands The section includes the coastal areas of South and South East Asia with the exception of the Malay Peninsula, but including Sri Lanka, Hainan, and numerous smaller islands. The black-eared flying fox (Pteropus melanotus) is, as a species, found across a number of smaller islands in the Bay of Bengal and the eastern Indian Ocean. The nominate form (P. m. melanotus) is found in the Andaman and Nicobar islands, where it is threatened by loss of habitat and hunting. The Porbandar house gecko (Hemidactylus porbandarensis) is confined to the port area of the city of Porbandar in western India (Gujarat). The five-fingered cylindrical skink (Chalcides pentadactylus) is known only from a single specimen (since lost) collected in 1870 from the sandy banks of a tidal river in south-western India (Kerala). The Madras spotted skink (Barkudia insularis) is known only from two mangrove localities in north-eastern India (Odisha).

Sri Lanka Sri Lanka is a large island located off the south-eastern coast of India. A land bridge once existed between the two that was said to have been passable on foot up to the late fifteenth century, although today it consists only of a chain of limestone shoals slightly above sea level in places. The island itself consists mainly of flat to rolling coastal plains, although with high mountains rising in the south-central part. The latter is responsible for the two main climatic divisions of the island, a very humid one in the south-west and a very dry one in the north-east. The south-western monsoon sweeps the south-west with moisture and nourishes it with a high precipitation, but almost all the rainfall is released there because the oceanic winds are stopped by the high mountain barrier. More than 2500 years of cultivation have deprived Sri Lanka of almost all its virgin vegetation. However, much of it was still covered by more or less dense forests up to the 1850s. During the last 150 years these forests have been cut down to make way for plantations. This threatens many of Sri Lanka’s animals with

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extinction, although the establishment of national parks and reserves has saved some of them, at least temporarily. That said, what remains of the biodiversity of this island ranks among the largest and most important in the world. The Sri Lankan elephant (Elephas maximus maximus) is one of the three currently recognized subspecies of Asian elephant. Historically it was found throughout the island from sea level to the highest montane forests. Until 1830 they were so plentiful that their destruction was actually encouraged by the colonial government, with rewards paid for any that were killed. Many bull elephants were also ruthlessly hunted for sport. A lone British officer was credited with shooting over 1500, and two others are reputed to have shot half that number each. Many other trophy hunters claimed 250–300 during this period. It is estimated that about 10,000 elephants in all were killed or captured between 1831 and 1900. Yet by the turn of the twentieth century the animals were still widely distributed and relatively abundant in both the wet and dry lowlands. With a rapidly growing human population and resulting development, however, these large herbivores became increasingly restricted to fragmented areas of the lowland dry zone in the north, east and south-east, where by 1970 no more than 2500 still survived. With better protection and management, however, numbers began to grow once more. Unfortunately, the network of national parks where many now live are too small and provide insufficient habitat during the dry period, and human–elephant conflict outside the confines of these protected areas has increased considerably, to the detriment of the latter. In addition, during the recent civil war many of the animals were maimed or killed by poaching and land mines. Today an effort is being made to ameliorate conflicts and to preserve as many viable populations as possible, in as wide a range of suitable habitats as is feasible, both inside and outside protected areas. The Sri Lankan sloth bear (Melursus ursinus inornatus) historically occurred throughout the island, but is now confined to the remaining lowland dry forests of the north and east. There are no reliable population figures available, but the animals are considered to be threatened. The Sri Lankan leopard (Panthera pardus kotiya) is still fairly widespread on the island, but everywhere threatened by habitat destruction and fragmentation as well as by poaching. The toque macaque (Macaca sinica) is found throughout most of Sri Lanka where it is divided into three subspecies that will be discussed below. All are threatened by habitat destruction, persecution and collection for use as pets. The purple-faced langur (Semnopithecus vetulus) is found throughout much of Sri Lanka, where it is divided into four subspecies. Philbrick’s purple-faced langur (S. v. philbricki) is found in northern and eastern Sri Lanka, where it is threatened by loss of habitat. The red slender loris (Loris tardigradus) is a type of nocturnal prosimian confined to the rainforests of Sri Lanka, where it is divided into two subspecies which will be discussed below. The grey slender loris (L. lydekkerianus), previously

The Oriental Region

discussed, also has at least two subspecies endemic to Sri Lanka. The foothill grey slender loris (L. l. grandis) is known only from the East Matale Hills, but probably occurs throughout the lower foothills of central Sri Lanka (Central province). All are threatened by loss of habitat and other factors. Layard’s palm squirrel (Funambulus layardi) and the dusky palm squirrel (F. obscurus) are both found patchily in central and south-western Sri Lanka, where they are threatened by loss of habitat. The Ohiya rat (Srilankamys ohiensis) is found patchily in central and south-western Sri Lanka. Fernandon’s spiny mouse (Mus fernandoni) is known only from a few localities scattered throughout Sri Lanka. Mayor’s mouse (M. mayori) is known only from a few localities in central and south-western Sri Lanka. Both are threatened by loss of habitat, pesticides and introduced domestic predators. The Sri Lankan long-tailed shrew (Crocidura miya) is known only from a few localities in central and south-western Sri Lanka (Central and Subaragamuwa provinces). The jungle pygmy white-toothed shrew (Suncus zeylanicus) is known only from a few localities in central and southwestern Sri Lanka (Central, Subaragamuwa and Southern provinces). The red-faced malkoha (Phaenicophaeus pyrrhocephalus) is a large cuckoo known to have been widespread on the island in the late nineteenth century. With the loss of much of its preferred moist forest habitat it is now largely confined to the Sinharaja Forest Reserve and surrounding areas in southwestern Sri Lanka, where it numbers no more than a few thousand and perhaps as low as several hundred. The Sri Lankan wood pigeon (Columba torringtoniae) is found patchily in central and south-western Sri Lanka (Central, Subaragamuwa, Uva, and Southern provinces). The Sri Lankan blue magpie (Urocissa ornata) is confined to central and south-western Sri Lanka, where it is threatened by loss of habitat. The white-faced starling (Sturnornis albofrontatus) is confined to central and south-western Sri Lanka. Kelaart’s toad (Adenomus kelaartii) is found patchily in central and south-western Sri Lanka (Central, Subaragamuwa, Uva, and Southern provinces). Kotagama’s dwarf toad (Duttaphrynus kotagamai) is known only from three scattered localities in south-western Sri Lanka (Kitulgala, Messana, and Sinharaja forests). The Rumassala shrub frog (Pseudophilautus zimmeri) is known only from a single specimen collected in 1927 from south-western coastal Sri Lanka (Southern province). All habitats in the region have since been destroyed by urbanization, and the species is almost certainly extinct. The Sinharaja Forest shrub frog (P. silvaticus) is confined to a small area of south-western Sri Lanka (Subaragamuwa province), where it is threatened by loss of habitat. The long-snouted foam-nesting frog (Taruga longinasus) is known only from a few scattered localities in central and

south-western Sri Lanka (Central, Subaragamuwa, and Southern provinces). Karunaratne’s rice frog (Microhyla karunaratnei) is known only from two localities in southern Sri Lanka (Subaragamuwa province). The Pattipola caecilian (Ichthyophis orthoplicatus) is confined to a small area of south-central Sri Lanka (Uva province). The pseudo-angular caecilian (Ichthyophis pseudangularis) is found patchily in central and south-western Sri Lanka (Central, Subaragamuwa, and Southern provinces). The Central Highlands The Central Highlands are located in the south-central interior of Sri Lanka. During the first half of the nineteenth century most of the montane rainforests were cleared for the largescale planting of coffee and tea, resulting in the extinction of a great many species. What remains is still a biological hotspot of global importance, and is protected within three main protected areas (the Peak Wilderness Sanctuary, Horton Plains National Park, and Knuckles Conservation Forest). The Central Highlands toque macaque (Macaca sinica opisthomelas) is confined to the Central Highlands. The Central Highlands purple-faced langur (S. v. monticola) is confined to the Central Highlands. The Horton Plains red slender loris (Loris tardigradus nycticeboides) is confined to a small area of the Central Highlands. The Central Highlands rat (Rattus montanus) is confined to four localities within the Central Highlands (Central and Subaragamuwa provinces). The Central Highlands tree mouse (Vandeleuria nolthenii) is confined to a few fragmented localities in the Central Highlands (Central and Uva provinces). Fellowes-Gordon’s pygmy white-toothed shrew (Suncus fellowesgordoni) is known only from a few localities within the Central Highlands (Central province). Pearson’s long-clawed shrew (Solisorex pearsoni) is known only from a few localities within the Central Highlands (Central province). The Sri Lankan whistling thrush (Myophonus blighi) is a rare species confined to the Central Highlands (Central, Uva and Subaragamuwa provinces). The spineless forest lizard (Calotes liocephalus) is known only from an undefined area of central Sri Lanka. The Deignan tree skink (Lankascincus deignani) is confined to montane rainforest within the Central Highlands, where it is threatened by loss of habitat. Cope’s rough-sided snake (Aspidura copei) is known only from a few specimens collected from the Central Highlands. The Kandy dwarf toad (Adenomus kandianus) was long known only from its original collection in 1872 from an undefined locality in Sri Lanka. It was rediscovered in 2009 within the Peak Wilderness Sanctuary of the Central Highlands, and a second population was located in 2014 from the Pidurutalagala Forest Reserve.

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The Rawkana foam-nesting frog (Taruga fastigo) is known only from a single montane forest fragment in south-western Sri Lanka (Subaragamuwa province). Günther’s foam-nesting frog (T. eques) is confined to a few localities in central and south-western Sri Lanka (Central, Uva and Subaragamuwa provinces). The shrub frogs (Pseudophilautus) are a large genus from the Western Ghats of south-western India and Sri Lanka. Sadly, a great many species from the latter became extinct before anything could be learned about them, although it is logical to suppose that many lived in the Central Highlands. These include Günther’s shrub frog (P. adspersus), the Dimbulla shrub frog (P. dimbullae), Shreve’s shrub frog (P. eximius), the blunt-snouted shrub frog (P. extirpo), the Pattipola shrub frog (P. halyi), Malcolm Smith’s shrub frog (P. malcolmsmithi), the white-nosed shrub frog (P. leucorhinus), the sharp-snout shrub frog (P. nasutus), the pygmy shrub frog (P. nanus), the white-blotched shrub frog (P. zal), the temporal shrub frog (P. temporalis), the variable shrub frog (P. variabilis), the leopard shrub frog (P. pardus), the Farnland shrub frog (P. rugatus), the Maia shrub frog (P. maia), and the sharp-snouted shrub frog (P. oxyrhynchus). The starry shrub frog (P. stellatus) was thought to be extinct for 156 years, until unexpectedly rediscovered in the Peak Wilderness Sanctuary in 2009. Other still-extant threatened species include the golden-eyed shrub frog (P. ocularis), the bigfoot shrub frog (P. macropus), the elegant shrub frog (P. decoris), the Morningside shrub frog (P. simba), the cheeky shrub frog (P. procax), the papillated shrub frog (P. papillosus), Asanka’s shrub frog (P. asankai), the golden shrub frog (P. auratus), the blue-thighed shrub frog (P. caeruleus), the hollow-snouted shrub frog (P. cavirostris), the round-snout shrub frog (P. femoralis), Frankenberg’s shrub frog (P. frankenbergi), the small-eared shrub frog (P. microtympanum), Halliday’s shrub frog (P. hallidayi), the Handapan Ella shrub frog (P. lunatus), Annandale’s shrub frog (P. semiruber), Steiner’s shrub frog (P. steineri), the pug-nosed shrub frog (P. silus), Schmarda’s shrub frog (P. schmarda), Sarasinors’ shrub frog (P. sarasinorum), Poppy’s shrub frog (P. poppiae), the webless shrub frog (P. hypomelas), and the Horton Plains shrub frog (P. alto). The dull-green bush frog (Raorchestes viridis) is known only from a small area of south-central Sri Lanka (Central province). Green’s frog (Zakerana greenii) is confined to a small area of south-central Sri Lanka (Central and Uva provinces). The Central Highlands balloon frog (Uperodon palmatus) is confined to a small area of south-central Sri Lanka (Central and Uva provinces). The Sri Lanka rice frog (Microhyla zeylanica) is confined to a small area of south-central Sri Lanka (Central and Uva provinces). The Knuckles Range –– The Knuckles Range (Dumbara Kanduvetiya in Sinhalese) is located in central Sri Lanka

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(Central province). It takes its name from a series of recumbent folds and peaks in the west of the massif, which resemble the knuckles of a clenched fist. Tennent’s leaf-nosed lizard (Ceratophora tennentii) appears to be confined to the Knuckles Range. The Knuckles pygmy lizard (Cophotis dumbara) is known only from the Knuckles Range, where it is seriously threatened by loss of habitat. Stuart’s shrub frog (Pseudophilautus stuarti) is known only from the Corbett’s Gap region in the southern Knuckles Range, and from nearby Bambarella Peak. Moores’ shrub frog (P. mooreorum) has been recorded from Corbett’s Gap, Hunnasgiriya and Bambarella Peak. Hoffmann’s shrub frog (P. hoffmanni) is known from the Corbett’s Gap region and Bambarella Peak, but may occur in other cloud forests of the Knuckles Range. The tawny shrub frog (P. fulvus) is confined to the northern Knuckles Range. All are threatened by logging and pesticides. The marbled streamlined frog (Nannophrys marmorata) is a largely aquatic species confined to hillstreams within the Knuckles Range. Lowland Rainforests Areas of lowland rainforest (i.e. below 1000 m elevation) are to be found in central and south-western Sri Lanka. During the colonial period from 1505 to 1948 this ‘wet zone’ was largely converted to agriculture, and became heavily settled. Today little remains outside of protected areas. The lowland wet zone toque macaque (Macaca sinica aurifrons) is confined to south-western Sri Lanka. The southern purple-faced langur (Semnopithecus vetulus vetulus) and the regal purple-faced langur (S. v. nestor) are both confined to the lowland wet zone. The wet zone red slender loris (Loris tardigradus tardigradus) is confined to south-western Sri Lanka. The Serendib scops owl (Otus thilohoffmanni) is known only from a few localities in south-western Sri Lanka (Western, Southern, and Subaragamuwa provinces). The ashy-headed laughingthrush (Garrulax cinereifrons) is found patchily in south-western Sri Lanka. The green-billed coucal (Centropus chlororhynchos) is a type of cuckoo found patchily in south-western Sri Lanka (Central, Western, Southern, and Subaragamuwa provinces). The rough-nose horned lizard (Ceratophora aspera) is found patchily in south-western Sri Lanka (Central, Western, Subaragamuwa and Southern provinces). Ranawane’s cat snake (Boiga ranawanei) is known only from two localities in central Sri Lanka (Central province). Noellert’s toad (Duttaphrynus noellerti) is known only from a few scattered localities in central and south-western Sri Lanka (Central, Subaragamuwa and Southern provinces). Several species of shrub frog (Pseudophilautus) are threatened by loss of habitat and agrochemical pollution. The whistling shrub frog (P. nemus) is confined to the Haycock Hill (Hiniduma) Forest Reserve in south-western Sri Lanka

The Oriental Region

(Southern province). The Gannoruwa shrub frog (P. zorro) is known only from a small area of central Sri Lanka (Central province). The side-striped shrub frog (P. pleurotaenia) was long known only from now-lost type specimens, but has since been found in two disjunct localities in central and southwestern Sri Lanka (Central province). The slender shrub frog (P. tanu) is known only from the Beraliya and Kanneliya forest reserves in south-western Sri Lanka (Southern province). Mittermeier’s shrub frog (P. mittermeieri) is known only from two disjunct localities in south-western Sri Lanka (Southern province). The reticulated shrub frog (P. reticulatus) is known from a few scattered localities in central and south-western Sri Lanka (Central, Western, Subaragamuwa, and Southern provinces). The short-horned shrub frog (P. singu) and the leafdwelling shrub frog (P. folicola) are both known only from a few widely scattered localities in south-western Sri Lanka (Subaragamuwa and Southern provinces). The sharp-snouted shrub frog (P. cuspis) is known only from a small area of south-western Sri Lanka (Subaragamuwa and Southern provinces). Nagao’s balloon frog (Uperodon nagaoi) is known only from a few specimens collected in the Kanneliya Forest Reserve and Hiyare Forest Reserve in south-western Sri Lanka (South province). The Sinharaja Forest –– The Sinharaja Forest is located in a hilly region of south-western Sri Lanka (Southern province). The country’s last viable area of primary rainforest, it was saved from commercial logging by its inaccessibility and is now nominally protected within the Sinharaja Forest Reserve. In 1988 it was declared a UNESCO World Heritage Site. The Sinharaja shrew (Crocidura hikmiya) is known only from two localities within the Sinharaja Forest Reserve. Lowland Dry Forests Lowland dry forests are to be found across most of Sri Lanka with the exception of the south-western ‘wet zone’, the Central Highlands, and the thorn scrubs of the northern Jaffna Peninsula. The dry zone toque macaque (Macaca sinica sinica) is confined to northern and eastern Sri Lanka. The southern sacred langur (Semnopithecus thersites) is confined to Sri Lanka’s dry-zone. The dry-zone grey slender loris (Loris lydekkerianus nordicus) is found throughout north-central and eastern Sri Lanka, possibly extending into the south-east.

Lakes, Rivers, and Marshes Sri Lanka has over 100 rivers draining from the Central Highlands, giving rise to a number of spectacular waterfalls. The largely aquatic streamlined frogs (Nannophrys) are endemic to Sri Lanka, where they are threatened by loss of habitat and pollution. Günther’s streamlined frog (N. guentheri) is known only from its original collection from an undefined locality in Sri Lanka. It is now considered to be

extinct. Naeyaka’s streamlined frog (N. naeyakai) is known only from a few areas of east-central Sri Lanka. The Ceylon streamlined frog (N. ceylonensis) is confined to south-western Sri Lanka. The Asoka barb (Systomus asoka) is largely confined to the upper reaches of the Sitawaka River and its tributaries in south-western Sri Lanka, with an additional population within the Kelani River drainage. It is seriously threatened by habitat destruction. The Wilpita rasbora (Rasbora wilpita) is confined to a handful of shallow streams within lowland rainforest areas of south-western Sri Lanka. The redfin labeo (Labeo lankae) is confined to the Malvathu, Kala Oya, and Perararu river drainages of central and north-western Sri Lanka. The lipstick goby (Sicyopus jonklaasi) is confined to a few fast-flowing streams within lowland rainforest areas of southwestern Sri Lanka, where it is threatened by loss of habitat and agrochemical pollution. Jonklaas’ loach (Lepidocephalichthys jonklaasi) is confined to a handful of shallow streams within lowland rainforest areas of south-western Sri Lanka. The Mahaweli River –– The Mahaweli River rise in Horton Plains National Park in central Sri Lanka and flows northeast. At 335 km in length it is the longest river in the country. Martenstyn’s barb (Systomus martenstyni) is confined to a few localities in the mid-Mahaweli River drainage. Fischer’s labeo (Labeo fisheri) is known only from a few streams in the Mahaveli River drainage, where it may already be extinct. The Kelani River –– The Kelani River is located in southwestern Sri Lanka. The Bandula barb (Pethia bandula) was long confined to a single small tributary of the Kelani River, although in recent years two other populations have been established. The Nilwala River –– The Nilwala River is located in southwestern Sri Lanka (Southern province). It rises in the Sinharaja Forest Reserve and flows south. The barred danio (Devario pathirana) is confined to the Nilwala River drainage, where it is threatened by pollution and overcollection for the international aquarium trade.

The Maldives The Maldives consist of almost 1200 coral islands grouped in a double chain of 26 low-lying atolls located south-west of India. None are more than 2.4 m above sea level, and are thus at high risk of being submerged by rising seas due to climate change. Addu Atoll Addu Atoll (also known as Seenu Atoll), located just south of the equator, is the southernmost of the Maldives. The Maldives flying fox (Pteropus hypomelanus maris) is known only from Addu Atoll.

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The Indo-Malaysian Realm

The Andaman Islands

The Nicobar Islands

The Andamans are an archipelago of 235 islands located in the Bay of Bengal between India and Myanmar. The Andaman rat (Rattus stoicus) is known only from Henry Lawrence Island, South Andaman, and Middle Andaman. Tytler’s black-eared flying fox (Pteropus melanotus tytleri) is confined to South Andaman and Rutland Island. The Andaman horseshoe bat (Rhinolophus cognatus) is fairly widespread within the Andaman Islands, being known from North Andaman, Interview Island, Baratang, Little Andaman, and a number of smaller islands. It is threatened mainly by human disturbance of its roosting caves. The Andaman teal (Anas albogularis) is a type of duck confined to the Andaman Islands and Great Coco Island. The Andaman krait (Bungarus andamanensis) is a type of venomous snake known for certain only from South Andaman Island, but is likely to occur on Middle and North Andaman as well. Charles Darwin’s oriental frog (Ingerana charlesdarwini) is known only from a few localities on South Andaman, North Andaman, and Long Island. The Andaman pipefish (Microphis insularis) is confined to rivers and streams within the Andaman Islands, where it is threatened by habitat destruction and degradation.

The Nicobar Islands are an isolated archipelago located in the eastern Indian Ocean off the northern tip of Sumatra. Unlike the Andaman Islands to the north they were never connected by a land bridge to the mainland, and are still largely covered by rainforests. The Nicobar long-tailed macaque (Macaca fascicularis umbrosus) is confined to Great Nicobar, Little Nicobar, and Katchall Island, where it is threatened mainly by hunting and trapping. The Nicobar tree shrew (Tupaia nicobarica) is confined to Great Nicobar and Little Nicobar. The Nicobar rat (Rattus burrus) is confined to Great Nicobar, Little Nicobar, and Trinkat Island. The palm rat (Rattus palmarum) is confined to Car Nicobar and Great Nicobar. The Nicobar flying fox (Pteropus faunulus) is currently known from six of the Nicobar Islands (Nancowry, Teressa, Camorta, Bompoka, Katchall, and Trinkat). It historically occurred on Car Nicobar as well, but appears to have been extirpated from there. The Nicobar sparrowhawk (Accipiter butleri) is known only from Great Nicobar, Little Nicobar, Pilo Milo, Kamorta, Teressa, Bompoka, Tillanchong, Katchall, and Nancowry islands. The Nicobar scops owl (Otus alius) was originally known only from two specimens collected on Great Nicobar in 1998, and another trapped and photographed in 2003 on Nancowry Island. In recent years it has been seen more frequently and may occur on other islands in the southern Nicobar group, in particular Little Nicobar. The Nicobar scrubfowl (Megapodius nicobariensis) is a chicken-like bird divided into two subspecies. The nominate form (M. n. nicobariensis) is found on Camorta, Trinkat, Nancowry, Katchall, Teressa, Bompoka, and Tillanchong islands. Abbott’s Nicobar scrubfowl (M. n. abbotti) is confined to Great Nicobar, Little Nicobar, Kondul, Menchal, Treis, and Meroe. Both were considerably affected by the 2004 tsunami but have recovered, although loss of habitat continues to be a threat. The Nicobar cat snake (Boiga wallachi) is confined to Great Nicobar and Little Nicobar. The Nicobar frog (Minervarya nicobariensis) is confined to the central and northern Nicobar Islands (Car Nicobar, Teressa, Camorta, Trinkat, and Nancowry).

North Andaman and Middle Andaman North Andaman and Middle Andaman are separated only by a narrow strait. The Middle Andaman shrew (Crocidura hispida) is known only from a single specimen collected on Middle Andaman. The Middle Andaman rice frog (Microhyla chakrapani) is known only from its original collection on Middle Andaman Island in the 1970s. South Andaman South Andaman is the southernmost and most populous of the Andaman Islands. The South Andaman shrew (Crocidura andamanensis) was long known only from a single specimen collected on South Andaman is 1902. A second has since been found on Mount Harriet. Narcondam Narcondam is a small, volcanic island located north-east of the main chain of the Andamans. It is still largely forested. The Narcondam hornbill (Rhyticeros narcondami) is confined to Narcondam, where it has a small but stable population of around 300. The Coco Islands The Coco Islands are a small group of islands located north of the Andaman Islands. Oates’ blind snake (Argyrophis oatesii) is known only from tiny, uninhabited Table Island in the Coco Islands.

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Great Nicobar Great Nicobar is the southernmost and largest of the Nicobar Islands. The vast majority of it has been set aside as the Great Nicobar Biosphere Reserve. The Great Nicobar shrew (Crocidura nicobarica) is known only from the southern tip of Great Nicobar. The Great Nicobar tree frog (Polypedates insularis) is confined to Great Nicobar.

Hainan The island of Hainan is located in the South China Sea and is separated from the Chinese mainland to the north by a shallow, narrow strait. A relatively stable climate over a long period of time

The Oriental Region

has led to the development of a very diverse flora and fauna, but habitat destruction and excessive hunting combined with a high population density have taken their toll. Almost no large animals or lowland forests remain, and the island’s lakes and rivers are threatened by introduced carp and catfish. The Hainan crested gibbon (Nomascus hainanus) is one of the world’s most endangered primates. Prior to the 1960s it was widely distributed on the island, but has since been confined to the Bawangling Nature Reserve in western Hainan. In 1993 there were only three groups remaining in remnant forest with less than 20 individuals in total. By 2003 only 13 were left. Since then, the population has increased somewhat but is still thought to be no more than 20. Sadly, some hunting pressure remains. The Hainan sambar deer (Rusa unicolor hainana) is confined to Hainan. The Hainan hare (Lepus hainanus) currently survives only in two artificially deforested lands in western Hainan used for deer ranching, where it is still heavily poached for its meat and skin. It also faces competition from introduced feral rabbits. The Hainan lesser marmoset rat (Hapalomys delacouri marmosa) is confined to Hainan. The Hainan peacock-pheasant (Polyplectron katsumatae) is found patchily in central and southern Hainan, where it has undergone significant declines due to habitat destruction and hunting. The Hainan partridge (Arborophila ardens) is found patchily in central and southern Hainan, where it has undergone significant declines due to habitat destruction and hunting. The Hainan magpie (Urocissa whiteheadi) is confined to the remaining forests of Hainan, where it is threatened by hunting. The Hainan leaf warbler (Phylloscopus hainanus) is a littleknown species known only from a few localities in central and southern Hainan. The Hainan odd-scaled snake (Achalinus hainanensis) is known only from two specimens collected on Mount Jianfengling in south-western Hainan. The Hainan toad (Parapelophryne scalpta) is confined to central and southern Hainan. The Hainan gliding tree frog (Rhacophorus yinggelingensis) is confined to the montane forests of the Yinggeling National Nature Reserve in central Hainan, where the total population is estimated to be around 3000. The red-headed gliding frog (Buergeria oxycephala) is confined to central and southern Hainan. The ocellated shrub frog (Liuixalus ocellatus) is known only from a few localities in central and southern Hainan. The Hainan large-eyed litter frog (Leptobrachium hainanense) is known only from a few localities in central and southern Hainan. The fine-spined frog (Sylvirana spinulosa) is known only from a few localities in central and southern Hainan. The Hainan wart frog (Limnonectes fragilis) is known only from a few localities in central and southern Hainan. The Diaoluo shrub frog (Liuixalus hainanus) is known from a few localities in central and western Hainan.

Boulenger’s cascade frog (Amolops hainanensis) and the little cascade frog (A. torrentis) are both confined to central and south-western Hainan, where they are threatened by loss of habitat and overharvesting for food. The Hainan knobby newt (Tylototriton hainanensis) is known only from three localities in central and south-western Hainan, all of which fortunately lie within protected areas.

Islands of Rach Gia Bay Rach Gia Bay is located off southern Vietnam. Its 92 small, rocky islands are protected as part of the Kien Giang Biosphere Reserve. Hon Son Hon Son is composed of large granite boulders and has virtually no remaining vegetation. The Hon Son pit viper (Cryptelytrops honsonensis) is confined to the island. Hon Khoai Hon Khoai is one of the largest islands in Rach Gia Bay. The psychedelic day gecko (Cnemaspis psychedelica) is confined to Hon Khoai and Hon Tuong, the latter a small islet off Hon Khoai.

Miscellaneous Islands Con Son is the largest island of the Con Dao Archipelago, off the coast of southern Vietnam. It has been designated a National Park. The Con Son long-tailed macaque (Macaca fascicularis condorensis) is confined to Con Son Island. Phu Quoc is located off southern Vietnam in the Gulf of Thailand. The Phu Quoc shrew (Crocidura phuquocensis) is confined to the island’s undisturbed lowland rainforests. Phangan Island (Ko Phangan in Thai) is located in the Gulf of Thailand. The Phangan spectacled langur (Trachypithecus obscurus seimundi) is confined to Phangan Island. Khram Island (Ko Khrom in Thai) is an uninhabited island located in Bangkok Bay, Thailand. The dark-crowned longtailed macaque (Macaca fascicularis atriceps) is confined to Ko Khram Island.

Balance for the Oriental Region The Oriental Region was already inhabited by Homo erectus approximately 1.5 million years ago. Modern humans (H. sapiens) first arrived on the coastal areas of the Indian subcontinent from Africa by 55,000 years ago, and throughout mainland South East Asia by around 60,000 years ago. Evidence for the domestication of food crops and animals and the construction of permanent settlements there date from around 6500 B . C .. The latter gradually developed into the Indus Valley Civilization, the first urban culture in South Asia, which flourished during 2500–1900 B . C . in what is now Pakistan and western India. Others would follow, gradually expanding east and south. South Asia’s subsequent history would be an inevitable progression of ever-rising human population and

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The Indo-Malaysian Realm

environmental destruction. Much of the latter has been driven there by the presence of entirely unproductive domestic cattle, which Hindu religious precepts prohibit the killing of. European exploration of the Oriental Region was dominated by the Portuguese. In the early 1490s Pêro da Covilhã visited the Malabar Coast of India, and would make another journey there in 1497–98. In 1506 Lourenço de Almeida reached the Maldives and Sri Lanka, and in 1511 Duarte Fernandes led a diplomatic mission to Siam (present-day Thailand). That same year Rui Nunes da Cunha undertook a similar mission to what is now Myanmar, and in 1516 Portuguese traders landed in what is now Vietnam, naming it Cochinchina. The British, French and Portuguese would subsequently proceed to set up trading posts along the South and South East Asian coast, before ultimately colonizing the mainland areas well into the twentieth century. Hunting has since taken a particularly heavy toll on wild animals in India, particularly during the last two and a half centuries when the whole country was used as a happy hunting ground by both Asians and Europeans. There has also been a massive conversion of habitat for use as agriculture, although between the between the two World Wars the situation for India’s wildlife at least became a little improved by the establishment of national parks and reserves. It was not until the end of the colonial period, during the second half of the twentieth century, that environmental conditions in this region truly began to deteriorate. Indeed, after the end of World War II a massacre of wild animals and a destruction of their habitats, particularly in

lowland areas, began on an unprecedented scale. Forest clearance and a largely uncontrolled trade in wild animals for their meat or use in ‘traditional medicine’ is still going on. Diseases contracted from domestic cattle are a particular threat to Asia’s wild ungulates, and several species of vulture have suffered catastrophic declines due to feeding on animal carcasses treated with the veterinary drug diclofenac. Attempts at banning the drug have thus far been unsuccessful. By removing all carcasses, the birds helped to decrease pollution, disease spread, and suppressed undesirable mammalian scavengers. In their absence, the population of feral dogs and rats, along with their zoonotic diseases, has increased considerably. In recent historical time (i.e. since A . D . 1500), the Oriental Region has lost at least 23 species/2 subspecies of vertebrates. Among the extinct forms 2 species/2 subspecies are mammals, 1 species is a bird, 18 species are amphibians, and 2 species are freshwater fishes. Another 13 species/1 subspecies are possibly extinct, and 1 species is currently extinct in the wild. In addition, there are 1002 species/51 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 103 species/38 subspecies are mammals, 62 species/9 subspecies are birds, 158 species/4 are reptiles, 244 species are amphibians, and 435 species are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

2 species

2 species

~ species

103 species

2 subspecies

1 subspecies

~ subspecies

38 subspecies

4 taxa

3 taxa

~ taxa

141 taxa

1 species

1 species

~ species

62 species

~ subspecies

~ subspecies

~ subspecies

9 subspecies

1 taxon

1 taxon

~ taxa

71 taxa

~ species

1 species

1 species

158 species

~ subspecies

~ subspecies

~ subspecies

4 subspecies

~ taxa

1 taxon

1 taxon

162 taxa

18 species

~ species

~ species

244 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

18 taxa

~ taxa

~ taxa

244 taxa

2 species

9 species

~ species

435 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

2 taxa

9 taxa

~ taxa

435 taxa

23 species

13 species

1 species

1002 species

2 subspecies

1 subspecies

~ subspecies

51 subspecies

25 taxa

14 taxa

1 taxon

1053 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

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The Sundaic Region

The Sundaic Region The Sundaic Region, as here defined, includes all of the Malay Peninsula; the Greater Sunda Islands of Borneo, Sumatra, and Java; and numerous smaller islands. It corresponds to a larger ancient landmass (known as Sundaland) that was exposed at various times throughout the last 2.6 million years during periods when sea levels were lower. Before sea levels rose again at the end of the last Ice Age Borneo had been part of the mainland of Asia, forming, with Java and Sumatra, the upland regions of a peninsula that extended east from present-day Indochina. The South China Sea and the Gulf of Thailand now submerge the former low-lying areas of the peninsula. Deeper waters separating Borneo from neighbouring Sulawesi prevented a land connection, creating, as previously noted, the divide known as Wallace’s Line between Asian, Australian and Papua-Melanesian zoogeographic regions. The climate of this region is, of course, entirely tropical, with diverse habitats, large forests, and high mountains, and the fauna shares affinities with mainland Asia. The Sundaic lesser one-horned rhinoceros (Rhinoceros sondaicus sondaicus) was historically found on Sumatra and Java, but was exterminated everywhere on the former island during the 1940s except in Loeser National Park, where it hung on a few decades more before ultimately being wiped out. It may have represented a distinct species or subspecies, but we will likely never know. The surviving Javan population will be dealt with separately below. The western two-horned rhinoceros (Dicerorhinus sumatrensis sumatrensis) occurred in the Malay Peninsula and Sumatra. It is thought to have survived in very small numbers in the Tenasserim Mountains up to perhaps the twenty-first century, but is now certainly extirpated from the Asian mainland. The subspecies persists in Sumatra, however, where it will be discussed in greater detail. The Malayan tapir (Tapirus indicus) is the largest of the living tapir species and the only Asian representative. Once found throughout the tropical rainforests of Cambodia, southern and central Sumatra, Laos, Malaysia, Myanmar, Thailand and Vietnam, at all elevations, it is now largely restricted to the Malay Peninsula (parts of Myanmar, Thailand and Malaysia) and to Sumatra, where populations have been drastically reduced by hunting and loss of habitat. The siamang (Symphalangus syndactylus) is a large black gibbon confined to lowland and montane rainforests in southern peninsular Malaysia and the island of Sumatra, where it is threatened by loss of habitat. The agile gibbon (Hylobates agilis) is found in both lowland and montane forests in southern peninsular Malaysia and Sumatra. Geoffroy’s white-handed gibbon (H. lar entelloides) is found in southern Myanmar and south-western Thailand. Both are threatened by habitat destruction and overcollection for the illegal pet trade. The southern pig-tailed macaque (Macaca nemestrina) is found in southern peninsular Malaysia, Sumatra, Borneo, and

Bangka, where it is threatened by loss of habitat, hunting for food, and collection for use as pets. The Sunda slow loris (Nycticebus coucang) is a small, nocturnal prosimian found in the southern Malay Peninsula, central and southern Sumatra, the Riau Archipelago, the Natuna Islands, and Singapore, where it is threatened mainly by collection for the international pet trade. The southern serow (Capricornis sumatraensis) is a type of goat-antelope that lives in the mountains of Sumatra and peninsular Malaysia, where it has been much-hunted. The Sunda bearded pig (Sus barbatus) is divided into two subspecies, both of which are threatened by loss of habitat. The nominate form (S. b. barbatus) was historically found throughout Borneo and some of its nearshore islands, the Malay Peninsula, Singapore and the Riau Islands (although it has been extirpated from the latter islands, and much of its former range elsewhere). The Sunda clouded leopard (Neofelis diardi) is divided into two subspecies found, respectively, on Borneo and Sumatra, although fossil evidence indicates that the species formerly occurred on Java as well. They will be discussed below. The flat-headed cat (Prionailurus planiceps) is a small wild cat found patchily on the Malay Peninsula, Sumatra and Borneo. It is threatened by loss of its wetland habitat. The Sunda wild dog (Cuon alpinus sumatrensis) is largely confined to protected area on Sumatra and Java. The Sunda binturong (Arctictis binturong penicillata) is a large viverrid known from Sumatra and Java. The otter civet (Cynogale bennettii) is a semi-aquatic species found in the Malay Peninsula, Sumatra and Borneo, where it is threatened by loss of habitat and pollution. The smoky flying squirrel (Pteromyscus pulverulentus) is fairly widespread in the southern Malay Peninsula, Sumatra and Borneo, but has undergone significant population declines due to habitat destruction. The arrow flying squirrel (Petinomys sagitta) is known only from a single specimen collected on Java in 1766, although it may also occur on the island of Bangka. Hagen’s flying squirrel (P. hageni) is known only from a few specimens collected in northern Sumatra and western Borneo, but is almost certainly more widespread. Vordermann’s flying squirrel (P. vordermanni) is known from a few scattered localities on the Malay Peninsula, Borneo and the Riau Islands off eastern Sumatra. The whiskered flying squirrel (P. genibarbis) is found in Peninsular Malaysia, Sumatra, northern Borneo and eastern Java. All are presumably threatened by deforestation. The Niobe ground squirrel (Lariscus niobe) is known from the mountains of western Sumatra and eastern Java, where it has likely been impacted by loss of habitat. The Rajah spiny rat (Maxomys rajah) and Whitehead’s spiny rat (M. whiteheadi) are both found in the southern Malay Peninsula, Sumatra, and Borneo, where they are threatened by loss of habitat.

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The bare-backed rousette fruit bat (Rousettus spinalatus) is a cave-roosting species known only from northern Sumatra and northern Borneo (Sabah, Sarawak and Kalimantan Utara). Ridley’s leaf-nosed bat (Hipposideros ridleyi) is found patchily in peninsular Malaysia, Singapore and northern and western Borneo (Sarawak and Sabah). The orbiculus leafnosed bat (H. orbiculus) is known from a few localities in peninsular Malaysia and central Sumatra. Both are threatened by loss of habitat. Vordermann’s pipistrelle bat (Hypsugo vordermanni) is known from a few sporadic records originating in coastal Borneo, Banggi Island, and Belitung. Andersen’s horseshoe bat (Rhinolophus nereis) is known only from Siantan Island in the Anambas Islands and from the North Natuna Islands, in the South China Sea. The Krau woolly bat (Kerivoula krauensis) is found widely but patchily in the southern Malay Peninsula, Borneo, and Sumatra. The bronze tube-nosed bat (Murina aenea) is found in the Malay Peninsula and Borneo. The gilded tube-nosed bat (M. rozendaali) is found patchily in the Malay Peninsula, Borneo, and southern Sumatra. Both are threatened by loss of habitat. The false serotine bat (Hesperoptenus doriae) is known only from a few specimens collected in southern peninsular Malaysia and north-western Borneo (Sarawak). The northern free-tailed bat (Chaerephon johorensis) is known only from a few specimens collected in the southern Malay Peninsula and from Sumatra. Herman’s mouse-eared bat (Myotis hermani) is known only from three individuals collected from the Malay Peninsula and Sabang Island north of Sumatra. The Malayan tailless leaf-nosed bat (Coelops robinsoni) is a cave-roosting species known only from southern peninsular Malaysia and eastern Borneo. Storm’s stork (Ciconia stormi) is found patchily on the Malay Peninsula, Sumatra and Borneo, where it lives primarily in swamp forests. One of the rarest of the storks, the total population is thought to number less than 500 and continues to be threatened by deforestation and dam construction. Wallace’s hawk-eagle (Nisaetus nanus) remains widespread across the Malay Peninsula, Sumatra and Borneo, but is everywhere rare and declining due to loss of habitat. The helmeted hornbill (Rhinoplax vigil) is found widely but patchily in the Malay Peninsula, Sumatra and Borneo, where it is has become very rare due to severe hunting pressure and habitat destruction. The plain-pouched hornbill (Rhyticeros subruficollis) is found in southern Myanmar, western Thailand and the Malay Peninsula, where the population is small and declining due to hunting and loss of habitat. Vieillot’s crested fireback pheasant (Lophura rufa) occurs patchily in peninsular Malaysia, Sumatra and Bangka. The crestless fireback pheasant (L. erythrophthalma) is found in peninsular Malaysia, Singapore, Sumatra and Borneo. Both are threatened by habitat destruction and hunting.

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The black partridge (Melanoperdix niger) occurs patchily in lowland as well as lower montane areas of southern peninsular Malaysia, Sumatra and Borneo, where it is threatened by loss of habitat. The chestnut-necklaced partridge (Tropicoperdix charltonii) is divided into three subspecies, all of which are threatened by habitat destruction and collection for the cage-bird trade. The nominate form (T. c. charltonii) is found patchily in southern peninsular Malaysia and Thailand. The black and red coucal (Centropus nigrorufus) is a type of cuckoo confined to a few coastal areas on Java and perhaps Sumatra. The short-toed coucal (C. rectunguis) is found in the Malay Peninsula, Borneo and Sumatra. Both are threatened by habitat destruction and trapping. The Sunda nightjar (Caprimulgus concretus) is a rare species from Sumatra, Borneo and Belitung. The silvery pigeon (Columba argentina) appears always to have had a highly restricted range. It was recorded during the late nineteenth and early twentieth centuries from a few offshore islands in the Natuna Sea (west of Borneo) and west of Sumatra, and possibly on the adjacent mainland areas as well. Last recorded in 1931 and thought to be extinct, beginning in 2008 a small number of wild birds have been photographed within the Mentawai Islands. The reason for the species’ decline is thought to have been loss of habitat, hunting and the introduction of mammalian predators. The large green pigeon (Treron capellei) is found in the Malay Peninsula as well as on Sumatra and Borneo, but is everywhere threatened by loss of habitat. The southern pied starling (Gracupica jalla) is historically known from Java, Bali and eastern Sumatra (Lampung province), but appears to have been completely extirpated from the wild within the past few decades due to trapping for the illegal cage-bird trade. Large numbers are being bred in commercial bird farms in Java, although imports of a related species and the apparent interbreeding of the two make it unlikely that these might serve as a source for captive breeding. A population of genetically pure southern pied starlings may survive, however, within the Bali Bird Park. The ruby-throated bulbul (Rubigula dispar) is a passerine bird found on Sumatra, Java and Bali, where it is declining rapidly due to trapping for the international cage-bird trade. The hook-billed bulbul (Setornis criniger) is found in eastern Sumatra and Borneo, where it is threatened by loss of habitat. The greater green leafbird (Chloropsis sonnerati) is widespread in lowland rainforest areas of the Malay Peninsula, Borneo, Sumatra, Java, and Bali, but is everywhere threatened by overcollection for the cage-bird trade and loss of habitat. The Sunda white-eye (Zosterops flavus) is a passerine bird found patchily in the coastal areas of southern Borneo and northern Java, where it is seriously threatened by loss of habitat and overcollection for the cage-bird trade. The large-billed blue flycatcher (Cyornis caerulatus) is found patchily in lowland rainforest areas of Sumatra and Borneo.

The Sundaic Region

The false gharial (Tomistoma schlegelii) is a type of crocodilian found sporadically in swamp forest and wetland areas of peninsular Malaysia, Borneo, Sumatra and (possibly) far western Java, where it is threatened by loss of habitat. The Indochinese box turtle (Cuora amboinensis kamaroma) occurs in Vietnam, Laos, Cambodia, the Malay Peninsula and Borneo. The South Indonesian box turtle (C. a. couro) is found in Sumatra, Java, Bali and Sumbawa. Both are seriously threatened by overcollection for use as food and in ‘traditional medicine’, as well as for the international pet trade. The South East Asian narrow-headed softshell turtle (Chitra chitra) is divided into two subspecies. The nominate form (C. c. chitra) is found in Malaysia and Thailand, while the Sunda narrow-headed softshell turtle (C. c. javanensis) is confined to Java and Sumatra. Both are threatened by hunting and habitat destruction. The Sunda giant turtle (Orlitia borneensis) is found disjunctly in southern peninsular Malaysia, eastern Sumatra, and southern Borneo. It has been seriously depleted by overharvesting. The painted roofed turtle (Batagur borneoensis) is confined to the Malay Peninsula and Borneo. The Bengkulu cat snake (Boiga bengkuluensis) is known only from isolated records in southern peninsular Thailand and Sumatra. The spotted coral snake (Calliophis gracilis) is known only from a few historical collections in east-central Sumatra, the southern Malay Peninsula, Singapore, and Penang Island. The goldstripe rasbora (Rasbora chrysotaenia) is a popular aquarium fish that nevertheless has never been scientifically recorded in the wild state. It is believed to originate somewhere in the East Indies. The tricolour sharkminnow (Balantiocheilos melanopterus) is found widely but patchily in peninsular Malaysia, Sumatra and Borneo. The Sundaic knight goby (Stigmatogobius borneensis) is known only from a handful of specimens collected from brackish streams on Borneo. One was also collected from Singapore in 1878 although the species is believed to be extirpated there now, and there is a further (unverified) record from Sumatra. The smooth-thorn sheatfish (Ompok leiacanthus) is confined to peat swamps in Sumatra and Bangka. Bleeker’s pipefish (Doryichthys heterosoma) is known only from a few specimens collected from the Sambas and Lupar rivers of western Borneo and from the Natuna Islands.

The Malay Peninsula The Malay Peninsula (Semenanjung Tanah Melayu in Malay) runs almost directly north to south and, at its terminus, is the southernmost point on the Asian mainland. It includes southern Thailand, southern Myanmar, peninsular Malaysia and satellite islands such as Singapore.

The Malayan tiger (Panthera tigris jacksoni) was historically found in forested habitats throughout the Malay Peninsula, from montane areas to peatlands. It formerly occurred in Thailand and Singapore as well, but is now extinct there (the last wild tiger in Singapore was shot dead in 1930). The total estimated population declined from roughly 3000 in the 1950s to around 500 by the end of the century, and continues to fall. The latest estimate is around 250–340, mainly confined to protected areas of remote montane forest. Even there they continue to be threatened by poachers. The Malaysian white-handed gibbon (Hylobates lar lar) is confined to the Malay Peninsula. The banded surili (Presbytis femoralis) is a type of leafeating monkey divided into three subspecies, two of which are endemic to the Malay Peninsula. The Malaysian banded surili (P. f. femoralis) is confined to the southern Malay Peninsula and the island of Singapore. Robinson’s banded surili (P. f. robinsoni) is found in southern Thailand, southern Myanmar and Peninsular Malaysia. Fea’s muntjac (Muntiacus feae) is a little-known species from the Malay Peninsula (southern Myanmar and southern Thailand), where it is likely threatened by hunting and habitat destruction. The Malaysian tree rat (Pithecheir parvus) is confined to southern peninsular Malaysia (Pahang and Johore states). The Burmese pipistrelle bat (Hypsugo lophurus) is known only from its original collection in 1915 from southern peninsular Myanmar. The Selangor leaf-nosed bat (Hipposideros nequam) is only known from a single specimen collected during the nineteenth century in south-western peninsular Malaysia (Selangor state). The Chiku bent-toed gecko (Cyrtodactylus hidupselamanya) is confined to a single karst outcrop situated within a limestone formation in central peninsular Malaysia. The Larut Hills caecilian (Ichthyophis larutensis) is known from two nineteenth-century records and one recent one from the border region of southern peninsular Thailand and northwestern peninsular Malaysia. Supachai’s caecilian (I. supachaii) is known only from a small number of specimens collected in southern peninsular Thailand (Nakhon Si Thammarat and Trang provinces).

The Tenasserim Range The Tenasserim Range (Thio Khao Tanao Si in Thai/Banjaran Tanah Seri in Malay) is a roughly 1700-km chain of mountains and hills located in southern Myanmar, south-western Thailand and peninsular Malaysia. Tickell’s brown hornbill (Anorrhinus tickelli) is a rare species confined to the Tenasserim Mountains, where it is threatened by loss of habitat. The Chong Mountains The Chong Mountains (Khao Chong in Thai) are located in southern peninsular Thailand.

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The Chong stream toad (Ansonia siamensis) appears to be confined to the Chong Mountains. The Titiwangsa Range The Titiwangsa Range (Banjaran Titiwangsa in Malay/Khao Sankalakhiri in Thai) is the southernmost section of the Tenasserim Mountains. It runs some 480 km along the Malay Peninsula in southern Thailand and Malaysia. The mountain peacock-pheasant (Polyplectron inopinatum) is known for certain only from scattered localities within central peninsular Malaysia, but may also occur in extreme southern Thailand. Robinson’s whistling-thrush (Myiophonus robinsoni) is found patchily in peninsular Malaysia, where it is threatened by loss of habitat. Waterstradt’s bullfinch (Pyrrhula waterstradti) is a rare species from peninsular Malaysia (Pahang, Perak, and Selangor states). Jeet Sukumaran’s stream toad (Ansonia jeetsukumarani) is known only from a few localities in peninsular Malaysia (Pahang and Kelantan states). The Larut Hills rice frog (Microhyla annectens) has only been recorded with certainty from Bukit Larut, the Cameron Highlands, and the Genting Highlands of peninsular Malaysia. The Tanah Rata wart frog (Limnonectes nitidus) is known from a few localities in peninsular Malaysia (Pahang state). The Cameron Highlands (Tanah Tinggi Cameron in Malay) is a plateau located in peninsular Malaysia (Pahang, Perak and Selangor states). The Cameron Highlands white-bellied rat (Niviventer cameroni) is confined to the Cameron Highlands. The convex horseshoe bat (Rhinolophus convexus) is known only from a few specimens collected in the Cameron Highlands. The Cameron Highlands pit viper (Popeia nebularis) is known only from a single locality in the Cameroon Highlands, where it is threatened by illegal collection for the international exotic pet trade. The Genting Highlands (Tanah Tinggi Genting in Malay) is a plateau located in peninsular Malaysia (Pahang state). A large and expanding resort complex, the continued construction of hotels and other developments is having a serious impact on the environment. The precious stream toad (Ansonia smeagol) is confined to the Genting Highlands. Mount Jerai (Gunung Jerai in Malay, and formerly known as Kedah Peak) is located in peninsular Malaysia (Kedah state). The Jerai odorous frog (Odorrana monjerai) is confined to montane streams and their environs on Mount Jerai. Mount Tahan (Gunung Tahan in Malay) is located in central peninsular Malaysia (Pahang state), within Taman Negara National Park. Robinson’s sticky frog (Kalophrynus robinsoni) is known only from Mount Tahan, where it was last reported in 1922.

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Fraser’s Hill is located in peninsular Malaysia (Pahang state). The Fraser’s Hill day gecko (Cnemaspis flavolineata) is known only from three specimens collected from Fraser’s Hill.

Lowland Tropical Rainforests and Swamp Forests The once-extensive lowland rainforests within the Malay Peninsula have been largely replaced by oil palm plantations. The Selangor pygmy flying squirrel (Petaurillus kinlochii) is known only from a few specimens collected in south-western peninsular Malaysia. The social pipistrelle bat (Arielulus societatis) is known only from lowland forest fragments within southern peninsular Malaysia. The Malayan peacock-pheasant (Polyplectron malacense) is found in peninsular Malaysia and, at least historically, in southern peninsular Thailand as well. The species has been heavily impacted by hunting and habitat destruction, and is now largely confined to isolated forest patches within protected areas. The white-fronted scops owl (Otus sagittatus) is found patchily in lowland and foothill forests patches in the Malay Peninsula. Gurney’s pitta (Hydrornis gurneyi) is an extremely rare passerine bird now confined to a few remaining patches of lowland forest in peninsular Myanmar, having been extirpated from adjacent areas of Thailand where it was once common. Sworder’s bent-toed gecko (Cyrtodactylus sworderi) is confined to a few scattered localities in peninsular Malaysia (Johor and Pahang states), where it is threatened by loss of habitat. Groundwater’s keelback (Hebius groundwateri) is a type of snake confined to a small area of peninsular Thailand (Ranong and Chumphon provinces). Jintakune’s kukri snake (Oligodon jintakunei) is known only from a single specimen collected from southern peninsular Thailand (Krabi province). The banded green cat snake (Boiga saengsomi) is known only from a single locality in southern peninsular Thailand (Krabi province), where it is threatened by oil palm expansion. Shäfer’s spinejaw snake (Xenophidion schaeferi) is known only from a single specimen collected in south-western peninsular Malaysia (Selangor state). Klemmer’s blind snake (Malayotyphlops klemmeri) is known only from a single specimen collected near Kuala Lumpur in peninsular Malaysia (Selangor state). The Gollum toad (Ingerophrynus gollum) is known only from two small, disjunct localities in peninsular Malaysia (Johor and Selangor states), but presumably occurs more widely. The Kumquat toad (I. kumquat) is confined to peat swamps in south-western peninsular Malaysia, where it is threatened by habitat destruction and degradation. The Endau stream toad (Ansonia endauensis) is known only from Endau-Rompin National Park in peninsular Malaysia (Johor state). Grismer’s stream toad (A. latirostra)

The Sundaic Region

is known only from two widely separated localities in peninsular Malaysia (Pahang state). Latiff’s stream toad (A. latiffi) is confined to a few localities in peninsular Malaysia (Pahang and Terengganu states). All are threatened by loss of habitat. The ashy shrub frog (Philautus cinerascens) is known only from its original collection during the late nineteenth century from peninsular Myanmar. Kemp’s shrub frog (P. kempii) and the Kobo shrub frog (P. microdiscus) are both known only from their original collection during the early twentieth century from north-eastern India (Arunachal Pradesh). Kiew’s sticky frog (Kalophrynus palmatissimus) is confined to a small area of peninsular Malaysia (Pahang, Selangor, Negeri Sembilan, and Johor states). The minute balloon frog (Glyphoglossus minutus) is known only from two localities within Taman Negara National Park and a third in the Krau Wildlife Reserve in peninsular Malaysia (Pahang state). The Hala Bala dwarf litter frog (Leptobrachella solus) is known only from Hala Bala Wildlife Sanctuary in peninsular Thailand (Narathiwat province). It may occur more widely.

Lakes, Rivers, and Marshes The Malay Peninsula features a number of rivers draining either side of the central mountains, along with biologically important peat bogs. Two little-known species of water snake (Enhydris) are known only from southern peninsular Malaysia. The Pahang water snake (E. pahangensis) is known only from a single specimen collected in the 1940s. The Selangor water snake (E. indica) is known only from a few specimens collected from Selangor state and a further, rather questionable old record from Singapore. The Siamese perch (Badis siamensis) is confined to peninsular Thailand and the island of Phuket, where it is threatened by loss of habitat and overcollection for the international aquarium trade. Hale’s barb (Discherodontus halei) is known for certain only from the Pahang River of western peninsular Malaysia. Two other disjunct populations, in the Tapi River of southern Thailand and in the Chao Phraya drainage of north-central Thailand, are likely referable to a different species. The strawberry rasbora (Boraras naevus) is known from swamps in peninsular Thailand (Surat Thani province), and may be found more widely within the lower Tapi River drainage. Harvey’s gourami (Parosphromenus harveyi) is confined to peat swamps in western peninsular Malaysia (Selangor and Perak states). Several gouramis of the genus Betta are threatened by habitat degradation, overcollection for the international aquarium trade, and hybridization with released captive betta species. The Songkhla betta (B. ferox) is confined to a single locality in southern peninsular Thailand (Songkhla province). The Persephone betta (B. persephone) is known only from a small area of peat swamp forest in peninsular Malaysia (Johor

state). The Selangor betta (B. livida) is confined to peat swamps in peninsular Malaysia (Selangor and Perak states). The hillstream betta (B. apollon) is confined to a small area of south-eastern peninsular Thailand. The pallid betta (B. pallida) is confined to a few coastal drainages in south-eastern peninsular Thailand and to Ko Samui Island. The blackwater betta (B. hipposideros) is known only from peat swamps in peninsular Malaysia (Selangor state) and from the Indragiri River drainage in central-eastern coastal Sumatra (Riau province). Tom’s betta (B. tomi) is confined to peninsular Malaysia (Johor and a few offshore islands). The species formerly occurred on Singapore as well, but was last recorded there in 1937 and now certainly extirpated. Kuehne’s betta (B. kuehnei) is confined to a few coastal streams in north-eastern peninsular Malaysia and extreme south-eastern peninsular Thailand. The Krabi betta (B. simplex) is found only in a small area of karst spring and wetlands near Khao Thong in southern peninsular Thailand (Krabi province). The armoured stickleback (Indostomus crocodilus) is confined to a small area of the south-eastern Malay Peninsula, specifically the Toh Daeng peat swamp forest in Thailand (Narathiwat province), and north-western Malaysia (Kelantan state). It is threatened by habitat destruction and overcollection for the international aquarium trade. The Toh Daeng catfish (Pseudeutropius indigens) is known only from the Toh Daeng peat swamp forest in south-eastern peninsular Thailand (Narathiwat province). The Bebar-Sedili airbreathing catfish (Encheloclarias kelioides) was historically known from peat swamps within the Bebar and Sedili River drainages of southern peninsular Malaysia. Last reported in 1993, it may be extinct. The Selangor airbreathing catfish (E. curtisoma) is known only from the North Selangor peat swamp forest in southern peninsular Malaysia (Selangor). The Tenasserim stream catfish (Akysis pictus) is confined to southern peninsular Myanmar. The Kra Isthmus stream catfish (A. pulvinatus) is confined to the Kra Isthmus in southern peninsular Thailand. The enigmatic loach (Ellopostoma mystax) is known only from the Tapi and Saiburi river drainages in peninsular Thailand. The tri-banded stone loach (Schistura balteata) is known only from a small area of the southern Malay Peninsula (Myanmar and Thailand). The elongated eel-loach (Pangio elongata) is confined to hillstreams in peninsular Myanmar. The Tapi River The Tapi River is located in peninsular Thailand. The Tapi stone loach (Nemacheilus ornatus) is confined to the Tapi River drainage. The Muar River The Muar River (Sungai Muar in Malay) is located in peninsular Malaysia (Johor, Negeri Sembilan and Pahang states).

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Duncker’s priapium fish (Phallostethus dunckeri) is known only from the Muar River.

Coasts and Satellite Islands Coastal areas within the Malay Peninsula, and in particular its mangroves, are highly threatened by development, conversion to shrimp ponds and pollution. The striped mangrove goby (Mugilogobius fasciatus) is known only from a few disjunct mangrove creeks and pools within peninsular Malaysia. Singapore Singapore and its associated islets lie just slightly north of the equator, at the southernmost tip of peninsular Malaysia. Its environment has changed drastically due to rapid urbanization and immense population density. In 1819, when a British trading post was first established there, the island was still nearly covered in rainforest. Before long massive deforestation had begun and was largely completed by the twentieth century. By some estimates Singapore has lost about 95 per cent of its natural habitat. The Singapore mouse-eared bat (Myotis oreias) is known only from a single specimen purported to have been collected on Singapore. The Singapore caecilian (Ichthyophis singaporensis) is known only from a single specimen collected in the midtwentieth century. Penang Penang (Pulau Penang in Malay) is located in the Malacca Strait off the north-western coast of peninsular Malaysia. The Penang stream toad (Ansonia penangensis) is known only from Penang Island. The collection of tadpoles on Penang Hill in 2004 represented the first record for the species in over a century. The Penang Hill frog (Fejervarya pulla) is known only from its original collection in 1870. Tioman Tioman Island (Pulau Tioman in Malay) is located off the eastern coast of peninsular Malaysia. The Tioman pit viper (Popeia buniana) is confined to Tioman Island, where it is threatened by overcollection for the international pet trade. The Tioman ground snake (Gongylosoma mukutense), Boo-Liat’s kukri snake (Oligodon booliati) and Inger’s reed snake (Calamaria ingeri) are all known only from a single forest fragment, which is still under threat from development. The Tioman stream toad (Ansonia tiomanica) is confined to three localities on Tioman Island. The Kajang slender litter frog (Leptolalax kajangensis) is known with certainty only from a single small cave atop Mount Kajang, although larvae thought to be conspecific have been found at lower elevations. The Tioman stone loach (Speonectes tiomanensis) is known only from a single cave on Tioman.

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Great Perhentian Great Perhentian (Perhentian Besar in Malay) is located in the Perhentian Islands off the eastern coast of Malaysia. The Perhentian spectacled langur (Trachypithecus obscurus styx) is confined to Great Perhentian. The Riau Islands The Riau Islands (Kepulauan Riau in Indonesian) are a group of 1796 small islands scattered between the Malay Peninsula, Sumatra, and Borneo. The Riau pale-thighed surili (Presbytis siamensis rhionis) is known for certain only from Bintan, although it may occur on the nearby islands of Batam and Galang as well. The Riau betta (Betta spilotogena) is a type of freshwater fish confined to the islands of Bintan and Singkep. Bintan (Pulau Bintan in Malay) is located south of Singapore. The Bintan betta (Betta miniopinna) is a type of fish confined to Bintan.

Sumatra Sumatra (Sumatera in Indonesian) is a large island located in western Indonesia. It lies close to the Asian mainland and differs biogeographically only slightly from the nearby Malay Peninsula. While seriously affected by human activities, it nevertheless serves as a refuge for many species either very rare or no longer to be found anywhere else. The Sumatran elephant (Elephas maximus sumatranus) was historically widespread on the island, but now survives only in highly fragmented populations. It is threatened by habitat destruction, poaching, and conflict with humans. The western two-horned rhinoceros (Dicerorhinus sumatrensis sumatrensis), previously discussed in this volume, historically occurred in the Malay Peninsula and on Sumatra. It is now extirpated from the Asian mainland, and today only 75–85 are thought to remain on Sumatra, mostly in the national parks of Bukit Barisan Selatan and Gunung Leuser but also in Way Kambas National Park. There is a small captive population. Two species of orangutan (Pongo) are endemic to Sumatra. The Aceh orangutan (P. abelii) is confined to northern Sumatra, with the majority found in Aceh province. It was formerly far more widespread, with a notable preference for lowland forest and peat swamps. Habitat loss due to logging and the creation of oil palm plantations has severely reduced the species, which in 2016 was estimated to number around 14,000. The Tapanuli orangutan (P. tapanuliensis) was only recently described in 2017 when a study showed that an isolated population living in the Batang Toru region, south of Lake Toba, is distinct from all others in both Sumatra and Borneo. By far the rarest of the great apes, it is threatened by logging, mining activities, agricultural plantations and a proposed hydroelectric dam. The Sumatran tiger (Panthera tigris sumatrae) is the smallest of the tigers and the last surviving of the three

The Sundaic Region Figure 5.5 Aceh orangutan. (Credit: Justin D’Agostino.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

Indonesian island subspecies. It still occurs in northern Sumatra and in the south-western mountains, where the wild population was estimated in 2014 at 400–500. The Sumatran clouded leopard (Neofelis diardi diardi) was historically found throughout Sumatra as well as in the Batu Islands off the western coast, but now appears to be confined to a few protected areas. The Sumatran white-handed gibbon (Hylobates lar vestitus) is confined to northern Sumatra. The North Sumatran black surili (Presbytis sumatrana) is a type of leaf-eating monkey found in north-central Sumatra and on Pulau Pini in the Batu Archipelago. Thomas’ surili (P. thomasi) is confined to northern Sumatra (Aceh and Sumatera Utara provinces), in both montane and lowland rainforest. Both are threatened by loss of habitat and trapping for use as pets. The western tarsier (Cephalopachus bancanus) is a type of nocturnal primate divided into a number of subspecies. The Sumatran tarsier (C. b. bancanus) is confined to south-eastern Sumatra and the island of Bangka. Hiller’s slow loris (Nycticebus hilleri) is a small, nocturnal prosimian known from northern Sumatra, north of the Batang Toru River. It may possibly occur as well on small surroundings islands, but this has yet to be determined. The Sumatran bearded pig (Sus barbatus oi) was historically found throughout Sumatra and offshore islands, but is now confined to a few areas of central Sumatra. The Sumatran sambar deer (Rusa unicolor equina) is confined to Sumatra. The Simeulue rat (Rattus simalurensis) is known from the islands of Simeulue, Siumat, Lasia, and Babi off the northwestern coast of Sumatra. Modigliani’s black-eared flying fox (Pteropus melanotus modiglianii) was originally found on Nias, the Mentawai Islands and Enggano, but was extirpated from the latter island after a severe typhoon. The Nias hill myna (Gracula robusta) is confined to Nias and the Banyak Islands, where it has become seriously threatened due to overcollection for the pet trade.

The Sumatran ground cuckoo (Carpococcyx viridis) is known only from eight specimens (the last of which was collected in 1916) and a few more recent sightings, the majority of which have come from the Barisan Mountains in the south-western part of the island. The graceful pitta (Erythropitta venusta) is a rare type of passerine bird found widely but patchily in western Sumatra, where it is threatened by loss of habitat. The Sumatran leafbird (Chloropsis media) is found widely but patchily in western Sumatra, where it is everywhere threatened by loss of habitat and collection for the cage-bird trade. The Sumatran blue-wattled bulbul (Brachypodius nieuwenhuisii inexspectatus) is known only from a single specimen collected in 1937 from an undefined locality on Sumatra. The spatula-toothed snake (Iguanognathus werneri) is known only from a single specimen collected in 1898 from an unspecified locality in Sumatra. Forcart’s reed snake (Calamaria forcarti) is known only from four specimens collected from two, widely separated localities (north-eastern Sumatra and the island of Nias). The Sumatran worm snake (Argyrophis hypsobothrius) is known only from two specimens collected from an unknown locality in Sumatra. Peters’ toad (Ingerophrynus claviger) is known only from two disjunct localities (south-western Sumatra and the island of Nias). The elongated caecilian (Ichthyophis elongatus) is known for certain only from a small area of central-western Sumatra (Sumatera Barat province), although specimens collected from Pagai, Siberut, Pini, and Tanah Masa in the Mentawai Islands may also refer to this species. The Kapahiang caecilian (I. paucidentulus) is known only from Kepahiang and Kerinci Seblat national parks in south-western Sumatra (Bengkulu province).

The Barisan Mountains The Barisan Mountains (Bukit Barisan in Indonesian) are located along the northern and central coast of western

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Sumatra, extending nearly 1700 km from north to south. They consist primarily of active volcanoes still shrouded in dense rainforest on the lower levels, and tropical pine forests higher up. The bicoloured mitred surili (Presbytis melalophos bicolor) is a type of leaf-eating monkey confined to the highlands of west-central Sumatra from the Hari River to the Inderagiri River. The Sumatran striped rabbit (Nesolagus netscheri) is confined to disjunct areas of remote montane rainforest in the Barisan Mountains. The Sumatran mountain muntjac (Muntiacus montanus) is a small deer first discovered in 1914 but unrecorded since 1930 until one was snared (and released) in Kerinci Seblat National Park, central-western Sumatra, in 2002. Two others have since been photographed within the park. The Sumatran flying squirrel (Hylopetes winstoni) is known only from a single specimen collected in northern Sumatra (Aceh province) in 1949. The dwarf gymnure (Hylomys parvus) is a hedgehog-like species known only from Mount Kerinci and a few other nearby mountains in central-western Sumatra (Sumatera Barat, Jambi, Bengkulu and Sumatera Selatan provinces). The Sumatran mountain spiny rat (Maxomys hylomyoides) is known only from a few disjunct localities in central-western Sumatra. The broad-nosed spiny rat (M. inflatus) is found widely but patchily within the Barisan Mountains. Both are threatened by loss of habitat. The Kerinci rat (Rattus korinchi) is known only from two specimens collected from Mount Kerinci and Mount Talakmau in central-western Sumatra during the early twentieth century. Hoogerwerf’s rat (R. hoogerwerfi) is known only from a few specimens collected in north-western Sumatra (Aceh province). The Sumatran pheasant (Lophura inornata) is divided into two subspecies, both of which are threatened by habitat destruction and hunting. Salvadori’s Sumatran pheasant (L. i. inornata) is found in the central and southern Barisan Mountains, while Hoogerwerf’s Sumatran pheasant (L. i. hoogerwerfi) occurs in the same range to the north. The Sumatran cochoa (Cochoa beccarii) is a type of thrush known only from four specimens and a few sight records across the length of the Barisan Mountains. The Sumatran laughingthrush (Garrulax bicolor) is found widely but patchily within the Barisan Mountains, where it has suffered significant declines due to habitat destruction and trapping for the cage-bird trade. Schneider’s pitta (Hydrornis schneideri) is a grounddwelling passerine bird found widely but patchily within the Barisan Mountains, where it is threatened by loss of habitat. The Sumatran mesia (Leiothrix laurinae) is a brightly coloured passerine bird found widely but patchily in the Barisan Mountains, where it is threatened by overcollection for the international pet trade. The Aceh bulbul (Pycnonotus snouckaerti) is confined to the Barisan Mountains of northern Sumatra (Aceh province).

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It is threatened by habitat destruction and trapping for the cage-bird trade. Manthey’s anglehead lizard (Gonocephalus lacunosus) is known only from two specimens collected from a single locality in north-central Sumatra (Sumatera Utara province). The Sumatran limbless skink (Larutia sumatrensis) is known only from three localities in central-western Sumatra (Sumatera Barat province). Werner’s kukri snake (Oligodon pulcherrimus) is known only from a few specimens collected at the beginning of the twentieth century from three localities in northern Sumatra (Aceh, Sumatera Utara and Sumatera Barat provinces). Ulmer’s reed snake (Calamaria ulmeri) is known only from two specimens collected during the mid-twentieth century from northern Sumatra (Aceh province). The thick reed snake (C. crassa) is known only from a handful of specimens collected from Mount Talakmau and the Padang Highlands of central-western Sumatra (Sumatera Barat province). The Sumatran toad (Duttaphrynus sumatranus) is known only from its original collection during the nineteenth century in central-western Sumatra (Sumatera Barat province). The horned shrub frog (Philautus cornutus) is known only from two localities within the region of Mount Kerinci in central-western Sumatra (Jambi province). The Leuser Range The Leuser Range is located in north-western Sumatra (Aceh province). It consists of three prominent, non-volcanic peaks (Mount Leuser, Mount Loser, and Mount Tanpa-Nama). It is protected within Gunung Leuser National Park. The Aceh rat (Rattus blangorum) is known only from two specimens collected in the foothills of the Leuser Range. The Battak Range The Battak Range is located in north-western Sumatra (Sumatera Utara province). Durheim’s kukri snake (Oligodon durheimi) is known only from a single specimen collected in 1913 from the Battak Range. The Battak frog (Pulchrana debussyi) is known only from a single locality in the Battak Range. The Toba Caldera Complex The Toba Caldera Complex is located in north-western Sumatra (Sumatera Utara province), surrounding Lake Toba. At least four cones, four stratovolcanoes, and three craters are visible. The Toba pit viper (Popeia toba) is known only from a few specimens collected in the early twentieth century. Mount Talakmau Mount Talakmau (Gunung Talakmau, and also known as Mount Ophir) is a complex volcano located in central-western Sumatra (Sumatera Barat province). The Talakmau shrub frog (Philautus similis) is known only from Mount Talakmau.

The Sundaic Region

Mount Kerinci Mount Kerinici (Gunung Kerinci in Indonesian) is a stratovolcano located in central-western Sumatra (Sumatera Barat province), and the highest peak in the Barisan Mountains. It is protected within Kerinci Seblat National Park. The Sumatran shrewlike mouse (Mus crociduroides) is currently known only from high-elevation montane forest on Mount Kerinci, but may occur more widely in the Barisan Mountains. Jacobson’s false cobra (Pseudoxenodon jacobsonii) is known only from a single specimen collected on Mount Kerinci in the early twentieth century. The Kerinci keelback (Hebius kerinciense) is a type of water snake known only from a single specimen collected on Mount Kerinci. Mount Sago Mount Sago (Gunung Malintang in Indonesian) is located in central-western Sumatra (Sumatera Barat province). Wegner’s glass lizard (Dopasia wegneri) is known only from Mount Sago. The Padang Highlands The Padang Highlands are located in south-western Sumatra (Sumatera Selatan province). The Sumatran water shrew (Chimarrogale sumatrana) is known only from a single specimen collected from the Padang Highlands in the early twentieth century.

Lowland Rainforests and Swamp Forests At the beginning of the twentieth century Sumatra was still largely forested, with lowland rainforest predominating throughout the centre of the island (as well as on the smaller offshore islands), and peat swamps along the eastern coast. Sadly, after more than a century of massive deforestation little of this survives today beyond a few isolated patches, generally within protected areas. The black-crested surili (Presbytis melalophos) is a type of leaf-eating monkey found in central-western coastal Sumatra, roughly from the Rokan River south to the Hari River. The mitred surili (P. mitrata) is found in southeastern Sumatra. Both are threatened by loss of habitat and trapping for use as pets. The East Sumatran banded surili (P. femoralis percura) is confined to a small area of centraleastern Sumatra. Brooks’ fruit bat (Dyacopterus brooksi) is known only from a few specimens collected from lowland rainforests throughout Sumatra. The Sumatran mastiff bat (Mormopterus doriae) is known only from its original description in 1907 from north-eastern Sumatra (Sumatera Utara province). The Sumatran chestnut-necklaced partridge (Tropicoperdix charltonii atjenensis) is confined to a few disjunct areas of coastal northern and southern Sumatra (Aceh and Sumatera Selatan provinces).

Rück’s blue flycatcher (Cyornis ruckii) is known only from two specimens collected in 1917 and 1918 in north-eastern coastal Sumatra (Aceh and Sumatera Utara provinces). It is most likely extinct, a victim of habitat destruction. Modigliani’s nose-horned lizard (Harpesaurus modigliani) is known only from Si-Rambé Forest in northern Sumatra (Sumatera Utara province). The Ujungpadang snake (Anoplohydrus aemulans) is known only from a single specimen collected in the early twentieth century from an imprecise locality in north-western Sumatra (Aceh province). Petronella’s kukri snake (Oligodon petronellae) is known only from a few specimens collected from central-western coastal Sumatra (Sumatra Barat province). Döderlein’s reed snake (Calamaria doederleini) is known only from a single specimen collected in 1902 from northeastern Sumatra (Sumatera Utara province). The Bengkulu reed snake (C. alidae) is known only from four specimens collected in central-western Sumatra. Mechel’s reed snake (C. mecheli) is known only from a few specimens collected at the beginning of the twentieth century from an imprecise lowland locality in north-central Sumatra. The stripe-necked reed snake (C. margaritophora) is known only from a handful of specimens collected during the 1940s over a relatively wide area of central Sumatra. Eiselt’s reed snake (C. eiselti) is known only from its original collection in 1965 from centralwestern Sumatra (Sumatera Barat province). While the possibility exists that these cryptic burrowing forms may be more widespread than is currently known, it seems much more likely that they have been significantly reduced by the loss of their lowland rainforest habitat. The Padang burrowing snake (Etheridgeum pulchrum) is known only from a single specimen collected in 1924 from central-western coastal Sumatra (Sumatera Barat province), in what is now the major port city of Padang. It is most likely extinct due to loss of habitat. The Sumatran golden-backed frog (Hylarana persimilis) is known only from two specimens collected from north-eastern Sumatra (Sumatera Utara province).

Lakes, Rivers, and Marshes A number of large rivers drain from the Barisan Mountains through the lowlands of eastern Sumatra, creating intermittent swamplands. The viper-like keelback (Hebius viperinum) is a type of water snake known only from a single specimen collected in 1901 from central Sumatra (Riau province). The Serokan sharkminnow (Osteochilus serokan) is known only from two localities in north-western Sumatra (Aceh and Sumatera Utara provinces). Lake Toba Lake Toba (Danau Toba in Indonesian) is large lake occupying the caldera of a supervolcano in central-northern Sumatra (Sumatera Utara province).

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Theinemann’s brook carp (Neolissochilus theinemanni) is confined to Lake Toba and its inlet upstream tributaries. Lake Tawar Lake Tawar (Danau Tawar in Indonesian) is an ancient volcanic caldera located in north-western Sumatra (Aceh province). The Tawar barb (Poropuntius tawarensis) is confined to Lake Tawar, where it is seriously threatened by pollution, invasive species, water extraction, and overfishing. The Tawar rasbora (Rasbora tawarensis) is confined to Lake Tawar, where it is seriously threatened by pollution, invasive species, water extraction, and overfishing.

Weh Weh (Pulau Weh in Indonesian) is a small, volcanically active island located off the far northern tip of Sumatra. It has been heavily degraded and there is practically no forest left. The Weh kukri snake (Oligodon praefrontalis) is confined to Weh Island. The Weh toad (Duttaphrynus valhallae) is known only from a single specimen, and is possibly extinct.

Simeulue Simeulue (Pulau Simeulue in Indonesian) is located about 150 km off the north-western coast of Sumatra. The Simeulue long-tailed macaque (Macaca fascicularis fuscus) is confined to Simeulue. The Simeulue scops owl (Otus umbra) is confined to Simeulue. The Simeulue parrot (Psittinus abbotti) is confined to the islands of Simeulue and the islet of Siumat. Jacobson’s day gecko (Cnemaspis jacobsoni) is known only from a few specimens collected on Simeulue.

Lasia Lasia (Pulau Lasia in Indonesian) is a small island located between Simeulue and Nias, on the western coast of Sumatra. The Lasia long-tailed macaque (Macaca fascicularis lasiae) is confined to Lasia.

Nias Nias (Pulau Nias in Indonesian) is a low-elevation island located off the north-western coast of Sumatra. The Nias binturong (Arctictis binturong niasensis) is a large viverrid confined to Nias. The Nias reed snake (Calamaria abstrusa) is known only from a few specimens collected from Nias.

The Mentawai Islands The Mentawai Islands (Kepulauan Mentawai in Indonesian) are a chain of about 70 islands and islets lying alongside the western coast of Sumatra, the four largest of which are Siberut, Sipora, Pagai Utara, and Pagai Selatan. Isolated from the mainland for more than half a million years has allowed the

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survival of relicts from an earlier Indo-Malaysian fauna as well as the evolution of a number of endemic species, including several primates. Kloss’ gibbon (Hylobates klossii) is an all-black species resembling a small siamang (Symphalangus syndactylus). It is still found on all four of the main islands (Siberut, Sipora, Pagai Utara, and Pagai Selatan), with the largest remaining population occurring in Siberut National Park. Threats include hunting and commercial logging. The Pagai macaque (Macaca pagensis) is confined to the smaller southern islands of Pagai Utara, Pagai Selatan, and Sipora. The pig-tailed langur (Simias concolor) is a large, heavily built species thought to be related to the proboscis monkey (Nasalis larvatus) of Borneo. There are two subspecies found throughout the Mentawai Islands. The Pagai pig-tailed langur (S. c. concolor) is found on Pagai Utara, Pagai Selatan, Sipora, and the nearby islets of Simalegu and Sinakak. The Mentawai surili (Presbytis potenziani) is a type of leafeating monkey found on Sipora, North Pagai, South Pagai and the islet of Sinakak. The Siberut surili (P. siberu) is confined to Siberut. Both are threatened by loss of habitat and hunting. The Mentawai palm civet (Paradoxurus hermaphroditus lignicolor) is confined to the Mentawai Islands. The Mentawai flying squirrel (Iomys sipora) is confined to Sipora and Pagai Utara. The Mentawai dwarf flying squirrel (Petinomys lugens) is confined to Siberut, Sipora and Pagai Utara. The Mentawai black-bellied squirrel (Callosciurus melanogaster) is found patchily within the Mentawai Islands, where it is threatened by loss of habitat. The fraternal squirrel (Sundasciurus fraterculus) is confined to Siberut, Sipora, Pagai Utara and Pagai Selatan. The Mentawai long-tailed giant rat (Leopoldamys siporanus) is confined to Siberut, Sipora, Pagai Utara and Pagai Selatan. The Mentawai spiny rat (Maxomys pagensis) is confined to Siberut, Sipora, Pagai Utara and Pagai Selatan. The Mentawai rat (Rattus lugens) is known only from a few specimens collected at the beginning of the twentieth century from Siberut, Sipora, Pagai Utara and Pagai Selatan. The golden-bellied tree shrew (Tupaia chrysogaster) is confined to Siberut, Sipora, Pagai Utara and Pagai Selatan. The Mentawai pencil-tailed tree mouse (Chiropodomys karlkoopmani) is known only from Siberut and Pagai Utara. Siberut Siberut is the largest and northernmost of the Mentawai Islands. Much of the island remains forested despite being subject to commercial logging. In 1993 the western half of the island was set aside as Siberut National Park. The Siberut macaque (Macaca siberu) is confined to Siberut. The Siberut pig-tailed langur (Simias concolor siberu) is confined to Siberut, where it is highly threatened by habitat destruction and hunting.

The Sundaic Region

Sipora Sipora is located south of Siberut. The most developed of the Mentawai Islands, only some 10–15 per cent of its original rainforest cover remains. The Sipora flying squirrel (Hylopetes sipora) is littleknown but believed to be threatened by deforestation. The Pagai Islands are the southernmost of the Mentawai Islands. They consist of Pagai Utara (North Pagai) and Pagai Selatan (South Pagai), which are only separated by a narrow channel. The short-headed leaf-nosed bat (Hipposideros breviceps) is known only from its original collection in 1941 from Pagai Utara.

Enggano Enggano (Pulau Enggano in Indonesian) lies about 100 km south-west of Sumatra. The forests of this remote and biologically interesting island, which was likely never connected to the Sumatran mainland, have been almost completely destroyed. The Indonesian government has recently proposed it as the possible location for a spaceport. Two species of rat (Rattus) were historically endemic to Enggano, but have most likely been driven to extinction by introduced black rats (R. rattus). Miller’s rat (R. enganus) is known only from the holotype collected during the nineteenth century. The burnished rat (R. adustus) is known only from a single specimen collected in 1940. Surveys of the island in the late 1980s failed to find evidence of either species. The Enggano scops owl (Otus enganensis) is a rare species confined to Enggano. The Enggano imperial pigeon (Ducula oenothorax) is confined to Enggano. The Enggano hill myna (Gracula enganensis) is confined to Enganno. The Enggano thrush (Zoothera leucolaema) is confined to Enganno, where it remains relatively common. The Enggano rat snake (Coelognathus enganensis) has not been recorded since it was first collected in the 1930s, despite repeated surveys.

Bangka Bangka (Pulau Bangka in Indonesian) is a large island off the south-eastern coast of Sumatra. Most of the original forest has been converted into oil palm plantations. The Bangka binturong (Arctictis binturong kerkhoveni) is a large viverrid confined to Bangka Island. The Bangka slow loris (Nycticebus bancanus) is a small, nocturnal prosimian known only from Bangka, where it was last recorded in 1937. It is likely extinct due to habitat destruction, although the species may occur on neighbouring Belitung. The Bangka shrew (Crocidura vosmaeri) is a little-known species confined to the island’s remaining forests. Two species of gourami, the Koba betta (Betta chloropharynx) and the Bikang betta (B. burdigala), are confined to peat

swamp forest fragments in southern Bangka, where they are seriously threatened by habitat destruction. The Bangka airbreathing catfish (Encheloclarias tapeinopterus) is confined to peat swamps in eastern Bangka.

Belitung Belitung (Pulau Belitung in Indonesian/known as Billiton in English) is located south-east of Bangka. It has undergone extensive deforestation and very little habitat remains. The Belitung tarsier (Cephalopachus bancanus saltator) is confined to Belitung. The Belitung reed snake (Calamaria javanica) is known only from three specimens collected in 1891 on Belitung, and one other of doubtful provenance (‘Java’). The Belitung caecilian (Ichthyophis billitonensis) is known only from a single specimen collected in the 1960s.

The Natuna Islands The Natuna Islands are a 272-island archipelago located between peninsular Malaysia and the north-western coast of Borneo. The Natuna silvered langur (Trachypithecus cristatus vigilans) is confined to the Natuna Islands. The Natuna crested serpent eagle (Spilornis cheela natunensis) is confined to the Natuna Islands.

Great Natuna Island Great Natuna Island (Pulau Natuna Besar in Indonesian; also known as Bunguran) is the largest of the Natuna Islands. The Great Natuna surili (Presbytis natunae) is a type of leaf-eating monkey confined to Great Natuna Island, where it is highly threatened by loss of habitat due to logging. The Great Natuna dwarf litter frog (Leptobrachella natunae) is known only from its original collection in 1895. The Great Natuna sticky frog (Kalophrynus bunguranus) is known only from a few specimens. Serasan Serasan Island (Pulau Serasan in Indonesian) is a large island located in the southern Natuna Islands. The Natuna Islands tarsier (Cephalopachus bancanus natunensis) is a type of nocturnal primate known only from Serasan Island and possibly nearby Subi Island.

Borneo Borneo (Pulau Borneo in Malay/Kalimantan in Indonesian) is the third largest island in the world and the largest in Asia. Straddling the equator, it is located north of Java, west of Sulawesi and east of Sumatra. Politically speaking the island is divided between Malaysia in the north and north-west and Indonesia in the south and south-east. The Malaysian portion is subdivided into three states (Sarawak, Brunei and Sabah), while the Indonesian part is comprised of five provinces (Kalimantan Barat, Kalimantan Tengah, Kalimantan Selatan, Kalimantan Timur, and Kalimantan Utara).

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The Bornean elephant (Elephas maximus borneensis) was long thought to be a population of feral descendants of elephants introduced in previous centuries. Genetic evidence, however, now suggests that they may be indigenous to the island. In any case the form is confined to northern and north-eastern Borneo, where the total population is estimated in the low thousands. The eastern two-horned rhinoceros (Dicerorhinus sumatrensis harrissoni) was once common throughout Borneo. Today it is thought that no more than 15 individuals still survive in eastern Borneo (Kalimantan Timur, Indonesia), with a few more perhaps in northern Borneo (Sarawak, Malaysia). Several species and subspecies of gibbon (Hylobates) endemic to Borneo are threatened by habitat destruction and hunting. The Bornean white-bearded gibbon (H. albibarbis) is confined to south-western Borneo south of the Kapuas River and west of the Barito River. The grey gibbon (H. muelleri) is usually divided into three subspecies, although some scientists consider the latter to be distinct species. Müller’s grey gibbon (H. m. muelleri) is confined to southeastern Borneo, roughly from south of the Mahakam River and east of the Barito River. Abbott’s grey gibbon (H. m. abbottii) is confined to western Borneo north of the Kapuas River (Sarawak, Malaysia and Kalimantan Barat, Indonesia). Geoffroy Saint-Hilaire’s grey gibbon (H. m. funereus) is confined to north-eastern Borneo. The proboscis monkey (Nasalis larvatus), so-named for the adult male’s distinctive outsized nose, was historically found in lowland forest, always near water, over the whole of coastal Borneo as well as on the satellite islands of Berhala, Sebatik, and Laut. Two rather ill-defined subspecies are recognized. The stripe-naped proboscis monkey (N. l. larvatus) is by far the more widespread, while the plain-naped proboscis monkey (N. l. orientalis) is confined to north-eastern Kalimantan. Ongoing hunting and habitat destruction have decimated both populations over the past few decades. Several species and subspecies of surili (Presbytis) endemic to Borneo are threatened by loss of habitat as well as by hunting for use as food and in ‘traditional medicine’. The cross-marked surili (P. chrysomelas), historically common, is now among the world’s rarest primates. Two subspecies are normally recognized. The western cross-marked surili (P. c. chrysomelas) is confined to a few localities in western Brunei, western Sarawak (Malaysia) and western Kalimantan Barat (Indonesia). The eastern cross-marked surili (P. c. cruciger) is confined to a few localities in north-eastern Sarawak and Sabah, Malaysia. The maroon surili (P. rubicunda) is divided into several subspecies collectively found throughout much of Borneo. Müller’s maroon surili (P. r. rubicunda) is found in south-eastern Borneo (Kalimantan Selatan), roughly east of the Barito River and south of the Mahakam River. The orange-backed maroon surili (P. r. chrysea) is confined to a small area of northern Borneo (eastern Sabah, Malaysia). Lyon’s maroon surili (P. r. rubida)

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is found in south-western Borneo, roughly south of the Kapuas River and west of the Barito River. The orange-naped maroon surili (P. r. ignita) is found in northern and western Borneo, roughly between the Kapuas and Baram rivers. The whitefronted surili (P. frontata) is found patchily in central and eastern Borneo, with a few additional populations in the west. Hose’s surili (P. hosei) is found in northern Borneo, where it is divided into two subspecies. The Niah Hose’s surili (P. h. hosei) appears to be confined to a small strip of forest along the lower Baram River in coastal Sarawak, Malaysia, where it is completely isolated by oil palm plantations. The Brunei Hose’s surili (P. h. everetti) is found in Brunei, Sabah and Sarawak (Malaysia) and Kalimantan Utara (Indonesia). Miller’s surili (P. canicrus) is confined to an area of eastern Borneo (Kalimantan Utara and Kalimantan Timur, Indonesia). The Sabah surili (P. sabana) is confined to far north-eastern Borneo (Sabah, Malaysia). The Bornean tarsier (Cephalopachus bancanus borneanus) is a type of nocturnal primate found throughout Borneo and the Karimata Islands, but is everywhere threatened by loss of habitat, collection for the pet trade, and persecution in the false belief that it is an agricultural pest. The Bornean slow loris (Nycticebus menagensis) is a small, nocturnal prosimian that, as a species, is widespread throughout Borneo and a few neighbouring islands. Three subspecies are recognized, which some scientists consider to be distinct species. The north Bornean slow loris (N. m. menagensis) is found in northern and eastern Borneo as well as on a few of the smaller islands of the Sulu Archipelago. The Kayan slow loris (N. m. kayan) is found in northern and central Borneo, while the south Bornean slow loris (N. m. borneanus) is found in central and south-western Borneo. All are threatened by loss of habitat and collection for use as pets. The Bornean clouded leopard (Neofelis diardi borneensis) is relatively widespread in Borneo but is everywhere rare and threatened by loss of habitat. The Bornean banteng (Bos javanicus lowi) is a type of wild cattle found disjunctly throughout Borneo. The Bornean sambar deer (Rusa unicolor brookei) is confined to Borneo. Hose’s palm civet (Diplogale hosei) is a cryptic and elusive species from northern and central Borneo, where it was long known only from a small number of museum specimens. In 1997 the first living specimen was obtained, and released after 2 months. The Bornean ferret-badger (Melogale everetti) is only known for certain from a small area of far northern Borneo (north-western Sabah). The tufted ground squirrel (Rheithrosciurus macrotis) is widespread in Borneo but is everywhere threatened by habitat destruction and trapping. The striped tree shrew (Tupaia dorsalis) is widely distributed within the rainforests of north-central Borneo, but is still known only from a few specimens and appears to be everywhere rare.

The Sundaic Region

The coppery pipistrelle bat (Arielulus cuprosus) is known only from a few scattered localities in northern and western Borneo (Sarawak and Sabah, Malaysia). Hartert’s frogmouth (Batrachostomus harterti) is a nocturnal, nightjar-like species known only from specimens collected from four localities in west-central Borneo (Sarawak, Malaysia and Kalimantan Barat, Indonesia). Bulwer’s pheasant (Lophura bulweri) was historically common throughout Borneo, particularly in the more remote areas of the interior, but has undergone a significant decline in recent decades due to loss of habitat and hunting. The blue-wattled bulbul (Brachypodius nieuwenhuisii) is a type of passerine bird divided into two naturally rare subspecies. The Bornean blue-wattled bulbul (B. n. nieuwenhuisii) is known only from a single specimen collected in north-eastern Kalimantan in 1900, along with a few other sight records in more recent years from from Brunei. The blue-headed pitta (Hydrornis baudii) is a passerine bird still found widely, although patchily, throughout Borneo. It has suffered significant declines due to loss of habitat, particularly in the lowland primary rainforests that it prefers. The Baram false cobra (Pseudoxenodon baramensis) is known only from a few specimens collected in northern Borneo (Sarawak, Malaysia). The Arqus keelback (Hebius arqus) is a type of snake known only from a single specimen, believed to have been collected in south-western Borneo (Kalimantan Barat, Indonesia). The bridled keelback (H. frenatum) is known only from two localities in north-western Borneo (central Sarawak, Malaysia). Battersby’s reed snake (Calamaria battersbyi) is known only from a single specimen collected during the 1960s from an imprecise locality in south-western Borneo (Kalimantan Barat, Indonesia). Lumholz’s reed snake (C. lumholtzi) is known only from three specimens collected from three widely separated localities in east-central Borneo. The Mulu flathead toad (Pelophryne api) is known only from Gunung Mulu National Park in Borneo (northern Sarawak). The masked gliding tree frog (Rhacophorus angulirostris) is largely confined to the Crocker Range and Mount Kinabalu National Park in northern Borneo (northern Sabah, Malaysia), with another small population known from central Borneo (Kalimantan Timur and Kalimantan Tengah, Indonesia). The Kerangas shrub frog (Philautus kerangae) is known only from two widely separated areas of northern Borneo (Sarawak, Malaysia), specifically Gunung Mulu National Park and Santubong National Park. The Lanjak shrub frog (P. refugii) is known only from a few scattered localities in western Borneo (south-western Sarawak, Malaysia). The Mendolong shrub frog (P. aurantium) is known only from a few localities in far northern Borneo (western Sabah, Malaysia). All are threatened by loss of habitat. The small rock frog (Staurois parvus) is known only from a few localities in northern Borneo (Sabah and Sarawak, Malaysia and Brunei).

The Kinabalu dwarf litter frog (Leptobrachella arayai) is confined to a few localities in far northern Borneo (Sabah, Malaysia). The striped dwarf litter frog (L. serasanae) has a highly disjunct distribution in west-central Borneo (Sarawak, Malaysia) and on Serasan Island off the south-western coast of Borneo. The Metang caecilian (Ichthyophis biangularis) was long known only from a single specimen collected in 1872 from Mount Metang in Borneo (Sarawak, Malaysia). It was rediscovered at the same locality in 2009.

The Central Highlands The Central Highlands stretch throughout the central interior of Borneo, with additional outlying mountains as well. They are among the highest in South East Asia and provide a habitat refuge for many species. The sculptor squirrel (Glyphotes simus) is confined to the mountains of northern Borneo (Sabah and Sarawak, Malaysia). The Bornean smooth-tailed tree shrew (Dendrogale melanura) is divided into two subspecies. The Sarawak smooth-tailed tree shrew (D. m. melanura) is known from a few localities in northern Sarawak (Mount Dulit, Mount Mulu, and the Kelabit Highlands). The Sabah smooth-tailed tree shrew (D. m. baluensis) is known from Mount Kinabalu and Mount Trus Madi. Neither has been recorded since the early 1970s. The grey-bellied pencil-tailed tree mouse (Chiropodomys muroides) is known only from a few specimens collected from Mount Kinabalu in northern Borneo (Sabah, Malaysia) and at another locality in eastern Borneo (Kalimantan Timur, Indonesia). The Bornean water shrew (Chimarrogale phaeura) is a semi-aquatic species that is only definitely known from the mountains of far northern Borneo (western Sabah, Malaysia), but may occur in suitable areas further south as well. Cox’s leaf-nosed bat (Hipposideros coxi) is known from a few localities in northern and western Borneo (Sarawak, Malaysia). The Bornean serpent-eagle (Spilornis kinabaluensis) is a rare species confined to the mountains of northern and central Borneo. Hose’s partridge (Rhizothera dulitensis) is known only from specimens collected during the nineteenth century from Mount Dulit and Mount Batu Song in Sarawak and from Mount Kinabalu in Sabah. The species has not been positively recorded since 1937, although the areas where it lives have been little surveyed. Guibe’s stream toad (Ansonia guibei) is confined to Mount Kinabalu and Mount Trusmadi in far northern Borneo (Sabah, Malaysia). Its habitat within the former locality may have been largely or entirely destroyed by landslides in 2015. Inger’s stream toad (A. latidisca) is known only from two localities in western Borneo (Sarawak, Malaysia and Kalimantan Barat, Indonesia). Last recorded in 1924, it was rediscovered in 2011. Inger’s shrub frog (Philautus ingeri) is known only from a few localities in northern Borneo (Sabah and Sarawak,

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Malaysia and Brunei), where it is threatened by loss of habitat. The Muruk Miau shrub frog (P. erythrophthalmus) was long known only from a single specimen collected on Mount Muruk Miau in western Sabah (Malaysia), but has since been discovered at a second locality in southwestern Sabah. The Crocker Range The Crocker Range (Banjaran Crocker in Malaysia) is located in northern Borneo (north-western Sabah, Malaysia). It is partially protected within Crocker Range National Park and Kinabalu Park. Mount Kinabalu (Gunung Kinabalu in Malay) is the fourth highest peak in South East Asia. It and the surrounding region have long been protected within Mount Kinabalu National Park, a World Heritage Site. The black pygmy white-toothed shrew (Suncus ater) is known only from a single specimen collected on Mount Kinabalu in the 1960s. The Kinabalu water snake (Hydrablabes praefrontalis) is known only from a single specimen collected on Mount Kinabalu in the 1890s. The white-striped reed snake (Calamaria lateralis) is known only from two specimens collected from Mount Kinabalu in the late nineteenth century. The Kinabalu giant blind snake (Anomochilus monticola) is known only from three specimens collected on Mount Kinabalu. The Kinabalu stream toad (Ansonia fuliginea) is confined to Mount Kinabalu. The Kinabalu shrub frog (Philautus gunungensis), Kamboranga shrub frog (P. amoenus), and Sauer’s shrub frog (P. saueri) are all confined to Mount Kinabalu. The Kinabalu large-eyed litter frog (Leptobrachium gunungense) is confined to Mount Kinabalu. The Tama Abu Range The Tama Abu Range is located in northern Borneo (northeastern Sarawak, Malaysia). Mount Mulu (Gunung Mulu in Malay) is protected within the boundaries of Gunung Mulu National Park, a World Heritage Site. The raspy shrub frog (Philautus acutus) and the pinnacle shrub frog (P. umbra) are both confined to submontane and montane forest on Mount Mulu. Dring’s dwarf litter frog (Leptobrachella brevicrus) is confined to Mount Mulu. The Kelabit Highlands The Kelabit Highlands are a vast highland plateau in northern Borneo (north-eastern Sarawak, Malaysia). The Eok sticky frog (Kalophrynus eok) is known only from a single specimen collected in the Kelabit Highlands. Mount Murud (Gunung Murud in Malay) is the highest mountain in Sarawak.

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The Linanit flathead toad (Pelophryne linanitensis) and Murud flathead toad (P. murudensis) are both confined to Mount Murud. The green-eyed tree frog (Polypedates chlorophthalmus) is known only from Mount Murud. Mount Dulit Mount Dulit (Gunung Dulit in Malay) is located in northern Borneo (north-central Sarawak, Malaysia). A western outlier of the Central Highlands, it is still largely covered with tropical rainforest. The Dulit caecilian (Ichthyophis dulitensis) is confined to Mount Dulit.

Lowland Rainforests and Swamp Forests The still considerable lowland rainforests of Borneo are among the oldest and most biologically important in the world. They consist mainly of primary and peat swamp forest, as well as Sundaland heath forest (also known as Kerangas forest), a type of lowland forest found on Borneo as well as the islands of Bangka and Belitung. The latter is characterized by a low, uniform canopy with thick underbrush and a rich growth of moss and epiphytes. All are threatened by massive deforestation due to logging, expansion of oil palm plantations and fires. The Bornean orangutan (Pongo pygmaeus) is a lowland forest specialist, rarely being found above 500 m. As a result of the enormous loss of these forests since the mid-twentieth century the species – along with its three subspecies, the north-west Bornean orangutan (P. p. pygmaeus), the northeast Bornean orangutan (P. p. morio) , and the south-west Bornean orangutan (P. p wurmbii) – has become seriously threatened. It is also illegally traded for use as pets, a practice with an inherently high mortality rate (i.e. the mother is usually shot in order to obtain the young, with a high percentage of the latter succumbing to mistreatment). The Bornean sun bear (Helarctos malayanus euryspilus) was once found throughout the island’s lowland rainforests, but has been eliminated from many areas. The Bornean bay cat (Catopuma badia), one of the world’s least-known wild cats, is found throughout the lowland, swamp and hill forests of Borneo but is rarely encountered. It is frequently trapped for its skin as well as for use in the international pet trade. The Bornean binturong (Arctictis binturong pageli) is large and little-known viverrid from the lowland rainforests of Borneo. The lesser pygmy flying squirrel (Petaurillus emiliae) was long known only from a single specimen collected in 1901 from northern Sarawak (Malaysia). It was rediscovered in 2013 from a second locality in western Borneo (Kalimantan Barat, Indonesia). Hose’s pygmy flying squirrel (P. hosei) is known only from a few specimens collected over a small area of northern Borneo (Sabah and Sarawak, Malaysia and Brunei). The Bornean tree rat (Pithecheirops otion) is known only from a single specimen collected in the Danum Valley of northern Borneo (north-eastern Sabah, Malaysia).

The Sundaic Region

Margaret’s ranee mouse (Haeromys margarettae) is known only from two old historical collections from two, widely separated localities in northern Sabah and south-western Sarawak. The Bornean peacock-pheasant (Polyplectron schleiermacheri) is a rare and elusive species found sporadically in dense lowland forests throughout the island. The Bornean crestless fireback pheasant (Lophura pyronota) is found widely but patchily throughout Borneo, where it is notably rare and localized. The Bornean crested fireback pheasant (L. ignita) is divided into two subspecies. The lesser Bornean crested fireback pheasant (L. i. ignita), and the greater Bornean crested fireback pheasant (L. i. nobilis) are both threatened by loss of habitat and hunting. Graydon’s chestnut-necklaced partridge (Tropicoperdix charltonii graydoni) is confined to a small area of northern Borneo (Sabah, Malaysia). The black-browed babbler (Malacocincla perspicillata) is a mysterious passerine bird, which, for 170 years, was known only from a single specimen collected from southern Borneo (Kalimantan Selatan, Indonesia). The species was thought to be extinct, an early victim of habitat destruction, until the unexpected discovery of a second specimen within the same province in 2020. The Bornean wren-babbler (Ptilocichla leucogrammica) remains widespread across the lowland forests of Borneo, but is generally rare and localized. The earless monitor (Lanthanotus borneensis) is a rare semi-aquatic lizard known only from northern Borneo in Sarawak (Malaysia) and from Kalimantan Utara and Kalimantan Timur (Indonesia). The Bornean spinejaw snake (Xenophidion acanthognathus) is known only from two disjunct localities in northern and western Sabah (Malaysia). Rebentisch’s reed snake (Calamaria rebentischi) is known only from a single specimen collected in 1860 from coastal western Borneo (Kalimantan Barat, Indonesia). With virtually no forest habitat now remaining in the area, the species is almost certainly extinct. Prakke’s reed snake (C. prakkei) is known only from its original collection in 1983 from Sandakan Bay in coastal northern Borneo (north-western Sabah, Malaysia). Habitat in the area has since been destroyed, and the species is most likely extinct. The slender reed snake (C. gracilima) is known only from four specimens collected from south-western Sarawak (Malaysia). The Engkari pipe snake (Cylindrophis enkariensis) is known only from a single specimen collected in southern Sarawak (Malaysia). The Akah River flathead toad (Pelophryne macrotis) is known only from a single specimen collected during the late nineteenth century from an imprecise locality in Sarawak (Malaysia). Günther’s flathead toad (P. guentheri) is known only from a few specimens collected over a wide area of Sabah and Sarawak (Malaysia). The Sarawak flathead toad (P. rhopophilia) is known only from two disjunct localities in north-

eastern and south-western Sarawak (Malaysia). All are presumably threatened by loss of habitat. The Gunung Gading gliding tree frog (Rhacophorus gadingensis) is known only from two protected areas in coastal south-western Sarawak (Malaysia). The disparate shrub frog (Philautus disgregus) is known only from three disjunct areas of eastern Sabah and western Sarawak (Malaysia), where it is threatened by habitat destruction. The Kenepai wart frog (Limnonectes kenepaiensis) is confined to a few localities in western Borneo (Sarawak, Malaysia and Kalimantan Barat, Indonesia). The large-eyed torrent frog (Meristogenys macrophthalmus) is known only from a single specimen collected in central Sarawak (Malaysia). The Matang torrent frog (M. jerboa) is known only from a small area of south-western Sarawak (Malaysia). The Sabah rice frog (Microhyla maculifera) is known only from the Danum Valley Conservation Area in northern Borneo (Sabah, Malaysia). The Lipaso dwarf litter frog (Leptobrachella palmata) is confined to a small area of northern Borneo (northcentral Sabah). Smith’s balloon frog (Glyphoglossus smithi) is known only from two or three disjunct areas of Sabah and Sarawak, Malaysia. The Mahakkam caecilian (Ichthyophis asplenius) is known only from two localities in south-western Sarawak (Malaysia). The black caecilian (I. monochrous) is known only from a few specimens collected from three widely separated localities in western Borneo (Sarawak, Malaysia and Kalimantan Barat, Indonesia). The Long Bloee caecilian (I. atricollaris) is known only from a few specimens collected during the 1960s from northern Borneo (Sarawak, Malaysia).

Isolated Caves, Springs, and Pools Borneo has a number of significant cave systems, primary in the northern and eastern parts of the island. Clearwater Cave in Sarawak features one of the world’s longest underground rivers, while Deer Cave is home to over three million bats, their guano having accumulated over 100 m deep. The Niah Caves in Sarawak is an important prehistoric site where c.40,000-year-old human remains along with paintings and implements have been discovered. Gunung Mulu National Park in Sarawak, also in Sarawak, and the SangkulirangMangkalihat Karst in eastern Borneo (Kalimantan Timur, Indonesia) both contain thousands of smaller caves. The cave bent-toed gecko (Cyrtodactylus cavernicolus) is known only from the Niah and Mulu cave systems in northeastern Borneo (Sarawak, Malaysia).

Lakes, Rivers, and Marshes Borneo has several major river systems and still extensive areas of peat swamp forest, although few lakes. The aquatic flat-headed toad (Barbourula kalimantanensis) is a fully aquatic, lungless species known only from a few

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localities in west-central Borneo (Kalimantan Barat and Kalimantan Tengah, Indonesia). It is threatened by habitat destruction and pollution. Schlueter’s frog (Fejervarya schlueteri) is known only from a single specimen collected during the nineteenth century from an unspecified locality in Borneo. Hose’s barb (Leptobarbus hosii) is confined to a few rivers in northern Borneo (Sarawak and Sabah, Malaysia). The spotfin betta (Betta macrostoma) is confined to a small area of northern Borneo (Brunei and northern Sarawak, Malaysia). Chin’s betta (B. chini) is confined to peat swamps and hillstreams in in northern Borneo (western Sabah (Malaysia). Both are threatened by loss of habitat and overcollection for the international aquarium trade. The Sarawak airbreathing catfish (Encheloclarias prolatus) is known only from an unspecified peat swamp locality in northern Borneo (Sarawak, Malaysia). The Sabah loach (Protomyzon borneensis) is found patchily in northern Borneo (Sabah, Malaysia), where it is threatened by habitat destruction and degradation. The Sabah stone loach (Sundoreonectes sabanus) is known only from a small area of north-eastern Borneo (western Sabah, Malaysia).

The Kapuas River The Kapuas River (Sungai Kapuas in Indonesian) is the longest river in Indonesia. It originates in the highlands of central Borneo and flows west through Kalimantan Barat (Indonesia) into the South China Sea, in the process creating an extended marshy delta. In recent years gold mining in the river has significantly increased environmental disturbances, in particular mercury pollution. The Kapuas water snake (Enhydris gyii) is known only from three specimens collected from the Kapuas River. Valliant’s chocolate gourami (Sphaerichthys vaillanti) is confined to the Kapuas River drainage. The Kapuas betta (Betta pinguis) is confined to forest streams in the middle Kapuas River drainage. Myers’ bumblebee catfish (Pseudomystus myersi) is confined to the Kapuas River drainage. The Kapuas stream catfish (Acrochordonichthys chamaeleon) is confined to the Kapuas River drainage. The Pinoh River is located in west-central Borneo (Kalimantan Barat, Indonesia). The Pinoh rasbora (Rasbora ennealepis) is confined to tributaries of the Pinoh River.

The Rajang River The Rajang River (Batang Rajang in Malay) is located in northern Borneo (Sarawak and Sabah, Malaysia, as well as Brunei). The Rajang sharkminnow (Osteochilus sarawakensis) is known only from the Rajang River.

The Sibau River The Sibau River (Sungai Sibau in Indonesian) is located in south-western Borneo (Kalimantan Barat, Indonesia). The Kalimantan sea snake (Hydrophis sibauensis) is known only from three specimens collected within a swampy area of the Sibau River, more than 1000 km upriver from the coast.

The Kinabatangan River The Kinabatangan River (Sungai Kinabatangan in Malay) is located in north-eastern Borneo (Sabah, Malaysia). The Bornean river shark (Glyphis fowlerae) is known only from 13 specimens collected from the Kinabatangan River in 1996. More recent expeditions failed to find any, and the species has not been seen by local fishermen for many years. The dwarf horsefaced loach (Acantopsis octoactinotos) is confined to the Kinabatangan River. It is a relatively common aquarium fish. The Kayan River The Kayan River is located in north-eastern coastal Borneo (Kalimantan Utara, Indonesia). The Bulungan sheatfish (Kryptopterus lumholtzi) is known only from a single specimen collected in 1914 from the Kayan River drainage. The Mahakam River The Mahakam River (Sungai Mahakam in Indonesian) is located in eastern Borneo (Kalimantan Timur, Indonesia). The Mahakam spineless eel (Chendol lubricus) is confined to the Mahakam River drainage.

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The Karimata Islands The Karimata Islands are a group of small islands located off the south-western coast of Indonesian Borneo. The Karimata maroon surili (Presbytis rubicunda carimatae) appears to be confined to the main island (Pulau Karimata) in the Karimata Islands.

Miang Miang (Pulau Miang in Indonesian) is located off the eastern coast of Borneo (Kalimantan Timur, Indonesia). Werner’s skink (Lipinia miangensis) is known only from Miang Island. Lorenz’s blind snake (Ramphotyphlops lorenzi) is known only from Miang Island, where it was last recorded in 1909.

Maratua Maratua (Pulau Maratua in Indonesian) is one of the Derawan Islands located off the eastern coast of Borneo. The Maratua long-tailed macaque (Macaca fascicularis tua) is confined to Maratua.

Bunyu Bunyu (Pulau Bunyu in Indonesian) is located off the northeastern coast of Borneo in the eastern Celebes Sea.

The Sundaic Region

Koekkoek’s blind snake (Malayotyphlops koekkoeki) is known only from Bunyu Island. An additional record from northern Sarawak is almost certainly a misidentification.

Java Java (Jawa in Indonesian) is a large, volcanically active island located south of Sumatra and Borneo. With a current figure of over 140 million people, it is the world’s most densely populated. This staggering increase from just three or four million in 1800 has led to massive forest loss and heavy industrialization, and there is not much left of the original environment. Most of the larger animals have already been exterminated, among them tigers and Asian elephants. The Sundaic lesser one-horned rhinoceros (Rhinoceros sondaicus sondaicus), previous discussed in this chapter and now better known simply as the Javan rhinoceros, was historically found on both Java and Sumatra. Since the 1930s, however, it has been entirely restricted to the lowland rainforest of the Ujung Kulon Peninsula in far western Java (Banten province), where the total population has remained stable at between 30 and 65. Wild, pure-bred populations of the Javan banteng (Bos javanicus javanicus), a type of wild cattle, are confined to a few protected areas on Java and, perhaps, Bali. The most important strongholds are in Ujung Kulon National Park in western Java (Banten province) and in Baluran National Park (Jawa Timur province). Other bantengs in Java are hybrids cross-bred with domestic cattle. The Javan tiger (Panthera tigris sondaica) was historically numerous throughout the island. By the 1940s it had been eliminated from most areas, including the important Ujung Kulon and Baluran reserves, and by 1972 only five individuals

were thought to remain. It was last positively recorded in 1976 in Meru Betiri National Park in eastern Java (Jawa Timur province). The subspecies has since been declared extinct, a victim of hunting and forest loss. The Javan leopard (Panthera pardus melas) is thought to be present in most forested regions of the island, although in very low numbers. The Javan or silvery gibbon (Hylobates moloch) was historically found throughout the island in both lowland and montane rainforest, but has been reduced to a few scattered patches mainly in protected areas. The largest viable population, about 1000, lives in the still extensive rainforests of Mount Halimun Salak National Park in western Java (Banten and Jawa Barat provinces). The Javan langur (Trachypithecus auratus) is a type of leafeating monkey found throughout Java as well as on Bali and a few other smaller islands. There are two subspecies. The eastern Javan langur (T. a. auratus) occurs on eastern Java, Bali, Sempu, and Nusa Barong, with an additional introduced population on Lombok. The western Javan langur (T. a. mauritius) is confined to western Java. Both are threatened by habitat destruction and hunting. The Javan surili (Presbytis comata) is confined to a few pockets in western and central Java. The Javan slow loris (Nycticebus javanicus) is a small, nocturnal prosimian found patchily in western and central Java, where it is threatened by loss of habitat and collection for the pet trade as well as use in ‘traditional medicine’. The Javan warty pig (Sus verrucosus) is divided into two subspecies. The nominate form (S. v. verrucosus) was historically found throughout Java and Madura, but has been extirpated from the latter island and is now found only in a few scattered pockets on the former. Figure 5.6 A Javan tiger photographed in Ujung Kulon in 1938. (Credit: Andries Hoogerwerf.)

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The Sunda rusa deer (Rusa timorensis) is believed to be native only to Java and Bali, but has been introduced to many other islands of the Indo-Pacific region. The Javan rusa deer (R. t. russa) is confined to a few scattered pockets across the island, where it is threatened by hunting and snaring even within protected areas. The red tree rat (Pithecheir melanurus) is known only from a small area of western Java (Jawa Barat province). The Javan giant rat (Sundamys maxi) is known only from two localities in western Java (Jawa Barat province). Sorensen’s leaf-nosed bat (Hipposideros sorenseni) is known only from a single cave in south-western coastal Java (Jawa Barat province). The Madura horseshoe bat (Rhinolophus madurensis) is known only from seven specimens collected from Madura and the Kangean Islands. Canut’s horseshoe bat (R. canuti) is known only from a few scattered localities on Java, Bali and the island of Nusa Barong, where it is threatened by loss of habitat. The pungent pipistrelle bat (Falsistrellus mordax) appears to be known only from nineteenth-century historical records from an undefined locality or localities on Java. The Javan hawk-eagle (Nisaetus bartelsi) is a rare species confined to forest patches scattered throughout Java. The white-faced partridge (Arborophila orientalis) is known only from a few localities in eastern Java (Jawa Timur province), where it is threatened by habitat destruction and hunting. Three starlings of the genus Acridotheres endemic to Java are seriously threatened by loss of habitat and capture for the cagebird trade. The black-winged myna (A. melanopterus) was historically common in western and central Java and on Madura, but is now virtually extinct in the wild. The grey-backed myna (A. tricolor) was formerly found throughout eastern Java, but now appears to be confined to the Meru Betiri and Baluran national parks. The white-vented myna (A. javanicus) is naturally found only on Java and Bali, but populations have been introduced to Taiwan, Singapore, Thailand, Sumatra, the Lesser Sunda Islands, Puerto Rico and possibly Japan. The Javan green magpie (Cissa thalassina) is a very rare species now known only from four protected areas in western and central Java, where the total population likely numbers fewer than 250. The species has suffered serious declines due to habitat destruction and capture for the cage-bird trade. Horsfield’s flameback (Chrysocolaptes strictus) is a rare type of woodpecker confined to eastern Java, Bali and the Kangean Islands. The white-rumped woodpecker (Meiglyptes tristis) is a rare species confined to western Java (Banten, Jawa Barat and Jawa Tengah provinces), where it is threatened by loss of habitat. The Javan blue-banded kingfisher (Alcedo euryzona) has, since the 1930s, been recorded only from the region of Mount Halimun in western Java (Banten and Jawa Barat provinces). The Javan finch (Padda oryzivora) is native to Java, Bali and probably Madura, where it was at one time abundant but has undergone a massive population decline in recent decades due to uncontrolled trapping for the cage-bird trade. The species has, however, been widely introduced in many parts of the world.

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Underwood’s bronzeback tree snake (Dendrelaphis underwoodi) is known only from three specimens collected from plantations in western Java (Jawa Barat province). The Javan keelback (Rhabdophis chrysargoides) is a type of snake known only from a few specimens collected from mostly undefined localities. Bernis’ caecilian (Ichthyophis bernisi) and Taylor’s caecilian (I. javanicus) are each known only from a single specimen collected from an undefined locality in Java.

The Javan Volcanic Chain Java was formed mainly by volcanic processes, with a chain of 31 active volcanoes and other mountains forming an east–west spine along the length of the island. Some of these still have some or all of their original cover of rainforest, providing a refuge for wildlife. The Javan scops owl (Otus angelinae) is found patchily throughout the Javan Volcanic Chain, where it is threatened by loss of habitat. The Javan blue-tailed trogon (Apalharpactes reinwardtii) is confined to a few forested mountains in western Java (Banten and Jawa Barat provinces), where it is threatened by habitat destruction and possibly trapping. The Javan cochoa (Cochoa azurea) is a type of thrush confined to a few montane forest localities in western and central Java. The rufous-fronted laughingthrush (Garrulax rufifrons) was historically common within the mountains of western and central Java, but is now seriously threatened by loss of habitat and overcollection for the cage-bird trade. All known wild populations are of the nominate subspecies (G. r. rufifrons). The bleeding toad (Leptophryne cruentata) is confined to a few montane localities in western and central Java (Jawa Barat and Jawa Tengah provinces). Mount Gede Pangrango Mount Gede Pangrango (Gunung Gede Pangrango in Indonesian) is a stratovolcano located in western Java (Jawa Barat province) containing two peaks (known as Mount Gede and Mount Pangrango). The area is protected within Mount Gede Pangrango National Park. The Javan bamboo rat (Kadarsanomys sodyi) is known only from a single collection on Gede Pangrango in the 1930s. Bartels’ flying squirrel (Hylopetes bartelsi) is confined to the forests of Mount Pangrango. The Javan thick-thumbed bat (Glischropus javanus) is known only from a few specimens collected nearly 70 years apart on Mount Pangrango. Barbour’s shrub frog (Philautus pallidipes) is known only from three specimens collected from Mount Pangrango in the early twentieth century. Mount Ungaran Mount Ungaran (Gunung Ungaran in Indonesian) is a deeply eroded stratovolcano located in central Java (Jawa Tengah province).

The Sundaic Region

Jacobson’s shrub frog (Philautus jacobsoni) is known only from a single specimen collected in 1912. All habitat from which it was obtained has now been completely deforested, and the species is likely extinct. Mount Slamet Mount Slamet (Gunung Slamet in Indonesian) is an active stratovolcano located in central Java (Jawa Tengah province). The Slamet rufous-fronted laughingthrush (Garrulax rufifrons slamatensis) was historically confined to Mount Slamet, where it has not been seen since 1925. A small captive population persisted up into the twenty-first century, although by 2018 only a single female remained. Mount Semeru Mount Semeru (Gunung Semeru in Indonesian) is an active volcano located in eastern Java (Jawa Timur province). Fruhstorfer’s mountain snake (Tetralepis fruhstorferi) is known only from Mount Semeru.

Lowland Rainforests On Java most forest below 1000 m, and in some areas up to 1500 m, has already been destroyed. Habitat destruction and degradation is driven primarily by agricultural expansion, logging, mining, development and uncontrolled fire, even within protected areas. The Ujung Kulon Peninsula, the westernmost tip of Java, has long contained the last remaining substantial areas of lowland rainforest on the island. The devastating 1883 eruption of the nearby island of Krakatoa and its resulting tsunami wiped out many settlements in the area and they were never repopulated. It was declared a national park in 1921 and harbours many highly threatened species. Outside Ujung Kulon only small fragments of lowland rainforest still exist. The Javan mastiff bat (Otomops formosus) is known only from four specimens collected in 1939 and in 1990 from western Java (Banten and Jawa Barat provinces). The Bantam caecilian (Ichthyophis hypocyaneus) was long known only from a single specimen collected in 1927 from western Java (Banten province). It was rediscovered in 2000 and is now known from three disjunct localities in Jawa Barat and Jawa Tengah provinces, and from some old museum specimens.

Lakes, Rivers, and Marshes Java is relatively poor in lakes and rivers, with rice fields serving as the most important wetland areas. The Javan lapwing (Vanellus macropterus) was a large, long-legged wading bird known for certain only from a few marshes and river deltas on the north-western and southeastern coasts of Java, although the species may have historically occurred on Sumatra, Timor and/or Belitung as well. On Java it appears always to have been rare. Last recorded in 1940, it is most likely extinct, although it is possible that it may survive in remote areas.

Bali The fabled island of Bali (Pulau Bali in Indonesian) lies just east of Java, on the western end of the Lesser Sunda Islands. Its central mountains include several peaks over 2000 m in elevation, the highest of which (Mount Agung) is an active volcano. At one time heavily forested, up until the early twentieth century it was home to a number of large mammals including the now extinct Bali tiger (see below), wild Javan bantengs (Bos javanicus javanicus), and Javan leopards (Panthera pardus melas). The banteng still occurs in its domestic form, but the leopard has long-since been extirpated. Up until the First World War the Bali tiger (Panthera tigris balica) was still rather common, but declined rapidly thereafter due to hunting and loss of habitat. The last definite record dates from 1937, when one was shot in the west of the island. Bali Barat National Park was established in 1941 for its protection, but it is likely that the subspecies became extinct by the end of World War II or possibly as late as the early 1950s. This was the smallest and least-known of the tigers, and was never photographed alive or kept in captivity. The Bali rusa deer (Rusa timorensis renschi) is confined to western Bali. The Bali myna (Leucopsar rothschildi) is now largely confined to West Bali National Park, with an additional small population on the offshore island of Nusa Penida (which is outside the species’ natural range). A beautiful and charismatic species; despite decades of conservation work it remains on the verge of extinction in the wild due mainly to unsustainable, illegal trapping for the global cage-bird trade. There is a considerable worldwide captive population. The grey-rumped myna (Acridotheres tertius) was historically found throughout Bali and the offshore islands of Nusa Penida and Nusa Lembongan; records from Lombok almost certainly represent either escapees from the cage-bird trade or vagrants. The species has undergone a significant decline in recent decades due to overcollection, and is now largely confined to Bali Barat National Park. The Bratan rasbora (Rasbora baliensis) is a type of freshwater fish that may or may not be confined to Lake Bratan, a crater lake located in north-eastern Bali.

Bawean Bawean (Pulau Bawean in Indonesian) is located approximately 150 km north of eastern Java. The island is hilly and was once covered by rainforest, although most of it was unfortunately replaced during the 1960s by teak plantations. Today, only five small pockets of forest remain, which is protected as the Bawean Nature Reserve. The Bawean deer (Axis kuhlii) is a small, relic form now confined to hilly areas of the island, although it appears to have once lived on Java as well. Reportedly common during the nineteenth century, it was almost exterminated by hunters during World War II. It recovered rapidly in the 1950s, only to find itself threatened once more in the 1960s and 1970s, this time by deforestation and persecution by locals, who regarded it as a nuisance for the damage it does to plantations. It is now

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fairly well protected, with a total population of between 250 and 500. The Bawean warty pig (Sus verrucosus blouchi) is confined to Bawean. The Bawean crested serpent eagle (Spilornis cheela baweanus) is confined to Bawean.

The Karimunjawa Islands The Karimunjawa Islands (Kepulauan Karimunjawa in Indonesian) are located north of Java. The Karimunjawa long-tailed macaque (Macaca fascicularis karimondjawae) is confined to the Karimunjawa Islands.

The Kangean Islands The Kangean Islands (Kepulauan Kangean in Indonesian) are located east of Madura in the Java Sea and consist of some 28 small islets, many of which are still uninhabited. The Kangean tit-babbler (Mixornis prillwitzi) is confined to remaining areas of lowland forest in the Kangean Islands.

The Masalembu Islands The Masalembu Islands (Kepulauan Mesalembu in Indonesian) are located in the Java Sea to the north of Madura. The three main islands are Masalembu, Masakambing, and Karamian. Abbott’s yellow-crested cockatoo (Cacatua sulphurea abbotti) is confined to the Masalembu Islands, where it has long been on the verge of extinction. The subspecies appears to have been extirpated from Masalembu in 1987 due to trapping and killing, and only about a dozen individuals were recorded on Masakambing in 2010.

Christmas Island Located south-west of Java, Christmas Island was uninhabited until the late nineteenth century. The island’s geographic isolation and history of minimal human disturbance had led to high levels of endemism. Large areas of monsoonal forest remain and much of the island has been set aside as a national park, although a wide range of introduced species have unfortunately decimated the native fauna. Two large rats (Rattus), historically endemic to Christmas Island, have been exterminated. The bulldog rat (R. nativitatis) has not been recorded with certainty since 1897–98. Maclear’s rat (R. macleari) was last observed in 1904, and extensive searches since then have failed to locate it. Both were likely wiped out by competition with introduced black rats (R. rattus) and disease. The Christmas Island shrew (Crocidura trichura) was described as ‘extremely common’ in 1900, but was already rare in 1909. Since then, there have been only four confirmed records (two individuals in 1958, and another two in 1985), although several unconfirmed reports occurred between 1996 and 1998. There has been nothing since despite intensive survey work, and the species is very likely extinct. The Christmas Island flying fox (Pteropus melanotus natalis) is confined to the island, where it has undergone significant declines in recent years.

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The Christmas Island pipistrelle (Pipistrellus murrayi) was a type of bat historically widespread and common on the island, but it underwent a dramatic decline most likely due to introduced species. Last recorded in 2009, it is now thought to be extinct. Abbott’s booby (Papasula abbotti), the largest of all booby species, was historically widespread in the Indian Ocean and probably the Pacific as well. Its only known breeding sites, however, were on Assumption Island in the Seychelles (where it was driven extinct by humans around 1936) and on Christmas Island, where it nests in tall emergent trees of the plateau rainforest. Today the population of around 6000 is highly threatened by logging activities. The Christmas Island frigatebird (Fregata andrewsi) is a type of seabird that breeds only on Christmas Island in four main colonies. The Christmas Island goshawk (Accipiter fasciatus natalis) was reported to be common all over the island in 1947, but by 1963–64 its numbers had dropped to a low of about 12. The population has since increased to about 100. The Christmas Island hawk-owl (Ninox natalis) is confined to the island, where the population is thought to be stable. The Christmas Island thrush (Turdus poliocephalus erythropleurus) is historically, and currently, endemic to Christmas Island. During the late nineteenth century it was introduced into the Cocos (Keeling) Islands, but was extirpated from there by the 1980s. The Christmas Island chained gecko (Lepidodactylus listeri) underwent a massive decline in the late twentieth century due to introduced species, and was last recorded in the wild in 2012. Fortunately, a captive breeding programme has been successful. The Christmas Island emo skink (Emoia nativitatis) was historically widespread on the island and considered common in 1979. By the 1990s, however, the population plummeted drastically, most likely due to introduced species, and was soon reduced to a few fragmented pockets. A projected captive breeding programme was organized too late, and the last known individual died in 2014. The species is now considered to be extinct. The Christmas Island blind snake (Ramphotyphlops exocoeti) is known only from a few localities on the island.

The Cocos (Keeling) Islands The Cocos (Keeling) Islands, located approximately midway between Australia and Sri Lanka, consist of two flat, low-lying coral atolls and 27 coral islands. The Cocos buff-banded rail (Gallirallus philippensis andrewsi) is now effectively confined to North Keeling Island, although it is still occasionally reported from other islands making up the Southern Atoll. The population is believed to number less than 1000. The Cocos Islands reed snake (Calamaria lautensis) is known only from three specimens collected on Cocos Island.

Balance for the Sundaic Region Fossilized remains of Homo erectus, popularly known as ‘Java Man’, suggest that the Sundaic Region was inhabited some 1.5

The Sundaic Region

million years ago. Another species, H. floresiensis, also lived in the area up until at least 50,000 years ago, after which it became extinct. Modern humans (H. sapiens) first reached the region between 70,000 and 50,000 B . C ., as evidenced by rock art and a skull found within the Niah Caves in northern Borneo. These early migrants would be joined by Austronesian peoples from Taiwan around 2000 B . C . During much of this period the present-day islands of western Indonesia were joined into a single landmass known as Sundaland due to lower sea levels. Ideal agricultural conditions and the mastering of wet-field rice cultivation as early as the eighth century B . C . allowed villages, towns and small kingdoms to flourish by the first century A . D ., and the archipelago’s strategic sea-lane position fostered trade as far away as India and China. Early European exploration of the region was undertaken by Portuguese mariners. In 1509 Diogo Lopes de Sequeira reached Sumatra and what is now the state of Malacca, on the southern coast of the Malay Peninsula. In 1511–12 Antonio de Abreu sailed through the Strait of Malacca between Sumatra and Bangka, and along the coasts of Java, Bali, Lombok, Sumbawa, and Flores to the ‘Spice Islands’ (i.e. the Moluccas). In 1518 Lourenço Gomes reached Borneo. In the following centuries Europeans would establish colonies, with the British on the Malay Peninsula, the Dutch in Indonesia and, for a brief period, the Japanese throughout during World

War II. Despite the establishment of large plantations human impact remained minimal. Sadly, the post-colonization period which began during the latter half of the twentieth century has been nothing short of disastrous for the wildlife of this region. A rapid and unsustainable population increase and subsequent spreading of agriculture and shifting cultivation had already begun to significantly decrease the rainforests of Indonesia and Malaysia, but it was not until 1966, when the forests were opened up to logging companies, that the cataclysmic damage truly began. Singapore, in particular, served as the nexus for a vast international trade in wildlife. In recent historical time (i.e. since A . D . 1500), the Sundaic Region has lost at least 12 species/3 subspecies of vertebrates. Among the extinct forms 5 species/2 subspecies are mammals, 2 species/1 subspecies are birds, 4 species are reptiles, and 1 species is an amphibian. Another 4 species are possibly extinct, and 2 species are currently extinct in the wild. In addition, there are 406 species/67 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 134 species/54 subspecies are mammals, 65 species/8 subspecies are birds, 70 species/5 subspecies are reptiles, 84 species are amphibians, and 53 species are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Mammals

5 species

~ species

~ species

134 species

2 subspecies

~ subspecies

~ subspecies

54 subspecies

7 taxa

~ taxa

~ taxa

188 taxa

2 species

~ species

1 species

65 species

1 subspecies

~ subspecies

~ subspecies

8 subspecies

3 taxa

~ taxa

1 taxon

73 taxa

4 species

1 species

1 species

70 species

~ subspecies

~ subspecies

~ subspecies

5 subspecies

4 taxa

1 taxon

1 taxon

75 taxa

1 species

1 species

~ species

84 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

1 taxon

1 taxon

~ taxa

84 taxa

~ species

2 species

~ species

53 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

2 taxa

~ taxa

53 taxa

12 species

4 species

2 species

406 species

3 subspecies

~ subspecies

~ subspecies

67 subspecies

15 taxa

4 taxa

2 taxa

472 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Threatened

Note: ~, not applicable.

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The Wallacean Region The Wallacean Region comprises a group of mainly Indonesian islands that includes the Philippines, Sulawesi, all of the Lesser Sunda Islands east of Bali, the Moluccas Archipelago, and many smaller islands. Essentially a transition zone between the Sundaic Region on the one hand and the Papua-Melanesian and Australian realms on the other, it consequently shares biogeographic elements of each in addition to many unique features of its own. It is named for Alfred Russel Wallace, the first to the first to draw a physical line between Asian and Australasian biogeographic regions. The fauna of Indonesia and Australasia was, for Wallace, what that of the Galápagos was for Charles Darwin: one of the foundations of the theory of natural selection. One hundred and fifty years ago he wrote that South America and Africa, although separated by the Atlantic, do not differ so much by far as do Asia and Australia. Modern humans are known to have lived in the Wallacean Region for tens of thousands of years, having first arrived from southern Asia by way of the sunken landmasses of Sundaland and Sahul. They were eventually assimilated by the Austronesian peoples who arrived by seaborne migration between 3000 and 1500 B . C ., most likely from Taiwan. The small Sulawesi cuscus (Strigocuscus celebensis) is a type of arboreal marsupial that remains widespread on Sulawesi, Wowoni, Buton, and the islands of Great Sangir and Siau in the Sangir Islands, but is everywhere threatened by hunting and habitat destruction. The Sulawesi giant squirrel (Rubrisciurus rubriventer) is widespread on Sulawesi and on the island of Great Sangir, where it is threatened by hunting. The grey flying fox (Pteropus griseus) is found on Sulawesi and its satellite islands, the Moluccas, and possibly on Flores in the Lesser Sunda Islands. It is threatened by bushmeat hunting on Sulawesi and by loss of habitat throughout its range. Keast’s tube-nosed fruit bat (Nyctimene keasti) is known for certain from the Tanimbar and Babar islands in the Moluccas, the Kai Islands in the Lesser Sunda Islands, and from Gag Island in the Raja Ampat Islands. It may also occur on Timor, Flores and other islands as well. The Sulawesi fruit bat (Acerodon celebensis) is found across Sulawesi as well as on Buton, the Togean Islands, Selayar Island, and the Sangir Islands, but is everywhere threatened by hunting for food. Many colonies have been extirpated, and the species remains unprotected. Peters’ horseshoe bat (Rhinolophus keyensis) is known sporadically from the Lesser Sunda Islands and the Moluccas. The Indonesian tomb bat (Taphozous achates) is known sporadically from the Lesser Sunda Islands and the Moluccas. Bernstein’s scrubfowl (Megapodius bernsteinii) is confined to the Banggai and Sula Islands east of Sulawesi, where it has suffered significant declines due to loss of habitat and hunting pressure. The Wallacean box turtle (Cuora amboinensis amobinensis) is found throughout Sulawesi, the Moluccas, the Lesser Sunda

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Islands and possibly the Philippines, but is everywhere threatened by overcollection. The Sulawesi black rat snake (Ptyas dipsas) is a littleknown species from an undefined area of Sulawesi and, possibly, the island of Halmahera in the Moluccas.

The Philippines The Philippines (Pilipinas in Filipino) are an archipelago of over 7600 islands located north-east of Borneo and south of Taiwan. The islands are volcanic in origin and were originally covered by lush rainforest inhabited by a fascinating range of endemic species. At the beginning of the twentieth century this was largely intact, although since the period of World War II almost all forest has been destroyed and replaced by cultivation. Hunting is also a major problem to the island’s wildlife. Both of these factors must, of course, be seen in the light of an astonishing ongoing population increase, which has already risen to well over 100 million people. The Philippine long-tailed macaque (Macaca fascicularis philippinensis) is still found throughout the Philippines, but is everywhere threatened by hunting and habitat destruction. The Philippine tarsier (Carlito syrichta) is a type of nocturnal primate found in the central and southern Philippines. The nominate form (C. s. syrichta) is found on Leyte, Samar, Dinagat, Biliran, and Maripipi in the south-eastern Philippines. The Mindanao tarsier (C. s. carbonarius) occurs patchily on Mindanao and on Basilan in the Sulu Archipelago. Both are threatened by loss of habitat and collection for use as food or pets. The Philippine brown deer (Rusa marianna) still occurs on many islands (with introduced populations occurring as well in the Mariana and Caroline Islands), but is threatened by intense hunting pressure and habitat destruction. There are four subspecies, which will be discussed below. The nominate form, the Luzon brown deer (R. m. marianna), occurs on Luzon and other islands of the northern Philippines. The Philippine spotted deer (R. alfredi) is widespread but rare and patchily distributed in the Western Visayan islands of the central Philippines. Historically it was found on Panay, Negros, Cebu, Masbate, Guimaras and probably the Ticao Islands, but today it is confined to the mountains of western Panay and in a few scattered remnants of forest on Negros. Several species and subspecies of wild boar (Sus) are endemic to the Philippines. The Philippine warty pig (S. philippensis) is still found in scattered pockets across the Philippines. Nevertheless, it continues to be threatened by hunting persecution by farmers (who consider it a pest) and habitat destruction. The North Philippine warty pig (S. p. philippensis) occurs on Luzon, Polillo, Catanduanes and, formerly, Marinduque. The South Philippine warty pig (Sus p. mindanensis) is found on Mindanao, Samar, Letye, Biliran, Bohol, Camiguin Sul, Basilan, and a few associated smaller islands. The Visayan warty pig (S. cebifrons) is native to the Western Visayan islands in the central Philippines, but has

The Wallacean Region

been extirpated from at least three of the six islands where it was historically known to occur, including Cebu. The Negros warty pig (S. c. negrinus) survives in remaining forest fragments on Negros, Panay, and possibly Masbate. The Visayan leopard cat (Prionailurus bengalensis rabori) is confined to Panay, Negros and Cebu, where it is threatened by loss of habitat. The Philippine binturong (Arctictis binturong whitei) is a large viverrid known from the islands of Calauit and Palawan in the south-western Philippines. The golden-crowned fruit bat (Acerodon jubatus) is divided into three subspecies. The nominate form (A. j. jubatus) is still found widely throughout the Philippines but is everywhere threatened by bushmeat hunting. The Philippine naked-backed fruit bat (Dobsonia chapmani) is historically known from the islands of Cebu and Negros in the south-central Philippines, where it was last recorded in 2003. It may possibly be extinct, a victim of habitat destruction and hunting. The Philippine tube-nosed fruit bat (Nyctimene rabori) is found on the islands of Cebu, Negros, Sibuyan, and possibly Panay in the south-central Philippines, where it is threatened by habitat destruction. The Philippine large leaf-nosed bat (Hipposideros coronatus) is known only from a handful of specimens collected throughout the Philippines. The Philippine eagle (Pithecophaga jefferyi) is the largest eagle in the world, having a wingspan of almost 2.5 m. It is also one of the most threatened species in the Philippines, occurring only in eastern Luzon, Samar, Leyte, and Mindanao, with Mount Apo National Park on the latter island supporting the majority of the population. It has long been a favourite target for trophy hunters.

Two species of hawk-eagle (Nisaetus) endemic to the Philippines are threatened by loss of habitat and hunting. The northern Philippine hawk-eagle (N. philippensis) is now confined to a few localities on Luzon and Mindoro, having been extirpated from a number of smaller islands. Pinsker’s hawk-eagle (N. pinskeri) remains relatively widespread if rare across the southern and eastern Philippines. The Philippine eagle-owl (Bubo philippensis) is a widespread but naturally rare species divided into two subspecies. The northern Philippine eagle-owl (B. p. philippensis) is found on Luzon and Catanduanes. The southern Philippine eagle-owl (B. p. mindanensis) occurs on Mindanao, Leyte, Samar and Bohol. Both are threatened by habitat destruction and hunting. The giant scops owl (Otus gurneyi) is confined to the islands of Samar, Mindanao, Dinagat, and Siargao in the south-eastern Philippines. The Visayan scops owl (O. nigrorum) is confined to Negros and Panay in the west-central Philippines. Both are threatened by habitat destruction. The northern rufous hornbill (Buceros hydrocorax) is confined to Luzon and, marginally, Marinduque. The southern rufous hornbill (B. mindanensis) is divided into two subspecies. The nominate form (B. m. mindanensis) is found on Dinagat, Siargao, Mindanao, Balut, Bucas, and Talicud. The yellow-billed southern rufous hornbill (B. m. semigaleatus) occurs on Samar, Leyte, Bohol, Panaon, Biliran, Calicoan, and Buad. All are threatened by habitat destruction and hunting. The Visayan hornbill (Penelopides panini) is divided into two subspecies, one of which is now extinct and will be discussed below. The nominate form (P. p. panini) is found on Panay, Negros, Masbate and Guimaras, where it is seriously threatened by habitat destruction and hunting.

Figure 5.7 Illustration of a Philippine eagle kept in London Zoo in 1909–10. (Credit: Henrik Grönvold.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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The rufous-headed hornbill (Rhabdotorrhinus waldeni) is historically thought to have occurred on Negros, Panay and Guimaras in the west-central Philippines. Owing to habitat destruction, hunting and nest poaching, it is now absent from Guimaras, and is now largely confined to protected areas on the other two islands. The Philippine cockatoo (Cacatua haematuropygia) was described as common throughout the Philippines during the mid-twentieth century, but has since undergone a catastrophic decline due to habitat destruction and trapping for the cagebird trade. Today, only around 1000 birds survive on Palawan and its satellite islands, Bohol, Samar and in the Sulu Archipelago. It has possibly been extirpated from Mindanao. The Philippine duck (Anas luzonica) has historically been recorded from all of the major islands of the Philippines and a number of smaller ones, but seems to be mainly found on Luzon and Mindanao. The species appears to have undergone a major decline in the mid-twentieth century, however, and today is found only in a few scattered freshwater and mangrove areas. The spotted imperial pigeon (Ducula carola) is divided into three subspecies, which are threatened by habitat destruction and hunting. The nominate form (D. c. carola) is found on Luzon, Mindoro and Sibuyan. The Visayan spotted imperial pigeon (D. c. nigorum) is confined to Negros and Siquijor. The dark-eared brown dove (Phapitreron brunneiceps) is a rare species known from Mindanao and Basilan in the southern Philippines, although it has not been recorded from the latter island since 1937. The Philippine collared dove (Streptopelia dusumieri) remains widespread throughout the Philippines, but is everywhere in decline owing to competition with two species of non-native dove. The species has also been introduced into the Mariana Islands and parts of eastern Malaysia. Keay’s bleeding-heart (Gallicolumba keayi) is a rare type of ground dove confined to a few localities on Negros and Panay in the west-central Philippines. Reichenbach’s bleeding-heart (G. crinigera) is divided into two subspecies. The nominate form (G. c. crinigera) is known from Samar, Leyte, Bohol, Dinagat and Mindanao, where it is threatened by loss of habitat. Whitehead’s swiftlet (Aerodramus whiteheadi) is found disjunctly in the mountains of the northern and southern Philippines, perhaps suggesting that more than one species may be involved. The southern Whitehead’s swiftlet (A. w. origenis) is known only from a few scattered records on Mindanao and Negros. The southern sooty woodpecker (Mulleripicus fuliginosus) is confined to lowland forests on Mindanao, Leyte and Samar in the south-eastern Philippines. The yellow-faced flameback (Chrysocolaptes xanthocephalus) is a type of woodpecker known from Negros and Panay, and at least formerly also on Guimaras, Masbate and Ticao in the central Philippines. The red-headed flameback (C. erythrocephalus) is found on Balabac, Palawan, Busuanga, and

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Calamian in the eastern Philippines. Both are threatened by loss of habitat. Steer’s wattled broadbill (Sarcophanops steerii) is a type of passerine bird that was historically widespread and common on Mindanao, Basilan, Dinagat and Siargao in the southern Philippines. Owing to extensive habitat destruction it is now confined to a handful of localities. The Visayan wattled broadbill (S. samarensis) is confined to Samar, Leyte and Bohol in the east-central Philippines, where it has only rarely been recorded since 1980. The Visayan flowerpecker (Dicaeum haematostictum) was historically widespread and common on Panay, Negros and Guimaras in the west-central Philippines. The species has undergone a significant decline due to loss of habitat, and appears to have been extirpated from Guimaras. The scarletcollared flowerpecker (D. retrocinctum) was long thought to be confined to Mindoro, where it was abundant up until the 1980s, but has since declined significantly. In the early 1990s it was unexpectedly discovered in small numbers on Panay and Negros. The streak-breasted bulbul (Hypsipetes siquijorensis) is a type of passerine bird divided into three subspecies, which will be discussed below. The rufous-lored kingfisher (Todiramphus winchelli) remains fairly widespread within the smaller islands and coastal lowland forests of the Philippines, but appears to be highly sensitive to habitat destruction. The Philippine dwarf kingfisher (Ceyx melanurus) is divided into three subspecies. The northern Philippine dwarf kingfisher (C. m. melanurus) is found on Luzon, Polillo, Alabat, and Catanduanes. The central Philippine dwarf kingfisher (C. m. samarensis) occurs on Samar and Leyte. The southern Philippine dwarf kingfisher (C. m. mindanensis) is confined to Mindanao and Basilan. All are threatened by loss of habitat. The ashy ground thrush (Geokichla cinerea) is confined to parts of Luzon and Mindoro in the northern Philippines. The Philippine leafbird (Chloropsis flavipennis) is historically known from Samar, Leyte, Cebu and Mindanao in the south-central Philippines. It was last recorded on Cebu in 1920, from Leyte in 1964, and from Samar in 1970, and now appears to be entirely confined to two localities on Mindanao. The Visayan miniature tit-babbler (Micromacronus leytensis) is confined to Samar, Biliran and Leyte in the southeastern Philippines. The flame-templed pygmy babbler (Dasycrotapha speciosa) is confined to a few localities on Panay and Negros in the west-central Philippines. The Visayan pygmy babbler (D. pygmaea) is confined to Samar and Leyte in the south-eastern Philippines, where it is rare. The white-throated jungle flycatcher (Vauriella albigularis) was historically known from Negros, Guimaras and Panay in the west-central Philippines. It was last recorded on Guimaras in 1887 and has presumably been extirpated from there, and has been much reduced on the other two islands due to habitat destruction.

The Wallacean Region

The ashy-breasted flycatcher (Muscicapa randi) is a rare species known disjunctly from Luzon, Negros and Samar. The little slaty flycatcher (Ficedula basilanica) is known from the islands of Mindanao, Samar, Leyte, Dinagat and Basilan. Formerly common, it now appears to be rare and largely confined to a few localities on Mindanao. The azure-breasted pitta (Pitta steerii) is a passerine bird known from the islands of Mindanao, Bohol, Leyte and Samar in the south-eastern Philippines. Seemingly always rare and localized, since 1980 it has only been recorded at two sites on Bohol and one on Mindanao. It was last recorded on Leyte in 1964, and from Samar in 1969. The celestial monarch (Hypothymis coelestis) is a passerine bird that has always been considered somewhat rare. The nominate form (H. c. coelestis) is known from Luzon, Samar, Dinagat, Mindanao, Tawi-Tawi, and Basilan, although it appears to have been extirpated from the latter island. The Visayan celestial monarch (H. c. rabori) was historically known from Sibuyan and Negros in the south-central Philippines. Last recorded in 1959, it may possibly be extinct. The bicoloured water-redstart (Phoenicurus bicolor) was historically known from Mindoro and Luzon in the northwestern Philippines, but is now confined to the mountains of the latter. The green-faced parrotfinch (Erythrura viridifacies) is confined to Luzon, Negros and Panay in the northern and central Philippines. The white-winged cicadabird (Edolisoma ostentum) is a type of cuckooshrike endemic to the Western Visayas, where it is known from Panay, Negros and Guimaras (although it is almost certainly extirpated from the latter island). It was formerly widespread on Negros as well but is now confined to a few localities. The black-bibbed cicadabird (E. mindanense) is divided into a number of subspecies found throughout the Philippines. The Mindanao black-bibbed cicadabird (E. m. mindanense) is confined to Mindanao and Basilan in the southern Philippines. Ripley’s black-bibbed cicadabird (E. m. ripleyi) is found on Samar, Biliran, Leyte and Bohol. Everett’s black-bibbed cicadabird (E. m. everetti) occurs on Jolo, Lapac, Tawitawi, and Bongao in the south-western Philippines. All are threatened by loss of habitat. The Visayan rhabdornis (Rhabdornis rabori) is a rare type of bird known only from a few forest patches on Panay and Negros in the west-central Philippines. The Philippine crocodile (Crocodylus mindorensis) is a relatively small freshwater species that historically occurred on all of the larger islands. A victim of relentless exploitation combined with unsustainable fishing methods (such as dynamite fishing), it is today known for certain only from a few areas of Luzon and Mindanao. The Philippine sailfin lizard (Hydrosaurus pustulatus) is a relatively large agama named for the sail-like structure of its tail and semi-aquatic habits. It occurs in suitable habitat throughout the Philippines, where it is threatened by habitat destruction and overcollection for the international pet trade.

The marbled crested lizard (Bronchocela marmorata) is known for certain only from Luzon and Sibuyan in the northern Philippines. The Philippine gliding dragon (Draco mindanensis) is confined to Mindanao, Leyte, Dinagat and Samar in the southern and eastern Philippines. The orange-spotted smooth-scaled gecko (Pseudogekko brevipes) is known from a few scattered localities across the islands of the central Philippines (Samar, Negros, Bohol, Siquijor, Cebu, Leyte, and Panay). Griffin’s keel-scaled tree skink (Dasia griffini) is known only from a few specimens collected from Palawan and Mindoro in the west-central Philippines. Semper’s skink (Lipinia semperi) is known only from historical records originating in Mindanao, Camiguin, and possibly Samar. Girard’s skink (L. vulcania) is known only from two specimens, the first collected from Mindanao in 1857 and the second from Luzon in 1956. The slender four-fingered burrowing skink (Brachymeles elerae) is a rare species with an undefined distribution within the Philippines. The spotted-bellied kukri snake (Oligodon modestum) is known from north-eastern Mindanao, Negros, Tablas, Panay, Cebu, and possibly Luzon. The Philippine bridal snake (Lycodon philippinus) was historically known from a number of islands, but in recent years has been reported only from northern Luzon, Marinduque, and Panay. Taylor’s keelback (Tropidonophis negrosensis) is a type of snake known sporadically from Cebu, Masbate, Negros, Panay, Basilan, and Mindanao. It may also occur on Luzon, Samar, and Lubang. The Philippine whip snake (Dryophiops philippina) was historically common throughout the Philippines, but has undergone a significant decline and is now rarely recorded. McNamara’s burrowing snake (Pseudorabdion mcnamarae) is known only from Negros, Panay, Masbate, Cebu, and Romblon in the west-central Philippines. The Cabilian blood frog (Sanguirana albotuberculata) is known only from a few localities on Leyte, Samar, and Mindanao in the south-eastern Philippines. The Polillo ground frog (Platymantis polillensis) is found patchily on Luzon and Polillo in the northern Philippines. Lawton’s wrinkled ground frog (P. lawtoni) is confined to Tablas and Sibuyan in the west-central Philippines. The Negros wrinkled ground frog (P. negrosensis) is confined to a few localities on Panay and Negros in the west-central Philippines. All are threatened by loss of habitat. The small-headed frog (Occidozyga diminutiva) is confined to western Mindanao, Basilan, and the Sulu Archipelago in the southern Philippines. The Visayan wart frog (Limnonectes visayanus) is found patchily on Masbate, Cebu, Negros, Guimaras, Panay and Siquijor in the central Philippines. The small-disked wart frog

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(L. parvus) is confined to Mindanao and Basilan in the southern Philippines. The Abungabung caecilian (Ichthyophis glandulosus) is known only from western Mindanao and Basilan in the southern Philippines. The spotted Hampala barb (Hampala lopezi) is a littleknown species endemic to the Philippines.

The Batanes Islands The Batanes Islands are located north of Luzon. They include Batan, Itbayat, Sabtang, and a number of uninhabited islets. The Batanes pit viper (Trimeresurus mcgregori) is known only from Batan and Sabtang.

Luzon Luzon is located in the northern Philippines, and is the largest and by far the most populous of the islands. Most of its original lowland rainforests were destroyed during the twentieth century, and what little remains in the higher elevations continues to be threatened by both legal and illegal logging. The southern Luzon cloud rat (Phloeomys cumingi) is found in southern Luzon and on the satellite islands of Catanduanes and Marinduque, where it is threatened mainly by hunting. The Luzon broad-toothed rat (Abditomys latidens) is confined to central and northern Luzon (Laguna and Mountain provinces). The Luzon short-nosed rat (Tryphomys adustus) is confined to a few localities in central and northern Luzon (Benguet, Laguna, and Tarlac provinces). The Luzon shrew-mouse (Crunomys fallax) is known only from a single specimen collected in the late nineteenth century from north-eastern Luzon. The brown-banded rail (Lewinia mirifica) is known only from Luzon, where it is subjected to heavy hunting. The green racquet-tail (Prioniturus luconensis) is a type of parrot that was historically widespread and common on Luzon and Marinduque. It has declined dramatically in recent decades due to habitat destruction and overcollection for the cage-bird trade, and is now largely confined to the Subic Bay Forest Reserve in western Luzon. The flame-breasted fruit dove (Ramphiculus marchei) is confined to Luzon, where it is threatened by loss of habitat and hunting. The Isabela oriole (Oriolus isabellae) was thought to be extinct until its rediscovery in 1993. Further populations have since been found in northern and eastern Luzon, but the species remains highly threatened by loss of habitat. The rufous-breasted blue flycatcher (Cyornis camarinensis) is confined to southern Luzon and Catanduanes Island, where it is threatened by habitat destruction. The Luzon black-bibbed cicadabird (Edolisoma mindanense lecroyae) is confined to Luzon, where it is very rare. The Luzon Whitehead’s swiftlet (Aerodramus whiteheadi whiteheadi) is confined to the mountains of Luzon.

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The white-browed jungle flycatcher (Vauriella insignis) is confined to the mountains of northern Luzon. Rabor’s ground warbler (Robsonius rabori) is known only from the mountains of north-western Luzon. Gray’s monitor (Varanus olivaceus) is confined to eastern Luzon and the satellite islands of Catanduanes and Polillo. It is threatened by habitat destruction, hunting, and overcollection for the international pet trade. Cuming’s fringed gecko (Luperosaurus cumingii) is confined to southern Luzon and the satellite island of Polillo. The San Roque skink (Parvoscincus laterimaculatus) is known only from a single specimen collected in 1961 from south-eastern Luzon (Sorsogon). Numerous surveys since have failed to detect the species, and it is most likely extinct. Knollman’s skink (Sphenomorphus knollmanae) is known from five scattered localities on Luzon and the island of Marinduque. The Manila wolf snake (Lycodon tessellatus) is known only from a single museum specimen collected in 1846 and labelled ‘Manila’. The Luzon wolf snake (L. solivagus) is known only from two localities in northern Luzon. The Manila blind snake (Malayotyphlops manilae) is known only from its original collection in the early twentieth century, most likely on Luzon. The light-collared blind snake (M. collaris) is known only from a single specimen collected in the early twentieth century from south-eastern Luzon. Moodie’s frog (Fejervarya moodiei) was a presumably aquatic species known only from a single specimen collected during the early twentieth century from what is now the city of Manila. It is most likely extinct. Taylor’s blood frog (Sanguirana igorota) is confined to the mountains of north-western Luzon, where it is threatened by habitat destruction and pollution. The montane wrinkled ground frog (Platymantis montanus) is confined to a few localities in south-central Luzon. The Cagayan wrinkled ground frog (P. cagayanensis) is confined to the northern coast of Luzon and the adjacent islet of Palaui. The Isarog wrinkled ground frog (P. isarog) is confined to Mount Isarog and Mount Malinao in south-eastern Luzon. The horned wrinkled ground frog (P. cornutus) is found patchily in northern Luzon. Taylor’s wrinked ground frog (P. taylori) is confined to north-eastern Luzon. All are threatened by loss of habitat. Walter’s narrow-mouthed frog (Kaloula walteri) is known from a few localities in south-eastern Luzon and the satellite island of Polillo, where it is threatened by loss of habitat. The dwarf freshwater goby (Gobiopterus stellatus) is a little-known species confined to Luzon. The Cordillera Central The Cordillera Central is a large mountain range located in north-central Luzon. The Luzon giant bushy-tailed cloud rat (Crateromys schadenbergi) is known only from the southern and central Cordillera Central.

The Wallacean Region

The short-footed Luzon tree rat (Carpomys melanurus) was long known only from a single specimen collected from Mount Data in the late nineteenth century. A possible second specimen was collected in 2008. The large-toothed hairy-tailed rat (Batomys dentatus) is known only from a single specimen collected in the early twentieth century from the Cordillera Central. The Luzon buttonquail (Turnix worcesteri) is known only from a few localities. Lawton’s skink (Sphenomorphus lawtoni) is known only from two specimens collected in the 1920s from the mountains of north-western Luzon. Wright’s burrowing skink (Brachymeles wright) is known only from two specimens collected in the 1920s. Barbour’s keelback (Rhabdophis barbouri) is a type of water snake known only from two specimens collected in 1917. Myer’s snake (Myersophis alpestris) is known only from two specimens collected in the 1960s from the mountains of north-central Luzon. The Luzon blind snake (Acutotyphlops banaorum) is known only from a single locality in north-western Luzon. The Cordillera Central wrinked ground frog (Platymantis subterrestris) is confined to the Cordillera Central, where it is threatened by loss of habitat. The Sierra Madre Mountains The Sierra Madre Mountains are located in north-eastern Luzon (Cagayan and Quezon provinces). Running in a north–south direction, they form the eastern backbone of the island. The Lataan fringed gecko (Luperosaurus kubli) is known only from Mount Lataan in the Sierra Madre Mountains. The Sierra Madre blood frog (Sanguirana tipanan) is confined to a few localities within the Sierra Madre Mountains. The Sierra Madre wrinkled ground frog (Platymantis sierramadrensis) is known for certain only from the Sierra Madre Mountains, but may range more widely. The Zambales Mountains The Zambales Mountains are located in north-western Luzon (Zambales province). They include Mount Pinatubo, Mount Negron, Mount Cuadrado, and Mount Tapulao. The Tapulao shrew-rat (Rhynchomys tapulao) is known only from Mount Tapulao. Mount Banahao Mount Banahao is a potentially active, three-peaked stratovolcano located in southern Luzon (Laguna and Quezon provinces). Naomi’s wrinkled ground frog (Platymantis naomii), the indiscoverable wrinkled ground frog (P. indeprensus), the false-backed wrinkled ground frog (P. pseudodorsalis), and the Banahao wrinkled ground frog (P. banahao) are endemic to Mount Banahao.

Mount Isarog Mount Isarog is a potentially active stratovolcano located in south-eastern Luzon (Camerines Sur province). It is nominally protected within Mount Isarog National Park. The Isarog shrew-rat (Rhynchomys isarogensis) is known only from Mount Isarog. The Isarog shrew-mouse (Archboldomys luzonensis) is known only from Mount Isarog. Malinao Volcano Malinao Volcano is a potentially active stratovolcano located in southern Luzon (Albay and Camarines Sur provinces). The Malinao wrinkled ground frog (Platymantis diesmosi) is confined to Malinao Volcano. Lake Taal Lake Taal (Lawa ng Taal in Tagalog, formerly known as Bombón Lake) is located in southern coastal Luzon (Batangas province). Originally an inlet of nearby Balayan Bay, a series of major eruptions during the eighteenth century combined with centuries of rain have converted it into a freshwater crater lake. Further eruptions have occurred since then, most recently in 2020. Semper’s sea snake (Hydrophis semperi), one of only two freshwater sea snakes in the world, is entirely confined to Lake Taal. The Taal herring (Sardinella tawilis) is a type of sardine that has evolved to live in freshwater, and is entirely confined to Lake Taal. It is threatened by overfishing, pollution, and competition from introduced fish species. Lake Bato Lake Bato is located in south-eastern Luzon (Camerines Sur province). The Bato pipefish (Microphis pleurostictus) is confined to Lake Bato as well as Yassot Creek and possibly other tributaries. Lake Buhi Lake Buhi is located in south-eastern Luzon (Camerines Sur province). The Buhi dog-faced water snake (Cerberus microlepis) is known only from Lake Buhi, where it was last recorded in the early 1990s. The Babuyan Islands The Babuyan Islands are a group of five main islands (Babuyan, Calayan, Dalupiri, Fuga, and Camiguin) along with their smaller satellites located north of Luzon. McGregor’s fringed gecko (Luperosaurus macgregori) is known only from the islands of Calayan, Barit, and Babuyan Claro. Ross’ wolf snake (Lycodon chrysoprateros) is only definitely known from three specimens collected in 1990. It may also be found on Calayan and Camiguin Norte Islands, but this has yet to be confirmed.

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Calayan is the largest of the Babuyan Islands. The Calayan rail (Gallirallus calayanensis) is confined to Calayan Island.

The Romblon Islands The Romblon Islands are located south of Luzon in the central Philippines. The Romblon hawk-owl (Ninox spilonotus) is known with certainty only from the islands of Sibuyan and Tablas, although there are unconfirmed reports from Romblon Island itself. The Romblon streak-breasted bulbul (Hypsipetes siquijorensis cinereiceps) is confined to Tablas and Rombon. The Romblon wrinkled ground frog (Platymantis levigatus) is confined to Romblon, Tablas, and Sibuyan. Tablas Tablas is the largest of the Romblon Islands. The Tablas drongo (Dicrurus menagei) is a type of bird confined to Tablas, where it was thought to have gone extinct by the end of the twentieth century. However, observations since that time have confirmed that it still survives, although in very small numbers. The Tablas fantail (Rhipidura sauli) is a type of passerine bird confined to mature forests on Tablas. Sibuyan Sibuyan is the second largest of the Romblon Islands. Owing to its geological formation it has never been connected with the rest of the Philippines, and while most of its lowland forests have been destroyed much remains in the higher elevations. The Sibuyan striped shrew-rat (Chrotomys sibuyanensis) is known only from a single specimen collected from Sibuyan in 1992.

Mindoro Mindoro is located in the west-central Philippines off the south-western coast of Luzon. Once covered with lush rainforests, most has already been destroyed by mining, agriculture and logging, with the remainder expected to disappear by 2030. The Siburan Forest/Sablayan Penal Colony holds the largest remaining area, and as such will be critically important for the survival of the island’s impressive number of endemic species. The tamaraw (Bubalus mindorensis) is a dwarf species of wild buffalo long found only on Mindoro, although it appears to have been historically present on Luzon as well and perhaps other islands. First recorded by Western science in 1888, the island’s problems with a particularly virulent strain of malaria deterred human settlers, allowing the species to roam at will. However, as the human population increased the tamaraw’s range shrank to just three known areas of dense montane forest with open glades for grazing, and the animals were forced to adapt nocturnal habits to avoid relentless hunting pressure (17 were killed in a single hunt by helicopter in 1968). By 1970 only around 100 remained, with perhaps a few other groups surviving in isolated areas. Since then the species and its main refuges have been given some protection, although enforcement is difficult. In 2016 the total population was estimated at 430. The Mindoro brown deer (Rusa marianna barandana) is confined to Mindoro, where it is seriously threatened by loss of habitat and hunting. The Mindoro warty pig (Sus oliveri) is nowadays confined to the mountains of central and north-western Mindoro, where it is heavily hunted and extremely rare. The Mindoro climbing rat (Anonymomys mindorensis) is known only from Ilong Peak in the Halcon Range of northcentral Mindoro, although it may be more widely distributed. Figure 5.8 Tamaraw (Bubalus mindorensis). (Credit: Unknown.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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The Mindoro shrew (Crocidura mindorus) is thought to be confined to the higher elevations of Mount Halcon in northcentral Mindoro. The Ilin bush-tailed cloud rat (Crateromys paulus) is known only from a single specimen said to have been collected on Ilin, a small island located just south of Mindoro, in 1953. The island’s forests have since been entirely destroyed. However, it seems possible that the species may still survive in southern Mindoro. The Mindoro stripe-faced fruit bat (Styloctenium mindorensis) is known only from a few localities in western Mindoro and from the islands of Ilin and Ambulong. The Mindoro wattle-winged fruit bat (Desmalopex microleucopterus) is known only from a few localities on Mindoro. The Mindoro hawk-owl (Ninox mindorensis) is confined to Mindoro, where it has been collected at all elevations. The Mindoro hornbill (Penelopides mindorensis) was historically widespread and abundant on Mindoro up until the 1970s, but has declined rapidly ever since due to habitat destruction and hunting. The Mindoro racquet-tail (Prioniturus mindorensis) is a type of parrot confined to Mindoro. The Mindoro imperial pigeon (Ducula mindorensis) is found patchily in the mountains of central and northwestern Mindoro. The Mindoro bleeding-heart (Gallicolumba platenae) is a type of ground dove confined to Mindoro, where it was reported to be widespread and common up to the midtwentieth century. Habitat destruction and hunting have seriously reduced its numbers in recent years. The black-hooded coucal (Centropus steerii) is a type of cuckoo that was historically widespread and common on Mindoro. Now confined to a few lowland forest patches, its total population is believed to be less than 400. The Mindoro black-bibbed cicadabird (Edolisoma mindanense elusum) is confined to Mindoro. The Mindoro golden-backed frog (Pulchrana mangyanum) is confined to Mindoro and Sibay. Schmacker’s shrub frog (Philautus schmackeri) is confined to high- and mid-elevation forests on Mindoro. The Mindoro barb (Barbodes hemictenus) is known only from the Sabaan, Mamboc, and Baco rivers and Lake Naujan.

Samar Samar is a large island located in the central Philippines. Ferron’s wolf snake (Lycodon ferroni) is known only from a single specimen collected on Samar. Marx’s blind snake (Ramphotyphlops marxi) is known only from north-western Samar. The Samar tree frog (Polypedates hecticus) is known only from a single locality on Samar.

Leyte Leyte is located just south of Samar in the central Philippines.

The Leyte goby (Sicyopus axilimentus) is confined to Lagu Lagu creek on Leyte.

Panay Panay is a triangular, mountainous island located in the central Philippines. The Panay busy-tailed cloud rat (Crateromys heaneyi) is confined to western Panay, where it has been collected from both lowland and montane localities. The Panay golden-crowned fruit bat (Acerodon jubatus lucifer) was an endemic subspecies now considered to be extinct, most like due to hunting. The Panay monitor (Varanus mabitang) is confined to remnant forests in the north-western and western mountains of Panay, where it is very rare and threatened by habitat destruction. Ernst Keller’s gecko (Gekko ernstkelleri) is known only from the north-western coastal area of Panay. Cornfield’s fringed gecko (Luperosaurus corfieldi) is known only from three specimens collected from far northwestern Panay. Sison’s skink (Parvoscincus sisoni) is known only from a few specimens collected in the central mountains of Panay. The Panay reed snake (Pseudorabdion talonuran) is known only from the western foothills of Mount Madja-as, within the island’s western coastal mountain range. The Panay stripe-lipped snake (Hologerrhum dermali) is known only from a few specimens collected in western Panay. Faust’s wolf snake (Lycodon fausti) is known only from a few specimens collected on Panay. Paeng’s wrinkled ground frog (Platymantis paengi) and the Panay forest wrinkled ground frog (P. panayensis) are both confined to Panay. The Gigantes Islands The Gigantes Islands are located north-east of Panay. The Gigantes narrow-disked gecko (Gekko gigante) is confined to Gigante Norte, Gigante Sur and a few nearby islets. It is threatened by mining activities. The Gigante wrinkled ground frog (Platymantis insulatus) is confined to the Gigantes Islands.

Palawan Palawan is a long, narrow island located in the south-western Philippines. Together with its numerous satellite islands it is notable for a flora and fauna that displays more affinities with certain islands in Indonesia, especially Borneo, than with the rest of the Philippines. Unfortunately, much has been lost or degraded. The Palawan bearded pig (Sus ahoenobarbus) occurs on Palawan and a number of satellite islands, where it appears to be fairly safe. The Philippine pangolin (Manis culionensis) is confined to Palawan, the Calamian Islands (Culion, Calauit, and a number of smaller islands) and possibly Balabac. It is seriously

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threatened by hunting both for its meat as well as use in ‘traditional medicine’. The Philippine porcupine (Hystrix pumila) is confined to Palawan and Busuanga. The Palawan montane squirrel (Sundasciurus rabori) is restricted to a few pockets of montane forest, where it is known from only a few specimens. The Palawan pencil-tailed tree mouse (Chiropodomys calamianensis) is confined to lowland forests on Palawan, the Calamian Islands, and Balabac. The Palawan fruit bat (Acerodon leucotis) is confined to Palawan, the Calamian Islands (Busuanga), and Balabac. The Palawan hornbill (Anthracoceros marchei) is confined to Palawan and satellite islands. The blue-headed racquet-tail (Prioniturus platenae) is a type of parrot confined to Palawan and satellite islands. The Palawan peacock-pheasant (Polyplectron napoleonis) is confined to Palawan. The Palawan flycatcher (Ficedula platenae) is confined to Palawan and satellite islands. The falcated wren-babbler (Ptilocichla falcata) is confined to lowland forest fragments on Palawan. The Palawan forest turtle (Siebenrockiella leytensis) was long known only from three specimens collected in 1921 and a fourth in 1988. Viable populations have since been discovered, but the species is seriously threatened by collection for food and the international pet trade. The Palawan fringed gecko (Luperosaurus palawanensis) is known only from a single specimen. Wright’s skink (Sphenomorphus wrighti) is known only from three specimens collected in the 1920s and 1940s from two localities in central Palawan. The Mount Victoria skink (Parvoscincus palawanensis) is known with certainty only from the original collection of three specimens in 1959 from Mount Victoria in central Palawan. The Palawan reed snake (Calamaria palavanensis) has not been reported since its initial collection in the 1960s. The white-striped flathead toad (Pelophryne albotaeniata) is known only from a few localities. The Palawan horned frog (Megophrys ligayae) is confined to Palawan and Balabac. The Philippine flat-headed toad (Barbourula busuangensis) is confined to Palawan, the Calamian Islands (Busuanga and Culion), and Balabac. The Palawan eastern frog (Alcalus mariae) is known only from Mount Balabag in southern Palawan. The Malatgan River caecilian (Ichthyophis weberi) is known only from a few localities in central Palawan. Lake Manguao Lake Manguao is located in the hills of northern Palawan. The Manguao barb (Barbodes manguaoensis) is confined to Lake Manguao. The Manguao sleeper goby (Bostrychus expatria) is confined to Lake Manguao.

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The Calamian Islands The Calamian Islands are located north of Palawan. They include four larger and numerous smaller islands and islets. The Calamian hog deer (Axis calamianensis) is today confined to three of the four larger islands in the Calamians (Busuanga, Calauit, and Culion), but is absent from Coron. It is reported to have occurred on at least nine other, smaller islands, but is now thought to survive only on Marily and Dimaquiat. Balabac Balabac lies to the south of Palawan, about 50 km from the northern tip of Borneo. The Balabac chevrotain (Tragulus nigricans) is a type of ‘mouse-deer’ originally found only on the islands of Balabac, Bugsuk, and Ramos, but has also been introduced to Apulit and Calauit. While the population appears stable, the species is everywhere subject to poaching for food.

Negros Negros is a relatively large, mountainous island located in the central Philippines. It too has been mostly deforested, particularly in the lowlands. The Negros shrew (Crocidura negrina) is known only from six specimens collected in southern Negros. The Negros fruit dove (Ptilinopus arcanus) is known only from a single specimen collected in 1953 on Mount Canlaon in north-central Negros. The Negros striped babbler (Zosterornis nigrorum) is confined to montane forest on Negros. The black slender skink (Lipinia rabori) is confined to southern Negros. The mountain burrowing snake (Pseudorabdion montanum) is known only from a few specimens. Taylor’s worm snake (Malayotyphlops canlaonensis) is known only from a single specimen collected on Mount Canlaon in 1917. All habitat at the type locality has since been completely deforested and the species is most likely extinct. The cave wrinked ground frog (Platymantis spelaeus) is confined to limestone karst areas and caves in southern Negros. Hazel’s wrinkled ground frog (P. hazelae) is confined to the mountains of Negros and possibly Masbate (although the latter population may represent a distinct form).

Ticao Ticao is a small island located in the central Philippines. The Ticao hornbill (Penelopides panini ticaensis) was historically endemic to Ticao, where it was abundant in the early twentieth century. There is little remaining habitat on the island, however, and the subspecies is almost certainly extinct.

Cebu Cebu is a long, narrow and mountainous island. Heavily populated, it has lost most of its original forests. The Cebu warty pig (Sus cebifrons cebifrons) was endemic to the island, where it is now believed to be extinct.

The Wallacean Region

The Cebu white-bellied woodpecker (Dryocopus javensis cebuensis) has not been seen in many years, and is now considered to be extinct. The Cebu burrowing skink (Brachymeles cebuensis) has only rarely been reported since its discovery in 1967. The Cebu blind snake (Malayotyphlops hypogius) is known only from a single specimen collected in the mid-twentieth century.

Bohol Bohol is located in the south-central Philippines. The Bohol tarsier (Carlito syrichta fraterculus) is a type of nocturnal primate confined to Bohol, where it is threatened by habitat destruction and collection for use as pets. Figure 5.9 Cebu four-coloured flowerpecker. (Credit: Joseph Smit.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The Cebu hawk-owl (Ninox rumseyi) was long known only from a single specimen collected in 1888, and not recorded again until 1998. Today, fewer than 200 pairs remain within a handful of forest fragments. The Cebu hanging parakeet (Loriculus philippensis chrysonotus) was long thought to be extinct and may indeed be so, although a report of the possible capture of a single specimen provides hope that it may still survive in a remote area of central Cebu. The Cebu brown dove (Phapitreron frontalis) was long thought extinct, but there have been a few reports of its continued survival in recent years. The Cebu amethyst-brown fruit dove (Phapitreron amethystinus frontalis) was most likely extinct by 1892. The Cebu four-coloured flowerpecker (Dicaeum quadricolor) is a type of passerine bird that was feared extinct in the early twentieth century, but rediscovered in very low numbers in 1992. The Cebu black shama (Kittacincla cebuensis) is a type of passerine bird that was long feared extinct. In recent years it has been found in a considerable number of localities, but remains highly threatened. The Cebu streak-breasted bulbul (Hypsipetes siquijorensis monticola) was long thought to be extinct but was rediscovered, in small numbers, in the late twentieth century. The Cebu bar-bellied cuckoo-shrike (Coracina striata cebuensis) was last seen in 1906, and was likely extinct by 1947. The Cebu blackish cicadabird (Coracina coerulescens altera) was last seen in 1906, and was likely extinct by 1947. The Cebu orange-bellied flowerpecker (Dicaeum trigonostigma pallidius) was last seen in 1906, and was likely extinct by 1947. The Cebu Everett’s white-eye (Zosterops everetti everetti) was last seen in 1906, and was likely extinct by 1947. The Cebu dark-throated oriole (Oriolus xanthonotus assimilis) was last seen in 1906, and was likely extinct by 1947.

Siquijor Siquijor is a hilly island located about 45 km north of Mindanao in the southern Philippines. The Siquijor hanging parakeet (Loriculus philippensis siquijorensis) is most likely extinct. The Siquijor streak-breasted bulbul (Hypsipetes siquijorensis siquijorensis) is confined to the island, where it is highly threatened.

Mindanao Located in the south-eastern Philippines, Mindanao is the second largest island in the archipelago. Most of the original lowland forest areas have long been cleared and only vestiges remain within the mountains. Two subspecies of Philippine brown deer (Rusa marianna), previously discussed, are endemic to Mindanao. The Mindanao uplands brown deer (R. m. nigella) is confined to a few isolated upland areas, while the Mindanao lowlands brown deer (R. m. nigricans) is found patchily at lower elevations. Both are threatened by habitat destruction and hunting. The Davao squirrel (Sundasciurus davensis) is known only from a single locality in eastern Mindanao. The Dinagat moonrat (Podogymnura aureospinula) is known from Dinagat and the nearby islands of Bucas Grande and Siargao, where it appears to be fairly common. The spiny long-footed rat (Tarsomys echinatus) is known only from the region of Mount Kitanglad and Mount Matutum. The greater lowland forest mouse (Apomys littoralis) is known only from the coastal plain of central Mindanao. The Mindanao golden-crowned fruit bat (Acerodon jubatus mindanensis) is found patchily on Mindanao, where it is threatened by bushmeat hunting. The Mindanao spotted imperial pigeon (Ducula carola mindanensis) is confined to Mindanao, where it is threatened by habitat destruction and hunting. The blue-capped kingfisher (Actenoides hombroni) was historically widespread on Mindanao but is now confined to a few scattered localities.

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The Mindanao jungle flycatcher (Rhinomyias goodfellowi) is known only from a few montane localities in central and southern Mindanao. The Mindanao miniature babbler (Micromacronus sordidus) is confined to a few montane localities. The Mindanao pygmy babbler (Dasycrotapha plateni) is a rare species found patchily throughout Mindanao. The Diwata skink (Sphenomorphus diwata) is known only from a few montane localities. The six-striped supple skink (Eutropis englei) is known only from south-central Mindanao. The Cotabato worm skink (Brachymeles pathfinderi) is known only from south-central Mindanao. The red-tailed emo skink (Emoia ruficauda) is known only from a few specimens collected on Mindanao. Taylor’s burrowing snake (Pseudorabdion taylori) is known only from a single specimen collected during the nineteenth century from an imprecise locality on Mindanao. The Tagibo wart frog (Limnonectes diuatus) is a semiaquatic species confined to northern Mindanao and Dinagat. The Zamboanga blood frog (Sanguirana everetti) is known from a few localities in western and southern Mindanao. The molted shrub frog (Philautus surrufus) is confined to a few scattered mountainous areas on Mindanao. The Cascade barb (Barbodes cataractae) is known only from the Cascade River and the brackish waters of Murcielagos Bay, where it is commercially important. The Tumba barb (B. tumba) is confined to a few small lakes and rivers. The Kitanglad Mountains The Kitanglad Mountains are located in north-central Mindanao (Bukidnon province). They include Mount Apo National Park, most of which has been deforested. The Kitanglad shrew mouse (Crunomys suncoides) is known only from a single specimen collected in 1993 from Mount Kitanglad. Mount Malindang Mount Malindang is an inactive volcano located on the Zamboanga Peninsula of western Mindanao (Misamis Occidental province). The greater Mindanao shrew (Crocidura grandis) was long known only from two specimens recorded from Mount Malindang in 1906. In recent years other specimens from the same locality, and which likely represent the species, have also been collected. Gary’s mountain stream snake (Opisthotropis alcalai) is known only from Mount Malindang. Dapitan Peak Dapitan Peak is located on the Zamboanga Peninsula of western Mindanao (Misamis Occidental province). The rusty skink (Lipinia zamboangensis) is known only from Dapitan Peak.

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Lake Lanao Lake Lanao is a large and ancient lake located in the mountains of central Mindanao (Lanao del Sur province). Overfishing and the introduction of common carp (Cyprinus carpio) and tank goby (Glossogobius giuris) have driven an endemic fish ‘species flock’ of the genus Barbodes within Lake Lanao and its associated waters to the point of extinction. Among those already believed to be lost are the Baoulan barb (B. baoulan), Clemens’ barb (B. clemensi), the katapa-tapa barb (B. flavifuscus), the Katolo barb (B. katolo), the kandar barb (B. lanaoensis), the manalak barb (B. manalak), the Agus Rapids barb (B. resimus), the bitungu barb (B. truncatulus), the pait barb (B. amarus), Herre’s barb (B. herrei), the disa barb (B. disa), the tras barb (B. tras), the rubberlip barb (B. pachycheilus), the prawn-eating barb (B. palaemophagus), and the palata barb (B. palata). The lindog barb (B. lindog) and the sirang barb (B. sirang) have not been recorded in many years and are possibly extinct. Lake Mainit Lake Mainit is located in north-eastern Mindanao (Surigao del Norte and Agusan del Norte provinces). At 223 m it is the deepest in the Philippines, and is also one of the largest. The Mainit priapum fish (Neostethus thessa) is confined to Lake Mainit, where it is threatened by pollution. The Dinagat Islands The Dinagat Islands are located off the northern tip of Mindanao. The Dinagat bushy-tailed cloud rat (Crateromys australis) is known only from a single specimen collected on Dinagat in the mid-1970s. Surveys since then have repeatedly failed to find others, and the species may be extinct. The Dinagat hair-tailed rat (Batomys russatus) is similarly known only from two individuals collected in 1975. Camiguin Camiguin is located off the northern coast of Mindanao in the Bohol Sea. It remains fairly well-forested, with an active stratovolcano on its south-western tip. The Camiguin forest rat (Bullimus gamay) is confined to Camiguin. The Camiguin forest mouse (Apomys camiguinensis) is known only from the central highlands of Camiguin. The Camiguin hawk-owl (Ninox leventisi) is confined to the island, where the remaining small population is threatened by habitat destruction. The Camiguin cross frog (Aphantophryne nana) is confined to Camiguin.

The Sulu Archipelago The Sulu Archipelago (Kapuluan ng Sulu in Filipino) is a chain of island groups located south-west of the main Philippine islands. Densely populated, most of the original forest has been cleared by logging.

The Wallacean Region

The Sulu hornbill (Anthracoceros montani) was historically found on three islands in the Sulu Archipelago (TawiTawi, Jolo and Sanga-Sanga), where it was described as common to abundant in the late nineteenth century. It has since disappeared from Jolo and Sanga-Sanga, and today survives only in mountain forest on Tawi-Tawi where it is threatened by loss of habitat. The total population is believed to be less than 50. The Sulu pygmy woodpecker (Picoides ramsayi) was historically widespread and abundant in the Sulu Archipelago. Already rare by the early 1970s, it is now confined to TawiTawi and fringing coralline islands. The limbless burrowing skink (Brachymeles vermis) is confined to Jolo, Tawi-Tawi and satellite islands. The Sulu leaf-litter skink (Tytthoscincus biparietalis) is confined to Basilan, Jolo, Tawi-Tawi and a few small satellite islands. The Sulu kukri snake (Oligodon meyerinkii) was historically found on Jolo, Tawi-Tawi and satellite islands. It was last recorded on Tawi-Tawi in the 1990s. The Sulu reed snake (Calamaria joloensis) is known only from a single specimen collected from an indistinct locality within the Sulu Archipelago (possibly Jolo). The Sulu blind snake (Ramphotyphlops suluensis) is known from Sibutu and Bubuan. The Papahag golden-backed frog (Hylarana melanomenta) is confined to Tawi-Tawi, Sibutu and a few satellite islands. Basilan Basilan is the largest and northernmost of the Sulu Islands. It comprises an extinct volcanic cone rising from a submarine ridge, surrounded by numerous satellite islands. Bartlett’s bleeding-heart (Gallicolumba crinigera bartletti) is a type of ground dove that became extinct in the wild during the mid-1990s, although a captive population exists in American zoos. The Basilan wart frog (Limnonectes micrixalus) is now confined to Basilan, although it may have formerly occurred on Mindanao as well. Jolo Jolo is an extinct volcanic cone rising from a submarine ridge, surrounded by numerous satellite islands. The Jolo fringed gecko (Luperosaurus joloensis) is known only from a small number of specimens collected on Jolo. A record from Mindanao appears to be erroneous. Tawi-Tawi The southernmost island in the Sulu Archipelago, Tawi-Tawi is covered by limestone and surrounded by numerous satellite islands. Virtually all of its original forests were destroyed by the end of the twentieth century. The Tawi-Tawi forest rat (Rattus tawitawiensis) is known only from three specimens collected on Tawi-Tawi in 1971.

The Tawi-Tawi hawk-owl (Ninox reyi) is confined to TawiTawi and satellite islands. The blue-winged racquet-tail (Prioniturus verticalis) is a type of parrot confined to Tawi-Tawi and satellite islands. The Tawi-Tawi brown dove (Phapitreron cinereiceps) is confined to Tawi-Tawi and adjacent Sanga-Sanga. The Tawi-Tawi bleeding-heart (Gallicolumba menagei) is a type of ground dove long known only from two specimens collected on Tawi-Tawi in 1891. Reported to be abundant prior to the 1970s, it declined dramatically thereafter and was last reported in 1995.

The Sangir Islands The Sangir Islands or Sangihe Islands (Kepulauan Sangir in Indonesian) are located north-east of Sulawesi. The elegant sunbird (Aethopyga duyvenbodei) is now confined to Great Sangir, although it is known to have formerly occurred on Siau as well.

Great Sangir Island The largest and northernmost island, Great Sangir (Sangir Besar in Indonesian, and sometimes called Sangihe or Sangir) is dominated by Mount Awu, an active volcano. Most of its original forest cover has been converted to agriculture. The Sangir tarsier (Tarsius sangirensis) is a type of nocturnal primate confined to Great Sangir. The red and blue lory (Eos histrio) was historically found in the Talaud and Sangir islands. The Sangir red and blue lory (E. h. histrio) was driven into extinction prior to the 1990s by hybridization with escaped trade individuals of the Talaud red and blue lory (E. h. talautensis). The Sangir shrike-thrush (Coracornis sanghirensis) was long known only from a single specimen collected in the late nineteenth century. It was rediscovered in the mountains of Great Sangir in 1995, where the total population is estimated at less than 100. The Sangir dwarf kingfisher (Ceyx sangirensis) is confined to Great Sangir, where it has not been recorded since 1997 and may indeed be extinct. The Sangir white-eye (Zosterops nehrkorni) is an extremely rare species long known only from a single specimen collected in the late nineteenth century. Rediscovered in 1996 and recorded again in 1999, after which it was not seen again until 2009. The total population is believed to number fewer than 50. The Cerulean paradise-flycatcher (Eutrichomyias rowleyi) is known only from a single specimen collected during the nineteenth century, and long presumed extinct. It was rediscovered in very small numbers in 1998. The Sangir pitta (Erythropitta caeruleitorques) is a type of passerine bird confined to Great Sangir. The Sangir golden bulbul (Thapsinillas platenae) is known only from a single locality on Great Sangir, where the total population is between 50 and 230.

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The Sangir gliding tree frog (Rhacophorus rhyssocephalus) is confined to Great Sangir.

Reports of specimens from Great Sangir Island probably refer to an as-yet undescribed form.

Siau

Miangas

Siau (Pulau Siau in Indonesian) is a tiny island with an active volcano on one end and a large human population on the other. Virtually all of its original forest has been destroyed and the remaining wildlife is heavily hunted. The Siau tarsier (Tarsius tumpara) is one of the world’s most endangered primates. It is confined to the few remaining forest fragments on Siau and, conceivably, on a few very small satellite islands. The Siau scops owl (Otus siaoensis) is known only from a single specimen collected in 1866. The Siau pitta (Erythropitta palliceps) is a type of passerine bird confined to Siau and the satellite islands of Tagulandang and Ruang.

Miangas is a small, low-lying island covered in coconut palms. The Miangas red and blue lory (Eos histrio challengeri) is an extinct form likely (but not positively) known from the island.

The Talaud Islands The Talaud Islands (Kepulauan Talaud in Indonesian) are a small group located north-east of the Sangir Islands. They are frequently beset by earthquakes and volcanic eruptions. Two small species of rodent of the genus Melomys, perhaps sharing a common ancestor, are endemic to Karakelang and Salibabu. The terrestrial Talaud mosaic-tailed rat (M. caurinus) is known only from four specimens, while the arboreal Talaud mosaic-tailed rat (M. talaudium) is similarly littleknown. The Talaud fruit bat (Acerodon humilis) is confined to Karakelang and Salibabu, where it was long known only from three specimens collected in 1909. It was rediscovered in 2002. The Talaud bush-hen (Amaurornis magnirostris) is known for certain only from Karakelang, although a small number may also survive on Salibabu. The Talaud red and blue lory (Eos histrio talautensis) was formerly widespread in the Talaud Islands, but is now largely confined to Karakelang. It is seriously threatened by illegal collection for the pet trade.

Karakelang Karakelang is the largest of the Talaud Islands. It has a notably steep coast except for the southern shore, which is fringed by a wide coral reef. Remaining forests are somewhat protected. The Karakelang rail (Gymnocrex talaudensis) is confined to Karakelang, where it has only been observed once since its discovery in the 1990s. The Karakelang pitta (Erythropitta inspeculata) is a type of passerine bird confined to Karakelang.

Salibabu Salibabu has been largely converted for agricultural purposes, although some remnant forest remains in its lower hills. The Salibabu bear cuscus (Ailurops melanotis) is a rare species of arboreal marsupial likely endemic to the island.

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Sulawesi Sulawesi is a large island located between Borneo and the Moluccas and north of Java. It consists of four long peninsulas joined to a mountainous central core. The volcanic and tectonic activities that created the island left in its wake a network of streams and ravines, along with the massive rifts and craters that later became rivers, lakes, and upland plains. Many species on Sulawesi can trace their origins to the continents of Asia and Australia, although the island’s geographic history and relative isolation have given it its own distinctive fauna. While most of Indonesia’s larger islands suffer from extreme deforestation, Sulawesi still supports relatively extensive tracts of both montane and lowland moist forests. Sulawesi’s most famous mammals are two species of anoa (Bubalus), dwarf buffalo that in the past were collectively distributed over the whole of the island. Quarles’ anoa (B. quarlesi) occurs patchily in both highland and lowland areas throughout Sulawesi as well as on the island of Buton, off the south-eastern coast. Uncontrolled hunting (mainly for food) and habitat destruction have reduced it to under 2500 individuals. Smith’s anoa (B. depressicornis) is also found throughout Sulawesi and the island of Buton, where it favours open, coastal areas. It too has been wantonly hunted, reportedly even by military units with machine guns. Several species of macaque (Macaca) are endemic to Sulawesi, where they are threatened by habitat destruction, hunting, trapping, poisoning, human persecution and capture for use as pets. The black crested macaque (M. nigra) is naturally confined to the Tangkoko Reserve at the tip of Sulawesi’s northern peninsula (Sulawesi Utara province), as well as to the small neighbouring islands of Manado Tua, Talise and (formerly) Lembeh. There is a fairly large introduced population on the island of Bacan in the Moluccas, where the species has become a crop pest. In Sulawesi itself the species is heavily hunted for its meat and has been extirpated from many areas. The Gorontalo macaque (M. nigrescens) is confined to the central section of Sulawesi’s northern peninsula (Gorontalo and Sulawesi Utara province), where it too is highly threatened by bushmeat hunting. It is protected within Bogani Nani Wartabone National Park. Heck’s macaque (M. hecki) is confined to the southern and western sections of the northern peninsula of Sulawesi (Gorontalo and Sulawesi Tengah provinces). The Tonkean macaque (M. tonkeana) is relatively widespread in central Sulawesi. A small population on Malenge in the Togean Islands is thought to have been introduced. The Moor macaque (M. maura) is

The Wallacean Region

largely confined to the highland and karst forests of the southwestern peninsula of Sulawesi (Sulawesi Selatan province), although a few populations remain in lowland areas as well. The booted macaque (M. ochreata) is divided into two subspecies. The Sulawesi booted macaque (M. o. ochreata) is found throughout the south-eastern peninsula of Sulawesi, extending to the north of the lakes region (Sulawesi Tengah, Sulawesi Selatan and Sulawesi Tenggara provinces). The Muna-Butung booted macaque (M. o. brunnescens) was historically found on Muna and Butung, although it may have been exterminated from the former as it is now almost completely deforested. It may also occur on the small island of Labuan Blanda off Sulawesi’s eastern coast. Several species of tarsier (Tarsius) are confined to small areas of Sulawesi and its satellite islands, where they are threatened by loss of habitat, agricultural pesticides, predation by domestic dogs and cats, hunting for use as food, and occasional collection for the pet trade. Dian’s tarsier (T. dentatus) occurs in the eastern portion of the central core of Sulawesi to the tip of the eastern peninsula (Sulawesi Tengah province). The Lariang tarsier (T. lariang) is confined to central-western Sulawesi (Sulawesi Barat province). The Makassar tarsier (T. fuscus) is found on the south-western peninsula of Sulawesi south of the Lake Tempe Depression (Sulawesi Selatan province). Wallace’s tarsier (T. wallacei) has a small, discontinuous range in north-western Sulawesi (Sulawesi Tengah province). The Manado tarsier (T. spectrumgurskyae) is found on the northern peninsula of Sulawesi from the Isthmus of Gorontalo to the tip (Sulawesi Utara province). The Bumbulan tarsier (T. supriatna) is found on the northern peninsula of Sulawesi from the Isthmus of Gorontalo westward at least as far as Sejoli and possibly Ogatemuku (Gorontalo province). The Sulawesi babirusa (Babyrousa celebensis) is a pig-like animal still found throughout Sulawesi as well as on the offshore islands of Muna, Buton and Lembeh. It is threatened by hunting and loss of its rainforest habitat. The Sulawesi palm civet (Macrogalidia musschenbroekii) is a rare species found patchily throughout Sulawesi and a few offshore islands, where it is threatened by habitat destruction and predator control efforts. The Sulawesi bear cuscus (Ailurops ursinus) is widespread in the forests of Sulawesi, Peleng in the Banggai Islands, the Togean Islands, Buton, and possibly Muna. It is everywhere threatened, however, by loss of habitat, hunting and collection for the pet trade. The Minahasa long-nosed squirrel (Hyosciurus ileile) is known only from a few localities within the northern peninsula of Sulawesi (Gorontalo and Sulawesi Tengah provinces). The Minahasa hill rat (Bunomys fratrorum) is confined to the north-eastern part of the northern peninsula of Sulawesi (Sulawesi Utara province). The south-eastern xanthurus rat (Rattus salocco) is known only from three specimens collected from the south-eastern peninsula of Sulawesi (Sulawesi Tenggara province), the last in

1932. The south-western xanthurus rat (R. bontanus) is known only from a few localities on the south-western peninsula of Sulawesi (Sulawesi Selatan province). The Minahasa taeromys rat (Taeromys taerae) is known only from a small area of the northern peninsula of Sulawesi (Sulawesi Utara and Gorontalo provinces), where it was last collected in 1932. The greater taeromys rat (T. callitrichus) is known only from a few localities in northern and west-central Sulawesi, where it is threatened by loss of habitat and hunting. The Sulawesi Utara spiny rat (Echiothrix leucura) is confined to the north-eastern tip of Sulawesi’s northern peninsula (Sulawesi Utara province). The Sulawesi harpy fruit bat (Harpyionycteris celebensis) is found throughout Sulawesi and on the island of Buton, but is everywhere rare due to overhunting for food. The small-toothed fruit bat (Neopteryx frosti) is known only from two disjunct localities in central and northern Sulawesi. It was last recorded in 1991. The Sulawesi rousette fruit bat (Boneia bidens) is widespread across Sulawesi except for the south-west, as well as the islands of Lembeh and Buton, but is everywhere threatened by hunting and habitat destruction. The crested leaf-nosed bat (Hipposideros inexpectatus) is known only from three historical collections in central and northern Sulawesi, the first two in the 1870s and the third at some time prior to 1925. Sclegel’s masked owl (Tyto inexspectata) is a rare species known only from a handful of historical specimens and a few more recent sightings originating from northern and northcentral Sulawesi. The maleo (Macrocephalon maleo) is a large gallinaceous bird from the rainforests of Sulawesi and Buton, where it has been much reduced by habitat destruction, hunting, and the unsustainable harvesting of its eggs. The knobbed hornbill (Rhyticeros cassidix) is found throughout Sulawesi as well as on Lembeh, Muna, Buton and the Togean Islands. It has undergone significant declines due to hunting and loss of habitat. Temminck’s hornbill (Rhabdotorrhinus exarhatus) is divided into two subspecies. The northern Temminck’s hornbill (R. e. exarhatus) is confined to northern Sulawesi and the island of Lembeh. The southern Temminck’s hornbill (R. e. sanfordi) is found in central, eastern and southern Sulawesi and on the islands of Muna and Buton. Both are threatened by habitat destruction, hunting for food and collection for use as pets. The Sulawesi yellow-crested cockatoo (Cacatua sulphurea sulphurea) was historically found throughout Sulawesi and a few satellite islands, but has undergone a massive decline due to overcollection for the pet trade. The blue-faced rail (Gymnocrex rosenbergii) is known from a few localities on the northern peninsula of Sulawesi and the islands of Peleng and Buton. The snoring rail (Aramidopsis plateni) is a secretive species known only from a handful of records, very few of them

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recent, originating throughout Sulawesi and the island of Buton. It is thought to be genuinely rare and threatened by loss of habitat. Heinrich’s nightjar (Eurostopodus diabolicus) was long known only from a single specimen collected from Mount Klabat on the northern peninsula of Sulawesi. Apart from a few unconfirmed reports in the following decades it was officially discovered again until 1996, and since then it has been observed in a few localities in northern and westcentral Sulawesi. Forsten’s tortoise (Indotestudo forstenii) is endemic to northern Sulawesi, where it is threatened by overcollection for food and the international pet trade. Conrad’s worm snake (Ramphotyphlops conradi) is known only from a single specimen collected during the late nineteenth century from an undefined locality at the northern tip of Sulawesi’s northern peninsula (Sulawesi Utara province).

Mountains The central part of Sulawesi is ruggedly mountainous, such that the island’s peninsulas have historically been remote from each other, with better connections by sea than by road. The Sulawesi mountain tarsier (Tarsius pumilus) is an enigmatic species of nocturnal primate from the mountains of central and southern Sulawesi. It was known only from three specimens until fairly recently. The first was collected in 1916 from Rano Rano, the second in 1930 from Mount Rantemario, and a third was found dead in a rat trap on Mount Rorekatimbu in 2000. It had never been seen alive by scientists until 2008, when a few individuals were captured in Lore Lindu National Park. The central mountains taeromys rat (Taeromys hamatus) is known only from Mount Lehio, Mount Kanino, and Mount Nokilalaki in west-central Sulawesi (Sulawesi Tengah province). The reddish-furred taeromys rat (T. punicans) is known only from its original collection in 1918 from westcentral Sulawesi (Sulawesi Tengah province). Sommer’s Sulawesi rat (Sommeromys macrorhinos) is known only from a few localities in west-central Sulawesi (Sulawesi Barat and Sulawesi Selatan provinces). The Sulawesi soft-furred rat (Eropeplus canus) is known only from a few localities in west-central Sulawesi. Dollman’s spiny rat (Maxomys dollmani) is known only from two disjunct localities in west-central and south-eastern Sulawesi. The Sulawesi mountain shrew-rat (Melasmothrix naso) is so far known only from Mount Rano Rano, Mount Nokilalaki, and Mount Gandangdewata, but likely also occurs in other mountains in west-central Sulawesi. The worm-eating shrew-rat (Paucidentomys vermidax) is known only from two specimens collected from Mount Gandangdewata and Mount Latimojong in westcentral Sulawesi. The long-tailed Sulawesi shrew-rat (Tateomys macrocercus) and Tate’s Sulawesi shrew-rat (T. rhinogradoides) are

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each known only from a few disjunct localities in westcentral Sulawesi. The Sulawesi shrew-mouse (Crunomys celebensis) is known only from three localities in west-central Sulawesi. The Linduan rousette fruit bat (Rousettus linduensis) is known only from four specimens collected in Lore Lindu National Park, west-central Sulawesi (Sulawesi Tengah province). The Minahasa pipistrelle bat (Pipistrellus minahassae) is known with certainty only from a single specimen collected from the Minahassa Peninsula (Sulawesi Utara province). The Cinnabar hawk-owl (Ninox ios) is known only from four records originating within the highlands of northern and west-central Sulawesi. The Matinan flycatcher (Cyornis sanfordi) is known only from four localities within the mountains of the northern peninsula of Sulawesi (Sulawesi Utara, Gorontalo and Sulawesi Tengah provinces). The southern hylocitrea (Hylocitrea bonthaina) is a type of passerine bird known only from the mountains of the southwestern peninsula of Sulawesi (Sulawesi Selatan province), where it has rarely been recorded in recent decades. The Minahasa cross frog (Oreophryne celebensis) is confined to a small area of the northern peninsula of Sulawesi (Sulawesi Utara and Gorontalo provinces). The variable cross frog (O. variabilis) is known only from Mount Lompobatang and Mount Latimojong in south-western Sulawesi, and from Mount Karua in the central-west. Both species are threatened by loss of habitat. The Djikoro wart frog (Limnonectes arathooni) is a rare species known only from the montane forests of Mount Lompobatang and Mount Latimojong in south-western Sulawesi (Sulawesi Selatan province). The Lompobatang wart frog (L. microtympanum) is confined to Mount Lompobatang and Bantimurung National Park in south-western Sulawesi (Sulawesi Selatan province). Mount Nokilalaki Mount Nokilalaki (Gunung Nokilalaki in Indonesian) is located in west-central Sulawesi (Sulawesi Tengah province). The lesser Margareta rat (Margaretamys parvus) is confined to montane forest on Mount Nokilalaki. Mount Rorekatimbo Mount Rorekatimbo (Gunung Rorekatimbo in Indonesian) is located in west-central Sulawesi (Sulawesi Tengah province). The mossy forest shrew (Crocidura musseri) is known only from montane forest on Mount Rorekatimbo, but may occur more widely. Mount Tambusisi Mount Tambusisi (Gunung Tambusisi in Indonesian) is located in east-central Sulawesi (Sulawesi Tengah province). It lies within the Morowali Nature Reserve. Watts’ spiny rat (Maxomys wattsi) is known only from a group of specimens collected on the forested slopes of Mount Tambusisi in 1980. It may occur more widely.

The Wallacean Region

The long-headed hill rat (Bunomys prolatus) is confined to Mount Tambusisi. The Mekongga Mountains The Mekongga Mountains (Pegunungan Mekongga in Indonesian) are located on the western side of the southeastern peninsula of Sulawesi (Sulawesi Tenggara province). The secretive dwarf squirrel (Prosciurillus abstrusus) is known only from a single collection in the 1950s from the Mekongga Mountains. The small-eared taeromys rat (Taeromys microbullatus) is known only from three specimens collected in 1932 from Mount Tanke Salokko. The plain-backed kingfisher (Actenoides regalis) is known only from two specimens collected from Mount Tanke Salokko. Zimmer’s cross frog (Oreophryne zimmeri) is known only from a single specimen collected in the 1930s from the Menkongga Mountains. Mount Moncong Lompobatang Mount Moncong Lompobatang (Gunung Moncong Lompobatang in Indonesian) is a prominent peak located on the tip of the south-western peninsula of Sulawesi (Sulawesi Selatan province). While protected within Gunung Lompobatang National Park, its forests are becoming highly fragmented due to selective logging and the creation of plantations. The Lompobatang soft-furred rat (Rattus mollicomulus) is confined to the upper slopes of Mount Moncong Lompobatang. The Lompobatang hill rat (Bunomys coelestis) is confined to Mount Moncong Lompobatang. The Lompobattang fruit dove (Ramphiculus meridionalis) is confined to Mount Moncong Lompobatang. The Lompobattang flycatcher (Ficedula bonthaina) is known only from two historical collections from Mount Moncong Lompobatang. Apparently common up until the 1930s, it has rarely been recorded since.

Lowland Rainforests While still relatively extensive compared to those of other islands in the region, Sulawesi’s lowland rainforests have suffered significant deforestation in recent decades. Weber’s dwarf squirrel (Prosciurillus weberi) is known only from a few specimens collected from a small area of south-central Sulawesi (Sulawesi Selatan province). The central Sulawesi spiny rat (Echiothrix centrosa) is known from a few scattered localities in northern and northcentral Sulawesi. The spiny lowland Margareta rat (Margaretamys beccarii) is known only from specimens collected in 1880 and 1976 from central and northern Sulawesi. The Minahassa ranee mouse (Haeromys minahassae) is known only from three disjunct localities in northern and central Sulawesi.

Gaskell’s false serotine bat (Hesperoptenus gaskelli) is known only from a single specimen collected in east-central coastal Sulawesi (Sulawesi Tengah province). Boeadi’s roundleaf bat (Hipposideros boeadii) is known only from a few specimens collected from Rawa Aopa Watumohai National Park, on the south-eastern peninsula of Sulawesi (Sulawesi Tenggara province). The Sulawesi forest turtle (Leucocephalon yuwonoi) is a little-known and highly threatened species from northern Sulawesi. Pak Djoko’s fringed gecko (Luperosaurus iskandari) is known only from the base of Mount Tompotika, in eastern Sulawsi (Sulawesi Tengah province). Boeseman’s reed snake (Calamaria boesemani) is known only from a single specimen collected in 1919 from the eastern peninsula (Sulawesi Tengah province). Heinrich’s wart frog (Limnonectes heinrichi) is confined to a small area of north-eastern Sulawesi (Sulawesi Utara and Gorontalo provinces).

Lakes, Rivers, and Marshes The old, isolated lakes of central Sulawesi host a very distinctive and highly endemic fauna and flora, in particular a large array of fishes, along with many rare aquatic plants such as floating ferns. Two species of large, semi-aquatic lizard, Peters’ sailfin lizard (Hydrosaurus celebensis) and the Makassar sailfin lizard (H. microlophus), are confined to wetland areas of Sulawesi, where they are threatened by habitat destruction and overcollection for the international pet trade. Boulenger’s water snake (Enhydris matannensis) is known only from Lake Matano and Muna Island. Ladiges’ rainbowfish (Marosatherina ladigesi) is known only from a few rivers on the south-western peninsula of Sulawesi (Sulawesi Selatan province), where it is threatened by pollution and overcollection for the international aquarium trade. Ladiges’ halfbeak (Nomorhamphus towoetii) is confined to Lake Towuti and Lake Poso. Lake Tondano Lake Tondano (Danau Tondano in Indonesian) is located in far northern Sulawesi (Sulawesi Utara province). The Tondano halfbeak (Tondanichthys kottelati) is confined to Lake Tondano. Lake Lindu Lake Lindu (Danau Lindu in Indonesian) is located in central Sulawesi (Sulawesi Tengah province), within Lore Lindu National Park. Sarasins’ buntingi (Xenopoecilus sarasinorum) is confined to Lake Lindu. Lake Poso Lake Poso (Danau Poso in Indonesian) is located in eastcentral Sulawesi (Sulawesi Tengah province). The third deepest

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lake in Indonesia, it has evolved a unique array of endemic fishes that have unfortunately been decimated by the introduction of exotic species such as Mozambique tilapia (Oreochromis mossambicus) and common carp (Cyprinus carpio), as well as by other factors. The duck-billed buntingi (Adrianichthys kruyti) is a type of ricefish that has not been recorded for many years, and is most likely extinct. Popta’s buntingi (Xenopoecilus poptae) and the egg-carrying buntingi (X. oophorus) are both confined to Lake Poso, where they are seriously threatened by overfishing and agrochemical pollution. The black ricefish (Oryzias nigrimas) and the sharp-jawed ricefish (O. orthognathus) are both confined to Lake Poso, where they are threatened by introduced species, pollution, and overfishing. The Poso halfbeak (Nomorhamphus celebensis) is confined to Lake Poso and tributary streams. The Poso mangrove goby (Mugilogobius amadi) was historically endemic to Lake Poso, where it was last recorded in 1978. It is almost certainly extinct, possibly due to volcanic and tectonic events that occurred in 1983 in addition to other factors. Sarasins’ goby (Tamanka sarasinorum) is confined to Lake Poso. The Malili Lakes System The Malili Lakes system is located in west-central Sulawesi (Sulawesi Selatan and Sulawesi Tenggara provinces). It consists of two large lakes (Matano and Towuti) and three smaller ones (Mahalona, Masapi, and Lontoa). Boulenger’s rainbowfish (Telmatherina celebensis) is known from Lake Towuti, the smaller lakes Mahalona and Lontoa, and the Tominanga River, where it is threatened by introduced predatory fish and pollution. Bont’s rainbowfish (T. bonti) is confined to the Malili Lakes system. Woltereck’s sailfin silverside (Paratherina wolterecki) is confined to Lake Towuti and the smaller Lake Mahalona. The marbled ricefish (Oryzias marmoratus) is confined to Lake Towuti and the smaller lakes Mahalona and Lontoa, as well as in a few tributary streams and creeks. Weber’s halfbeak (Nomorhamphus weberi) is known from Lake Matano and the smaller Lake Mahalona. Aurich’s goby (Glossogobius intermedius) is confined to Lake Towuti and Lake Mahalona. Weber’s goby (G. matanensis) is known from Lake Matano, Lake Towuti and Lake Mahalona. Both are threatened by introduced predatory fish, pollution, and sedimentation. The Malili Lakes mangrove goby (Mugilogobius latifrons) is known from Lake Matano, Lake Towuti, Lake Mahalona, and their tributaries. It is threatened by introduced predatory fish and sedimentation. Lake Matano (Danau Matano in Indonesian) is located in Sulawesi Selatan province. It is the deepest lake in South East Asia and the eighth deepest in the world.

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The Mengonuwai rainbowfish (Telmatherina obscura), Antonia Kottelat’s rainbowfish (T. antoniae), the opudi rainbowfish (T. opudi), Wahju’s rainbowfish (T. wahjui), Sarasins’ rainbowfish (T. sarasinorum), the prognathous rainbowfish (T. prognatha), Abendanon’s rainbowfish (T. abendanoni), and the white-lipped rainbowfish (T. albolabiosus) are all confined to Lake Matano, where they are threatened by introduced predatory fish and sedimentation. The Matano ricefish (Oryzias matanensis) is confined to Lake Matano. The Matano mangrove goby (Mugilogobius adeia) is confined to Lake Matano. Lake Towuti (Danau Towuti in Indonesian) is located in Sulawesi Selatan province. It is the largest lake in Sulawesi. The Towuti rainbowfish (Tominanga sanguicauda) is confined to Lake Towuti and its tributaries. The yellow-finned ricefish (Oryzias profundicola) is confined to Lake Towuti. The Towuti yellowfin goby (Glossogobius flavipinnis) is confined to shallow nearshore areas of Lake Towuti. Lake Mahalona (Danau Mahalona in Indonesian) is a small lake located between Lake Towuti and Lake Matano in Sulawesi Selatan province. The Mahalona rainbowfish (Tominanga aurea) is confined to Lake Mahalona. Lake Lontoa (Danau Lontoa in Indonesian, and also known as Lake Wawantoa) is a relatively small lake located to the north-east of Lake Towuti in Sulawesi Selatan province. The Lontoa sailfin silverside (Paratherina labiosa) is confined to Lake Lontoa, where it is threatened by increasing sedimentation due to deforestation.

The Togean Islands The Togean Islands (Kepulauan Togean in Indonesian) are an archipelago of 56 islands and islets located in the Gulf of Tomini, between the northern and eastern Sulawesi peninsulas. The Togean babirusa (Babyrousa togeanensis) is a type of wild swine confined to Batudaka, Togean, Talatakoh, and Malenge islands. However, the animals are excellent swimmers and are occasionally to be seen on offshore islets such as Pangempan. The Togean tarsier (Tarsius niemitzi) is a type of nocturnal primate confined to the Togean Islands.

The Banggai Archipelago The Banggai Archipelago (Kepulauan Banggai in Indonesian) is located off the eastern end of the central peninsula of Sulawesi. The Banggai fruit dove (Ramphiculus subgularis) is endemic to the Banggai Archipelago, where it is fairly common. Peleng Peleng (Pulau Peleng in Indonesian) is one of the principal islands of the Banggai Archipelago.

The Wallacean Region

The Peleng tarsier (Tarsius pelengensis) is currently known only from forest fragments on Peleng, but may be present on other islands in the Banggai Archipelago. Two species of rat (Rattus) are endemic to Peleng. Koopman’s rat (R. koopmani) is known only from a single specimen collected in 1938. The Peleng rat (R. pelurus) is known only from a few specimens collected in 1938. The Peleng scops owl (Otus mendeni) is confined to Peleng. The Banggai crow (Corvus unicolor) was long known only from two specimens collected from an unspecified island in the Banggai Archipelago. A sighting on the western slopes of Peleng in 1991 led to eventual surveys confirming that the species still survives there in small numbers. The Peleng jungle flycatcher (Cyornis pelingensis) is confined to Peleng.

Buton Buton (Pulau Buton in Indonesian) is a large island located off the south-eastern peninsula of Sulawesi. The Tuwa gliding tree frog (Rhacophorus georgii) appears to be confined to lowland rainforest on Buton, where it is rare.

The Tukangbesi Islands The Tukangbesi Islands are located immediately east of Buton in the Banda Sea. The Tukangbesi yellow-crested cockatoo (Cacatua sulphurea paulandrewi) is confined to the Tukangbesi Islands, where only a few individuals were reported to survive in 2009.

The Selayar Islands The Selayar Islands (Kepulauan Selayar in Indonesian) are located just south of the south-western peninsula of Sulawesi. Selayar Selayar is the largest of the Selayar Islands. The Selayar tarsier (Tarsius tarsier) is a type of nocturnal primate that was long thought to be widespread throughout Sulawesi and its satellite islands, but owing to considerable taxonomic splitting in recent years it now appears to be wholly or entirely confined to Selayar Island. Tanah Jampea Tanah Jampea (Pulau Tanah Jampea in Indonesian) is the second largest of the Selayar Islands. It has been designated an Important Bird Area by BirdLife International. The Tanah Jampea yellow-crested cockatoo (Cacatua sulphurea djampeana) is confined to Tanah Jampea, where in 2016 the total population was thought to be less than 100. The Tanah Jampea sunset lorikeet (Trichoglossus forsteni djampeanus) is (or was) confined to the island, where it is now most likely extinct. The white-tipped monarch (Symposiachrus everetti) is a type of passerine bird confined to Tanah Jampea.

Kalaotoa Kalaotoa (Pulau Kalaotoa in Indonesian) lies to the east of the Selayar Islands. The Kalaotoa sunset lorikeet (Trichoglossus forsteni stresemanni) is (or was) confined to the island, where it is now most likely extinct.

The Moluccas Archipelago The Moluccas (also known as the Maluku Islands), which lie between Sulawesi and New Guinea, include over 1000 islands ranging in size from fairly large to small and uninhabited islets of only a few hectares. Most are mountainous, some with active volcanoes. The largest two islands, Halmahera and Seram, are sparsely populated, while the most developed, Ambon and Ternate, are small. Rainforest covers most of the northern and central Moluccas, which, on the smaller islands, have been largely replaced by spice plantations. Four species of flying fox (Pteropus) are endemic to the Moluccas, where they are threatened by habitat destruction and hunting. The Moluccan flying fox (P. chrysoproctus) is fairly widespread in the islands and probably still not uncommon. Nevertheless, it has rarely been collected and is believed to be declining. The black-bearded flying fox (P. melanopogon) is confined to Seram, Buru and adjacent smaller islands in the central Moluccas, as well as in the Tanimbar Islands. Temminck’s flying fox (P. temminckii) is confined to Buru, Seram and Ambon in the central Moluccas. The spectacled flying fox (P. ocularis) is confined to Buru, Seram and (at least historically) Ambon. The lesser tube-nosed fruit bat (Nyctimene minutus) is confined to Buru and Seram in the central Moluccas. Rozendaal’s blossom fruit bat (Syconycteris carolinae) is confined to Halmahera and Bacan in the northern Moluccas. Stalker’s mouse-eared bat (Myotis stalkeri) is known only from a few specimens collected disjunctly from Gebe and the Kai Islands in the Moluccas, and from Waigeo and Batanta in the Raja Ampat Islands off the western coast of New Guinea. It is likely threatened by disturbance of its roosting sites. The Moluccan scrubfowl (Eulipoa wallacei) is still fairly widespread in the Moluccas, but has declined dramatically everywhere owing to the overcollection of its eggs. The Moluccan woodcock (Scolopax rochussenii) was long known only from eight specimens collected on Obi and Bacan in the northern Moluccas, the most recent in 1980. In 2010 and again in 2012 it was recorded from a few localities on Obi. Two species of cockatoo (Cacatua) endemic to the Moluccas are threatened by overcollection for the cage-bird trade. The white cockatoo (C. alba) is found on the islands of Halmahera, Bacan, Ternate, Tidore, Kasiruta, and Mandioli in the northern Moluccas, with an additional, introduced population on Taiwan. The salmon-crested cockatoo (C. moluccensis) appears to be nowadays confined to Seram and perhaps a single locality on Ambon. It formerly occurred on the smaller islands of Saparua and Haruku as well, but has been extirpated from there.

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The purple-naped lory (Lorius domicella) is confined to Seram, Ambon and perhaps Haruku and Saparua, where it is everywhere uncommon. The chattering lory (L. garrulus) is confined to Morotai, Rau, Halmahera, Widi, Ternate, Bacan, and Obi in the northern Moluccas. Both are threatened by overcollection for the cage-bird trade and by loss of habitat. Kuehn’s yellow white-eye (Zosterops kuehni) is confined to Ambon and Seram in the central Moluccas. Oort’s scaly toed gecko (Lepidodactylus oortii) is known from the Banda, Damar and Tanimbar islands in the southern Moluccas. Boulenger’s cylinder snake (Cylindrophis aruensis) is known only from the Aru Islands and Damar in the Barat Daya Islands. Rueppell’s big-eyed tree frog (Nyctimystes rueppelli) is confined to Halmahera and Morotai in the northern Moluccas, where it is threatened by loss of habitat.

Morotai Morotai (Pulau Morotai in Indonesian) is a rugged, forested island located north of Halmahera. The dusky friarbird (Philemon fuscicapillus) is a type of honeybird that was considered to be very common on Morotai prior to World War II, but is today threatened by the loss of its lowland forest habitat. The Moluccan bent-toed gecko (Cyrtodactylus deveti) is endemic to Morotai, where it is known only from a few specimens. Horst’s cross frog (Oreophryne frontifasciata) is known with certainty only from Morotai, but may occur on Halmahera, Selawati, and Geg as well.

Halmahera Halmahera (Pulau Halmahera in Indonesian) is the largest island in the Moluccas, and includes two active volcanoes. It was on here in 1869, between bouts of fever, that Alfred Russel Wallace formulated his own theory of evolution through natural selection. He wrote his ideas down in a letter that he later sent to Charles Darwin. The blue-eyed cuscus (Phalanger matabiru) is a type of arboreal marsupial confined to the offshore islands of Ternate and Tidore, where it is still relatively common although subject to hunting for food. The ashy-headed flying fox (Pteropus caniceps) is confined to Halmahera and the satellite islands of Btjan, Tidore, and Ternate. The drummer rail (Habroptila wallacii) is a flightless species known from only from a few specimens, although sightings have increased in recent years. The sombre kingfisher (Todiramphus funebris) is confined to Halmahera, where it is rare and localized. Weber’s sailfin lizard (Hydrosaurus weberi) is confined to Halmahera and Ternate. The Halmahera keelback (Tropidonophis punctiventris) is a type of snake known only from two specimens collected in 1895.

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The silver tree monitor (Varanus zugorum) is a mysterious species known only from a single specimen purportedly collected on Halmahera, as well as some possible photos taken by animal collectors. Boettger’s callulops frog (Callulops boettgeri) is known only from a single specimen collected in northern Halmahera in 1894. The dubious callulops frog (C. dubius) is similarly known only from its original collection during the late nineteenth century from an undefined locality in northern Halmahera. The Halmahera rainforest frog (Cophixalus montanus) is known only from a small area of northern Halmahera. Gebe Gebe (Pulau Gebe in Indonesian) is a small island located east of Halmahera. The Gebe cuscus (Phalanger alexandrae) is confined to Gebe, where it is still relatively common but threatened by hunting and habitat destruction.

The Sula Islands The Sula Islands (Kepulauan Sula in Indonesian) are a group located in the western Moluccas. The Sula jungle flycatcher (Cyornis colonus) is confined to Taliabu, Seho, Mangole, and Sanana. Taliabu Taliabu (Pulau Taliabu in Indonesian) is the largest island in the Sula group. The Taliabu masked owl (Tyto nigrobrunnea) is known only from a single specimen and a few subsequent sight records. Sanana Sanana is the southernmost of the Sula Islands. Kopstein’s callulops frog (Callulops kopsteini) is known only from a single specimen collected in 1930.

Obira Obira (Pulau Obira in Indonesian) is one of a small group of islands north of Buru and Seram known as the Obi Islands. The carunculated fruit dove (Ptilinopus granulifrons) is endemic to the island, where it remains common.

Buru Buru (Pulau Buru in Indonesian) is a large island to the west of Seram. Although its lowland forests have been largely degraded and cleared, two large, contiguous blocks remain. The Buru babirusa (Babyrousa babyrussa) is confined to Buru as well as on two of the Sula Islands (Mangole and Taliabu). It formerly also occurred on Sanana in the Sula Islands, but is believed to have been extirpated from there. The black-lored parrot (Tanygnathus gramineus) is a somewhat mysterious species that is rarely observed.

The Wallacean Region

The blue-throated lorikeet (Charmosyna toxopei) is confined to Buru, where it is known only from a few specimens and rarely observed. The rufous-throated white-eye (Madanga ruficollis) is known from four specimens collected prior to 1920, along with a few more recent records from remote mountainous areas.

Seram Seram (Pulau Seram in Indonesian; formerly known as Ceram) is the second largest island of the Moluccas. Manusela National Park, established in 1997, is located in the eastern part of the island. The Seram bandicoot (Rhynchomeles prattorum) is a type of terrestrial marsupial known only from seven specimens collected in 1920. The Seram mountain rat (Nesoromys ceramicus) is known from four specimens collected on Mount Mansuela. Four species of mosaic-tailed rat (Melomys) are endemic to Seram. The Seram mosaic-tailed rat (M. fulgens) is known only from two specimens collected in 1920. The Mansuela mosaic-tailed rat (M. fraterculus) is known only from a single specimen collected on Mount Mansuela in 1920. Pavel’s mosaic-tailed rat (M. paveli) is known only from a single specimen collected in 2003. The dusky mosaic-tailed rat (M. aerosus) is confined to a few montane areas on Seram. The Seram masked owl (Tyto almae) was first described in 2013, and known only from a couple of records. Boano Boano (Pulau Boano in Indonesian) is located at the western end of Seram. The black-chinned monarch (Symposiachrus boanensis) was long known only from a single specimen collected in 1918, but was rediscovered in 1991. It appears to be very rare and confined to a small area of mountain forest.

The Banda Islands The Banda Islands (Kepulauan Banda in Indonesian) are a group of 10 small volcanic islands in the Banda Sea, about 140 km south of Seram. The Banda Islands dtella (Gehyra barea) is a type of gecko endemic to the lowland rainforests of the Banda Islands.

The Kai Islands The Kai Islands (Kepulauan Kai in Indonesian) are located in the south-eastern Moluccas. The Kai flying fox (Pteropus keyensis) is confined to the Kai Islands. Great Kai Island Great Kai Island (Kai Besar in Indonesian) is the largest of the Kai Islands. It has been heavily logged. The Great Kai mosaic-tailed rat (Melomys bannisteri) is confined to Great Kai Island.

The Great Kai giant naked-tailed rat (Uromys siebersi) is confined to Great Kai Island.

The Barat Daya Islands The Barat Daya Islands (Kepulauan Barat Daya in Indonesian) are part of the same volcanic arc as the Banda Islands. Wetar Wetar (Pulau Wetar in Indonesian) is the largest of Barat Daya Islands. The Wetar bent-toed gecko (Cyrtodactylus wetariensis) is known only from a few specimens collected on Wetar. Damar Damar (Pulau Damar in Indonesian) is a small volcanic island flanked by four smaller islands. The Damar flycatcher (Ficedula henrici) was long known only from 14 specimens collected on Damar in 1898. Recent reports indicate that it remains relatively common.

The Tanimbar Islands The Tanimbar Islands are an archipelago of about 65 islands located in the central and southern Moluccas. The Tanimbar cockatoo (Cacatua goffiniana) is historically known from the islands of Yamdena, Larat, and Selaru in the Tanimbar Islands, where it is threatened by overcollection for the international pet trade, habitat destruction, persecution as a crop pest, and hunting for food. The species remains relatively common in any case, and has been introduced to the Kai Islands of Indonesia as well as to Puerto Rico and Singapore. Yamdena Yamdena is the largest of the Tanimbar Islands. Most of the original forest has been clear-cut. The Yamdena mosaic-tailed rat (Melomys cooperae) is known only from its original type series collected in 1995. The Saumlaki frog (Litoria capitula) is known only from its original collection in the 1960s. Riama Riama is a small island located south of Yamdena. The Riama mosaic-tailed rat (Melomys howi) is known only from Riama, but may occur on other islands within the Tanimbar Archipelago.

The Aru Islands The Aru Islands (Kepulauan Aru in Indonesian) are a group of about 95 low-lying, swampy islands located south-west of New Guinea in the Arafura Sea. Uniquely among the Moluccas they have a purely Papuan fauna including kangaroos, cassowaries, and birds of paradise. General threats include conversion of forests into sugar cane plantations. The Aru flying fox (Pteropus aruensis) has not been positively recorded since 1877 and is almost certainly extinct.

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Given its large size it is thought that hunting was the likely cause. The white-naped pheasant-pigeon (Otidiphaps aruensis) is confined to the Aru Islands, where it is considered to be rare. The black tree monitor (Varanus beccarii) is confined to the Aru Islands. The Aru gudgeon (Bostrichthys aruensis) is a fresh- and brackish water fish confined to the Aru Islands. Wokam Island Wokam Island (Tanah Besar in Indonesian) is one of the four main islands in the Aru Island group. The Wokam skink (Sphenomorphus rufus) is confined to Wokam Island.

The Lesser Sunda Islands The Lesser Sunda Islands (Kepulauan Nusa Tenggara – ‘southeastern archipelago’ – in Indonesian) consist of two geologically distinct archipelagos comprising numerous small islands, often divided by deep ocean trenches. The northern ‘inner’ group includes Bali (discussed in the Sundaic Region section), Lombok, Sumbawa, Flores and Wetar, and is volcanic in origin. The islands of the southern ‘outer’ group include Sumba, Timor, and Babar, and are non-volcanic. Movement of flora and fauna between the islands is limited, leading to the evolution of a high rate of localized species. The so-called Wallace’s Line, previously discussed, passes between Bali and Lombok and marks the boundary between the Sundaic and Wallacean zoogeographic regions. The Lesser Sunda rat (Komodomys rintjanus) is known only from Rintja, Padar, Lomblen, and Pantar, although it probably occurs on other islands in the Lesser Sunda chain. It is possibly under threat from introduced rats. The Lesser Sunda flying fox (Pteropus lombocensis) is known from Lombok, Sumbawa, Komodo, Flores, Lomblen, Pantar, Alor, and Timor. It is threatened by hunting, roost disturbance and habitat destruction. The Lesser Sunda fruit bat (Acerodon mackloti) is known from Lombok, Sumbawa, Sumba, Flores, Timor, Alor, and Moyo. It is threatened by hunting, roost disturbance and habitat destruction. The Lesser Sunda hawk-eagle (Nisaetus floris) is confined to Flores, Sumbawa, Lombok and Alor as well as two small satellite islands (Satonda near Sumbawa, and Rinca near Flores), with occasional vagrants on Komodo. The species is everywhere rare and seriously threatened by loss of habitat. The Lesser Sunda yellow-crested cockatoo (Cacatua sulphurea occidentalis) is confined to Lombok, Sumbawa, Komodo, Flores, Alor and various smaller islands, where it is threatened by overcollection for the international cage-bird trade and by loss of habitat. The Lesser Sunda green pigeon (Treron floris) is known from the islands of Sumbawa, Flores, Besar, Solor, Lembata, Pantar, and Alor, and may also occur on a few other islands as

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well, such as Komodo. The species is everywhere rare and threatened by loss of habitat and hunting. Hoedt’s ground dove (Pampusana hoedtii) is confined to Timor and Wetar, where it is threatened by habitat destruction and hunting. The Tenggara hill myna (Gracula venerata) is confined to the western Lesser Sunda Islands from Lombok and Sumbawa east to Pantar and Alor. It is heavily targeted by trappers throughout its range for use in the cage-bird trade. The Komodo monitor (Varanus komodoensis) is the world’s largest lizard, sometimes reaching 3 m in length. Found only on Komodo, Rinca, Gili Motang, Padar, and the north-western coastal part of Flores, this species has suffered declines over the past century mainly due to loss of habitat and hunters killing off its prey (generally rusa deer and wild boar). In 1966 all of Komodo Island, where the majority of the population is found, was declared a strict nature reserve, and has since been expanded into a national park encompassing it as well as Padar, Rinca, and 26 smaller islands. In recent years rising sea levels due to climate change has emerged as a new threat to the species. Schmutz’s worm snake (Typhlops schmutzi) is known only from the northern coast of Flores and Komodo. Everett’s frog (Litoria everetti) is known only from Timor, Sumba, Sawu, and Alor.

Lombok Lombok (Pulau Lombok in Indonesian) is a roughly circular island located east of Bali and west of Sumbawa. It is dominated by the highly active stratovolcano Mount Rinjani, the second highest volcano in Indonesia. The Lombok bent-toed gecko (Cyrtodactylus gordongekkoi) is known only from two specimens collected on Lombok in 1992.

Sumbawa Sumbawa is a grassy island located between Lombok and Flores. The Sumbawa sunset lorikeet (Trichoglossus forsteni forsteni) is confined to Sumbawa.

Flores Flores (Pulau Flores in Indonesian) is a large island located east of Sumbawa and north of Sumba and Timor. As already mentioned, the western coast of this island is one of the few places in the world where the Komodo monitor (Varanus komodoensis) is to be found in the wild, and is part of Komodo National Park. Hainald’s rat (Rattus hainaldi) is confined to Flores. Paula’s long-nosed rat (Paulamys naso) is confined to montane forest on Flores. The Flores pygmy white-toothed shrew (Suncus mertensi) is known only from a single specimen collected on western Flores in the 1970s. The Flores scops owl (Otus alfredi) was originally collected on Mount Repok in 1896 and not seen again until two

The Wallacean Region

specimens were collected in 1994 in the Ruteng Mountains. It is now seen fairly regularly by birdwatchers throughout Flores. The Flores hanging parrot (Loriculus flosculus) is confined to Flores and satellite islands. The Flores crow (Corvus florensis) is confined to Flores and Rinca. The Flores monarch (Symposiachrus sacerdotum) is confined to western Flores. The Flores ground snake (Stegonotus florensis) is known only from Flores, but may occur as well on Sumba and Semau. The Flores oriental frog (Occidozyga floresiana) is confined to Flores. The Flores cross frog (Oreophryne rookmaakeri) is confined to eastern Flores.

The Solor Archipelago The Solor Archipelago (Kepulauan Solor in Indonesian) is a group of islands lying to the east of Flores and west of the Alor Archipelago. Lembata Lembata (formerly known as Lomblen) is the largest island in the Solor Archipelago. The Lembata long-eared bat (Nyctophilus heran) is only known from a single specimen collected from Lembata.

The Alor Archipelago The Alor Archipelago (Kepulauan Alor in Indonesian) is located in the eastern Lesser Sunda Islands. Alor Alor (Pulau Alor in Indonesian, and also known as Ternate) is a rugged volcanic island and the largest of the Alor group. Johnstone’s mastiff bat (Otomops johnstonei) is known only from a single specimen collected on Alor Island in 1991.

Sumba Sumba (Pulau Sumba in Indonesian) is the westernmost of the southern group of Lesser Sunda Islands, and consists of low limestone hills. Most of the original forest has been cleared for the planting of crops, with only small isolated patches remaining. In 1998 two national parks were designated. The Sumba hornbill (Rhyticeros everetti) is confined to Sumba, where it is threatened by deforestation outside of protected areas. The Sumba yellow-crested cockatoo (Cacatua sulphurea citrinocristata) is confined to Sumba, where the total population was estimated at 562 in 2012. The Sumba buttonquail (Turnix everetti) is confined to dry, scrubby areas on Sumba. The red-naped fruit dove (Ptilinopus dohertyi) is confined to Sumba.

Timor Timor (Pulau Timor in Indonesian) is a large island at the eastern end of the Lesser Sunda archipelago southern group

and, like Sumba, non-volcanic. Much of the original dry deciduous forest has been cleared for agriculture and, apart from a large block in the centre of the island, only patches now remain. Buhler’s giant rat (Coryphomys buehleri) is known only from subfossil remains. One of four extinct giant rat species known to have lived on Timor, it is the only one to be formally described. The Timor forest rat (Rattus timorensis) is confined to montane forest on Timor. Although the subfossil record indicates it was formerly abundant, only a single living specimen has ever been collected (in 1990). The Timor shrew (Crocidura tenuis) is confined to Timor. The Timor leaf-nosed bat (Hipposideros crumeniferus) is confined to Timor. The Timor horseshoe bat (Rhinolophus montanus) is confined to Timor. The Timor yellow-crested cockatoo (Cacatua sulphurea parvula) is confined to Timor. The Timor green pigeon (Treron psittaceus) is confined to Timor and the satellite islands of Semau and Rote, where it is rare. McCord’s snake-necked turtle (Chelodina mccordi) is divided into two subspecies. The Timor snake-necked turtle (C. m. timorensis) is confined to Timor, where it is seriously threatened. The Timor skink (Eremiascincus timorensis) is confined to Timor, where it was first described in 1990. Roti Located south-west of Timor, Roti (Pulau Roti in Indonesian) is the southernmost island in Indonesia. Until recently, the Roti snake-necked turtle (Chelodina mccordi mccordi) occurred in three populations on this single small island. It is now extinct in the wild due to collection for the international pet trade, where it commands high prices.

Balance for the Wallacean Region There is evidence of early hominins living in what is now the Philippines as early as 709,000 years ago. The oldest modern human remains found on the islands are from the Tabon Caves of Palawan, which have been dated to around 47,000 years ago. This ‘Tabon Man’ was presumably a Negrito, who were among the archipelago’s earliest inhabitants and descendants of the first human migrations out of Africa via the coastal route along southern Asia to the now sunken landmasses of Sundaland and Sahul. The first Austronesians and perhaps other Asian peoples reached the Philippines around 2200 B . C . and quickly assimilated the existing Negritos. Other islands have a similar history. Early European exploration of the Wallacean Region was undertaken by Portuguese mariners. In 1511–12 Antonio de Abreu sailed through the Strait of Malacca between Sumatra

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Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

2 species

3 species

~ species

153 species

2 subspecies

~ subspecies

~ subspecies

12 subspecies

4 taxa

3 taxa

~ taxa

165 taxa

~ species

2 species

~ species

146 species

12 subspecies

2 subspecies

1 subspecies

21 subspecies

12 taxa

4 taxa

1 taxon

167 taxa

2 species

~ species

~ species

88 species

~ subspecies

~ subspecies

1 subspecies

2 subspecies

2 taxa

~ taxa

1 taxon

90 taxa

1 species

~ species

~ species

60 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

1 taxon

~ taxa

~ taxa

60 taxa

17 species

2 species

~ species

45 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

17 taxa

2 taxa

~ taxa

45 taxa

22 species

7 species

~ species

492 species

14 subspecies

2 subspecies

2 subspecies

35 subspecies

36 taxa

9 taxa

2 taxa

527 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

and Bangka, and along the coasts of Java, Bali, Lombok, Sumbawa, and Flores to the ‘Spice Islands’ (i.e. the Moluccas). In 1525 his countrymen reached ‘Celebes’ (modern Sulawesi), and explored the Moluccas around the same time. During the early 1520s Ferdinand Magellan and his Spanish expedition reached the Moluccas before continuing westward to what would eventually be the first circumnavigation of the globe. What is now the Philippines became a part of the Spanish East Indies during the sixteenth century. Since the independence of the countries within the region a massive surge in population and a corresponding increase in deforestation and hunting have been the hallmark. In recent historical time (i.e. since A . D . 1500), the Wallacean Region has lost at least 22 species/14 subspecies of vertebrates. Among the extinct forms 2 species/2 subspecies are mammals, 12 subspecies are birds, 2 species are reptiles, 1 species is an amphibian, and 17 species are freshwater fishes. Another 7 species/2 subspecies are possibly extinct, and 2 subspecies are currently extinct in the wild. In addition, there are 492 species/35 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of

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extinction). Of these, 153 species/12 subspecies are mammals, 146 species/21 subspecies are birds, 88 species/2 subspecies are reptiles, 60 species are amphibians, and 45 species are freshwater fishes.

Balance for the Indo-Malaysian Realm In recent historical time (i.e. since A . D . 1500), the IndoMalaysian Realm as a whole has lost at least 57 species/19 subspecies of vertebrates. Among the extinct forms 9 species/ 6 subspecies are mammals, 3 species/13 subspecies are birds, 6 species are reptiles, 20 species are amphibians, and 19 species are freshwater fishes. Another 24 species/3 subspecies are possibly extinct, and 3 species/2 subspecies are currently extinct in the wild. In addition, there are 1965 species/159 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 417 species/107 subspecies are mammals, 289 species/40 subspecies are birds, 330 species/12 subspecies are reptiles, 390 species are amphibians, and 539 species are freshwater fishes.

Balance for the Indo-Malaysian Realm

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

9 species

5 species

~ species

417 species

6 subspecies

1 subspecies

~ subspecies

107 subspecies

15 taxa

6 taxa

~ taxa

524 taxa

3 species

3 species

1 species

289 species

13 subspecies

2 subspecies

1 subspecies

40 subspecies

16 taxa

5 taxa

2 taxa

329 taxa

6 species

2 species

2 species

330 species

~ subspecies

~ subspecies

1 subspecies

12 subspecies

6 taxa

2 taxa

3 taxa

342 taxa

20 species

1 species

~ species

390 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

20 taxa

1 taxon

~ taxa

390 taxa

19 species

13 species

~ species

539 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

19 taxa

13 taxa

~ taxa

539 taxa

57 species

24 species

3 species

1965 species

19 subspecies

3 subspecies

2 subspecies

159 subspecies

76 taxa

27 taxa

5 taxa

2124 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

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Chapter

6

The Papua-Melanesian Realm

The Papua-Melanesian Zoogeographic Realm consists of the island of New Guinea along with its numerous satellites and the vast Melanesian archipelagos to the east. Primary among the latter are the Bismarck Archipelago, the Solomon Islands, Fiji, and New Caledonia. It shares affinities with Wallacean, Micronesian, and Australian fauna regions. Brown’s pademelon (Thylogale browni) is a small, wallabylike marsupial still widespread throughout north-central and north-eastern New Guinea, but heavily depleted by hunting. It was also introduced during prehistoric times to the islands of Bagabag, New Britain, New Ireland, and Umboi, and is known from the island of Yapen as well but has now possibly been extirpated there. The dusky pademelon (T. brunii) is found in south-central New Guinea, the Aru and Kai islands in the Moluccas, and (at least historically) in the grasslands surrounding Port Moresby in the south-east, although the latter population has most likely been extirpated. It is threatened mainly by hunting. McDonald’s long-tailed fruit bat (Notopteris macdonaldi) is confined to the Fiji Islands and Vanuatu, where its large cave-dependent colonies are threatened by hunting and disturbance. The species historically occurred in the Tonga Islands as well, but was extirpated following the arrival of Polynesians. Felten’s free-tailed bat (Chaerephon bregullae) is confined to a few roosting caves in Vanuatu (Malo and Espíritu Santo) and on Taveuni, Vanua Levu, and possibly Viti Levu in the Fiji Islands. The species formerly occurred in the Tonga Islands as well, but was extirpated there during prehistoric times. It is threatened by hunting and human disturbance. The collared petrel (Pterodroma brevipes) is a type of seabird widespread across the South Pacific during nonbreeding periods. The nominate subspecies (P. b. brevipes) is, however, currently known to breed only on Gau and Kadavu in the Fiji Islands. It formerly bred elsewhere in the Fiji Islands and on Raratonga in the Cook Islands, and may still breed on Makira in the Solomons, Tau in the Samoan Islands, and in other as-yet undiscovered localities in French Polynesia. It is threatened mainly by introduced rats, cats, and mongoose. Beck’s petrel (Pseudobulweria becki) was long known only from two specimens collected from the region of the Bismarck Sea, one taken east of New Ireland and north of Buka in 1928, and another the following year north-east of Rendova in the

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Solomon Islands. It was long feared extinct until 2003, when three birds most likely representing this species were observed off New Ireland. Others were observed in the same area in 2007, confirming the rediscovery, as well as in Vanuatu in 2010. It is thought that the birds breed in the mountains of New Ireland, although suitable habitat exists as well in the Solomons. The total population is believed to number only a few hundred. Heinroth’s shearwater (Puffinus heinrothi) is a type of seabird known from the Bismarck Archipelago and the seas around Bougainville and Kolombangara in the Solomon Islands. The few historical specimens are from Watom, near New Britain, suggesting breeding there. There have been no recent reports from this island, however, and only a few elsewhere. The palm lorikeet (Charmosyna palmarum) is confined to the Santa Cruz Islands in the Solomon Archipelago and to Vanuatu, where it is threatened by loss of habitat. The yellow-legged pigeon (Columba pallidiceps) is a rare species found patchily in the Bismarck and Solomon archipelagos, where it is threatened by loss of habitat. Mayr’s ground dove (Pampusana sanctaecrucis) is known for certain only from Tinakula in the southern Solomon Islands and from Espiritu Santo in Vanuatu. Most of the Tinakula population was destroyed in 2017 by a volcanic eruption, and the species continues to be under heavy threat from the caged-bird trade and introduced species. Mayr’s swiftlet (Aerodramus orientalis) is known for certain only from three specimens collected, respectively, from Guadalcanal in 1927, New Ireland in 1963, and Bougainville in 1979. Sightings of unidentified swiftlets have since occurred on New Britain, New Ireland and New Georgia, and may refer to this species. The little paradise kingfisher (Tanysiptera hydrocharis) is a rare species known only from a small number of localities in southern coastal New Guinea, the Aru Islands in the Moluccas, and possibly from islands in the Torres Strait. The sweetwater lyretail demoiselle (Neopomacentrus aquadulcis) is a type of fresh- and brackish-water fish known only from two localities in eastern Papua New Guinea and from Tetepare Island in the Solomons. It is threatened by habitat degradation. Guilbert’s loach goby (Rhyacichthys guilberti) was historically known from one river each on Grande Terre (New

New Guinea

Caledonia) and on Vanuatu, but may have been extirpated from the former. The Pentecost sicyopus goby (Smilosicyopus pentecost) is known from Pentecost Island and Malakula Island in Vanuatu, and from Grande Terre, New Caledonia. The slender pipefish (Microphis caudocarinatus) is known only from two specimens, the first collected from the Tawarin River of north-western New Guinea in 1903, and the other from Sumbawa in the Lesser Sunda Islands in 1995.

New Guinea Located to the north of Australia just below the equator, New Guinea is the world’s second largest island (after Greenland). Its shape has often been compared to that of a bird of paradise (one of the many groups largely endemic to the island), with the so-called Bird’s Head Peninsula (Vogelkop in Dutch) in the north-west and the Bird’s Tail Peninsula (also known as the Papuan Peninsula) in the south-east. A spine of east–west mountains, the New Guinea Highlands, are bordered to the north and south by vast lowlands containing areas of rainforest, grasslands, wetlands, and mangroves. Politically it is divided more or less equally between Indonesia (Papua and Papua Barat provinces, formerly Irian Jaya) and the independent country of Papua New Guinea to the east. New Guinea has been geographically isolated from Asia by water at least since the Tertiary Period, that is, about 60 million years. The break with Australia took place relatively recently, however, and the island accordingly functioned as a bridge for animal migration between the two. New Guinea is differentiated from its drier, flatter, and less-fertile neighbour by its much higher rainfall and active volcanism. Yet the two landmasses share a similar vertebrate fauna, among them marsupials such as wallabies and possums as well as the egg-laying monotremes, the echidnas; cassowaries (large ratite birds); reptiles, amphibians, and fish. Other than bats and some two-dozen endemic rodent genera there are no pre-human placental mammals. Despite having less than 0.5 per cent of the Earth’s total land surface New Guinea is thought to contain between 5 and 10 per cent of total species on the planet – an immense biodiversity comparable only to the Amazon or Madagascar. Moreover, it is one that, until just recently, remained relatively untouched, in spite of a human presence stretching back at least 40,000 years. The European colonists brought with them pigs, and new species of rat. The introduction of the cane toad (Rhinella marina) has resulted in the eradication of native lizards, while exotic carp and trout have exterminated many indigenous fishes. Several species of tree-kangaroo (Dendrolagus) are threatened by hunting and habitat destruction. Doria’s treekangaroo (D. dorianus) is confined to mid-elevation and montane forests in south-eastern Papua New Guinea. The ursine tree-kangaroo (D. ursinus) is known from the Vogelkop Peninsula and possibly the Fakfak Peninsula in western New Guinea (Papua Barat province, Indonesia). The grizzled tree-

kangaroo (D. inustus) is divided into two subspecies. The western grizzled tree-kangaroo (D. i. inustus) occurs in north-western coastal New Guinea on the Vogelkop Peninsula, the Fakfak Peninsula and the islands of Yapen, Waigeo, Misool Salawati, and possibly Batanta. Finsch’s grizzled tree-kangaroo (D. i. finschi) has a range centered on the northern coastal ranges of north-central coastal New Guinea. The ornate tree-kangaroo (D. goodfellowi) also has two subspecies (D. g. goodfellowi and D. g. buergersi) found widely but patchily in central Papua New Guinea. The black-spotted cuscus (Spilocuscus rufoniger) is an arboreal marsupial known historically from much of northern New Guinea, but now found only sporadically. It is threatened by hunting and habitat destruction. The southern groove-tailed moss-mouse (Microhydromys argenteus) is known only from three localities across a wide area of central and south-eastern Papua New Guinea. The mantled mastiff bat (Otomops secundus) is known only from six specimens collected from three localities in Papua New Guinea. The round-eared tube-nosed fruit bat (Nyctimene cyclotis) is known only from the Arfak Mountains and Mansuar Island in north-western New Guinea (Papua Barat province, Indonesia). The Papuan eagle (Harpyopsis novaeguineae) is found throughout New Guinea but in very low densities. It is threatened by loss of habitat and, in some areas, hunting. The chestnut-shouldered goshawk (Erythrotriorchis buergersi) is a rare species known only from a handful of records originating mainly from eastern New Guinea, although only one since the 1980s. Pesquet’s parrot (Psittrichas fulgidus) is still found across New Guinea, but is generally rare and has been extirpated from many parts of its former range due to hunting. The species is greatly in demand for its feathers in particular, which are used in ornamentation, but the meat is also consumed and in some areas the skins make up part of bride-price payments. The western crowned pigeon (Goura cristata) is found patchily in western New Guinea including the islands of Misool, Waigeo, Salawati and Batanta in the Raja Ampat Islands. The species also occurs on Seram in the Moluccas, where it was almost certainly introduced. At one time common, it has been extirpated from many areas mainly due to hunting for its meat and plumes. The Papuan swiftlet (Aerodramus papuensis) is known only from a handful of specimens collected over a wide area of northern and eastern New Guinea. Salvadori’s owlet-nightjar (Aegotheles affinis) is divided into two little-known subspecies. The western Salvadori’s owlet-nightjar (A. a. affinis) is known only from the Vogelkop Mountains in western New Guinea (Papua Barat province). The eastern Salvadori’s owlet-nightjar (A. a. terborghi) was known only from a single specimen collected in 1964 from east-central Papua New Guinea, but was sighted in 2016 in the Karimui Basin. Owlet-nightjars are naturally

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rare and cryptic, and it is likely that both subspecies range more widely. The blue-black kingfisher (Todiramphus nigrocyaneus) is a rare species found widely but patchily in northern, western and southern New Guinea and on the islands of Salawati, Batanta and Yapen. It is threatened by loss of habitat. The Fly River grassbird (Poodytes albolimbatus) is a type of warbler known only from a few wetland localities in southcentral New Guinea. Leopold’s dtella (Gehyra leopoldi) is a type of gecko known only from two localities on the Vogelkop Peninsula of western New Guinea (Papua Barat province, Indonesia). Oudeman’s dtella (G. interstitialis) is known only from a single specimen collected during the late nineteenth century from an undefined locality in western New Guinea (Papua province, Indonesia). Ahl’s emo skink (Emoia ahli) is known only from its original collection in 1932 from the upper Sepik River drainage in north-western Papua New Guinea. Cogger’s emo skink (E. coggeri) is known only from a small coastal strip in eastern Papua New Guinea. Vogt’s skink (Papuascincus phaeodes) is known only from its original collection in 1932 from the upper Sepik River drainage in north-western Papua New Guinea. The Setekwa River worm-eating snake (Toxicocalamus grandis) is known only from a single specimen collected in 1914 from an undefined locality in western New Guinea (Papua province, Indonesia). Jeud’s frog (Litoria jeudii) is known only from a single specimen collected in 1901 from an undefined locality in northern Papua New Guinea. The Spartacus frog (L. spartacus) is known only from a small area of west-central Papua New Guinea. Parker’s land frog (Austrochaperina parkeri) is known only from three localities in south-eastern coastal Papua New Guinea. The Morobe land frog (A. polysticta) is known only from a single specimen collected in 1901 from an imprecise locality on the Huon Peninsula in north-eastern Papua New Guinea. Gunnar’s narrow-mouthed frog (Choerophryne gunnari) is known only from a small area of south-eastern Papua New Guinea.

The New Guinea Highlands The New Guinea Highlands stretch for more than 1600 km from the ‘head’ to the ‘tail’ of the island, and dominate its geography. Many mountains rise to elevations of over 4000 m, and the tallest reach 4884 m and feature permanent glaciers. Apart from the central ranges various other, smaller ones occur both north and south. The luxurious, closed-canopy montane rainforests gradually give way at higher elevations to a belt dominated by southern beeches (Nothofagus), which in turn is replaced by mossy cloud forests, a subalpine zone dominated by rhododendrons, and finally by alpine tussock grasslands.

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The golden-mantled tree-kangaroo (Dendrolagus pulcherrimus) was historically fairly widespread in the mountains of northern New Guinea, but is now known for certain only from to two disjunct areas (the Torricelli Mountains in Papua New Guinea, and the Foja Mountains in Papua Barat province, Indonesia). There have also been unconfirmed reports from the Prince Alexander Mountains in Papua New Guinea. Seri’s tree-kangaroo (D. stellarum) is confined to the mountains of west-central New Guinea (Papua province, Indonesia and western Papua New Guinea). Scott’s tree-kangaroo (D. scottae) is confined to the Torricelli and Bewani mountains of north-western Papua New Guinea. The dingiso tree-kangaroo (D. mbaiso) is confined to the Tembagapura and Kwiyawagi mountains of west-central New Guinea (Papua province, Indonesia). The Huon tree-kangaroo (D. matschiei) is naturally confined to the mountains of the Huon Peninsula in north-eastern Papua New Guinea. A population on the island of Umboi in the Bismarck Archipelago was almost certainly introduced. The Ifola tree-kangaroo (D. notatus) is confined to the mountains of central Papua New Guinea. Calaby’s pademelon (Thylogale calabyi) is thought to have been historically widespread within the mountains of Papua New Guinea, but is now known only from one or perhaps two widely separated relic populations (Mount Albert Edward and Mount Giluwe). It has not been collected from the latter since the 1960s, and appears to have been extirpated from Mount Wilhelm sometime during the twentieth century. The decline of both species is thought to be due to feral pigs and hunting with dogs. The Huon pademelon (T. lanatus) is confined to the mountains of the Huon Peninsula in north-eastern Papua New Guinea. Menzies’ spiny bandicoot (Echymipera echinista) is a type of terrestrial marsupial known for certain only from two disjunct localities in central-western and south-western Papua New Guinea. There is a third, unconfirmed record from Mount Menawi. The Telefomin cuscus (Phalanger matanim) is an arboreal marsupial known only from six specimens collected from a mountainous area of forest in central-western Papua New Guinea, which was largely destroyed by fire in 1998. It is now most likely extinct. Echidnas of the genus Zaglossus are spiny monotremes, egg-laying mammals whose closest living relative is the platypus (Ornithorhynchus anatinus). All are seriously threatened by hunting. The western long-beaked echidna (Z. bruijnii) is known only from the Vogelkop Peninsula of western New Guinea and the adjacent land bridge island of Salawati. It is possible that the species also occurs on the islands of Batanta and Waigeo. Interestingly, a specimen was also reportedly collected in north-western Australia in 1901, although there is no evidence for its continued survival there. The eastern long-beaked echidna (Z. bartoni) is divided into four subspecies found over much of central and eastern New Guinea. The nominate form (Z. b. bartoni) is found in the highlands of central-eastern Papua New Guinea. The Huon

New Guinea

long-beaked echidna (Z. b. clunius) is confined to the Huon Peninsula of eastern Papua New Guinea. Diamond’s longbeaked echidna (Z. b. diamondii), the largest living monotreme, is found throughout the mountains of central New Guinea from the Paniai Lakes of Papua province, Indonesia to the Kratke Mountains of eastern Papua New Guinea. Tate’s three-striped dasyure (Myoictis wavicus) is a small, carnivorous marsupial known only from a handful of specimens collected from a small area of eastern Papua New Guinea. The glacier rat (Rattus richardsoni) is known only from the higher elevations of Mount Wilhelmina, Mount Jaya and the area around Lake Habbema in west-central New Guinea (Papua province, Indonesia), a range corresponding to the last remaining areas of glacial cover on the island. Arianus’ rat (R. omichlodes) is confined to a small area of west-central New Guinea (Papua province, Indonesia). Van Deusen’s rat (R. vandeuseni) is found patchily in the mountains of southeastern Papua New Guinea. Gressitt’s mosaic-tailed rat (Paramelomys gressitti) is known only from two localities in eastern Papua New Guinea, where it is threatened by mining activities and wood collection. The alpine woolly rat (Mallomys gunung) is known only from two high-elevation localities in west-central New Guinea (Papua province, Indonesia). Husson’s water rat (Hydromys hussoni) is known only from the Wissel Lakes area of western New Guinea (Papua province, Indonesia). The mountain water rat (Baiyankamys habbema) is known only from a small area between Lake Habbema and Mount Wilhelmina in west-central New Guinea (Papua province, Indonesia). The short-haired water rat (Paraleptomys wilhelmina) is known only from three widely separated localities in the Central Highlands of New Guinea. The northern water rat (P. rufilatus) is known only from the Cyclops, Princess Alexandra and Bewani mountains of central-northern coastal New Guinea. Champion’s tree mouse (Pogonomys championi) is known only from the Telefomin and Tifalmin valleys of west-central Papua New Guinea. The white-toothed mouse (Coccymys albidens) is known only from a few individuals collected in 1938 from Mount Wilhelmina and Lake Habbema in western New Guinea (Papua province, Indonesia). The western moss-mouse (Pseudohydromys occidentalis) is known only from two disjunct areas in the Central Highlands. The first is found in the region of Lake Habemma and Mount Trikora in the Snow Mountains of west-central New Guinea (Papua province, Indonesia), and the other in the Star and Victor Emmanual mountains of central-western Papua New Guinea. The northern groove-toothed moss-mouse (Microhydromys richardsoni) is known only from a small number of specimens collected over a wide area of northern New Guinea.

Bulmer’s fruit bat (Aproteles bulmerae) is a cave-roosting species known only from two widely separated localities in the mountains of Papua New Guinea. It is threatened by habitat destruction and hunting. The small-toothed long-eared bat (Nyctophilus microdon) is known only from a few localities in west-central and central Papua New Guinea. The Snow Mountain quail (Anurophasis monorthonyx) is confined to alpine grasslands atop the highest peaks of the Snow and Star mountains in west-central New Guinea (Papua province, Indonesia). Mayr’s forest rail (Rallicula mayri) is known only from a handful of specimens collected from the coastal mountain ranges of north-central coastal New Guinea. The blue bird-of-paradise (Paradisornis rudolphi) occurs patchily in the central and south-eastern mountains of Papua New Guinea, from Mount Sisa to the Owen Stanley Mountains. It is everywhere uncommon. MacGregor’s honeyeater (Macgregoria pulchra) is a large, crow-like bird that was historically widespread within the Central Highlands, but is now reduced to a few disjunct populations on the island’s highest mountains due to heavy hunting pressure. The long-bearded honeyeater (Melidectes princeps) is found patchily in central Papua New Guinea. The Papuan whipbird (Androphobus viridis) is a rare and cryptic passerine bird known from a handful of sites across the Central Highlands. The Papua Barat forest dragon (Hypsilurus tenuicephalus) is known only from a single specimen exported for commerce, reportedly collected somewhere within the highlands of western New Guinea (Papua Barat province, Indonesia). The Bundi bent-toed gecko (Cyrtodactylus arcanus) is known only from a small area of eastern Papua New Guinea. The mid-slope bent-toed gecko (C. medioclivus) is known only from a small area of central-western Papua New Guinea. The Wissel Lakes emo skink (Emoia paniai) is confined to an area of the Central Highlands in western New Guinea (Papua Barat province, Indonesia). The Wanagong skink (Lobulia glacialis) is confined to the area of Lake Wanagong in west-central New Guinea (Papua province, Indonesia). A number of frogs of the genus Litoria endemic to small, isolated montane areas. Pratt’s frog (L. pratti) is known only from two localities in north-western New Guinea (Papua Barat province, Indonesia). The Wapoga Alpha frog (L. scabra), Mack’s frog (L. macki), Wapoga River frog (L. wapogaensis) and the Derewo frog (L. fuscula) are each known only from a single locality in north-western New Guinea (Papua province, Indonesia). The Baliem Valley frog (L. umbonata), Brongersma’s tree frog (L. brongersmai), and the Wissel Lakes frog (L. wisselensis) are each known only from a small area of west-central New Guinea (Papua province, Indonesia). The West Sepik frog (L. leucova), the Telefomin frog (L. dorsivena), and Zweifel’s frog (L. zweifeli) are each known

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only from a small area of west-central Papua New Guinea. The Southern Highlands frog (L. kumae) and Beck’s frog (L. becki) are both known only from a small area of central Papua New Guinea. Sauron’s frog (L. sauroni) is known only from two localities in south-central Papua New Guinea. The Trauna frog (L. oenicolen) is known only from the Baiyer River Sanctuary in north-central Papua New Guinea. The Agaun frog (L. ollauro) is known only from a single locality in south-eastern Papua New Guinea. The Wapoga frog (L. megalops) is known only from a single locality in northwestern New Guinea (Papua province, Indonesia). The Mondo big-eyed tree frog (Nyctimystes gularis) is known only from Mount Obree and Mount Mondo in south-eastern Papua New Guinea. Grant’s big-eyed tree frog (N. granti) is known only from a single locality in west-central New Guinea (Papua province, Indonesia). Tyler’s big-eyed tree frog (N. tyleri) is known only from a single locality in south-eastern Papua New Guinea. Kuduk’s big-eyed tree frog (N. kuduki) is known only from a small area of west-central Papua New Guinea. The Wapoga Mawatta frog (Hylophorbus sextus) is known only from the headwaters of the Wapoga River in northwestern New Guinea (Papua province, Indonesia). The Huon Mawatta frog (H. rainerguentheri) is confined to the Huon Peninsula in north-eastern Papua New Guinea. Several species of fanged frog (Xenorhina) are endemic to small montane localities. The Wapoga fanged frog (X. macrodisca) is known only from a single locality in north-western New Guinea (Papua province, Indonesia). The giant fanged frog (X. gigantea), Parker’s fanged frog (X. minima), the Okbap fanged frog (X. multisica), the ocellated fanged frog (X. ocellata) and Scheepstra’s fanged frog (X. scheepstrai) are all confined to small areas of west-central New Guinea (Papua province, Indonesia). The Munggona fanged frog (X. schiefenhoeveli) and the Eipomek Valley fanged frog (X. eiponis) are both confined to the Eipomek Valley of central New Guinea (Papua province, Indonesia). The Huon fanged frog (X. huon) is known only from a small area of the Huon Peninsula in north-eastern Papua New Guinea. The warty callulops frog (Callulops glandulosus) is known only from Mount Kerewa, in central Papua New Guinea. The Idenburg cross frog (Oreophryne idenburgensis) is known only from a single locality in north-central New Guinea (Papua province, Indonesia). The Lake Habbema cross frog (O. habbemensis) and the dwarf cross frog (O. minuta) are each known only from a single locality in westcentral New Guinea (Papua province, Indonesia). The Bima cross frog (O. alticola) is known only from a small area of central New Guinea (Papua province, Indonesia). The Mount Michael rainforest frog (Cophixalus nubicola) is known only from Mount Michael in east-central Papua New Guinea. The Kaindi rainforest frog (C. kaindiensis) is known only from Mount Kaindi in eastern Papua New Guinea. The Arau Mawatta frog (Hylophorbus infulata) is known only from three disjunct montane localities in north-eastern Papua New Guinea.

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The Kosarek land frog (Austrochaperina kosarek) is known only from a single specimen collected in 1979 from westcentral New Guinea (Papua province, Indonesia). Mehely’s land frog (A. mehelyi) is known only from the mountains of north-eastern Papua New Guinea. The adamantine land frog (A. adamantina) is known only from the Bewani and Torricelli mountains of north-western coastal Papua New Guinea. The red land frog (Sphenophryne rubra) is known only from a few specimens collected in east-central Papua New Guinea. The yellow-throated Papua frog (Barygenys flavigularis) is known only from a small area of eastern Papua New Guinea. The spotted Papua frog (B. maculata) is known only from a small area of south-eastern Papua New Guinea. The Namosado barred frog (Mixophyes hihihorlo) is known only from a single locality in western Papua New Guinea. Burton’s narrow-mouthed frog (Choerophryne burtoni) is known only from a small area of west-central Papua New Guinea. Fafnir’s narrow-mouthed frog (C. fafniri) is known only from three localities in central Papua New Guinea. The Moiyokabip narrow-mouthed frog (C. rhenaurum) is known only from a single locality in central-western Papua New Guinea. The Elimbari narrow-mouthed frog (C. siegfriedi) is known only from Mount Elimbari in central Papua New Guinea. The long-nosed narrow-mouthed frog (C. longirostris) is known only from the Bewani and Torricelli mountains of north-western coastal Papua New Guinea. The Arfak Mountains The Arfak Mountains (Pegunungan Arfak in Indonesian) are located on the Vogelkop (‘Bird’s Head’) Peninsula in northwestern New Guinea (Papua Barat province, Indonesia). The Arfak pygmy bandicoot (Microperoryctes aplini) is known only from four specimens collected in the Arfak Mountains. Schlegel’s ringtail (Pseudochirulus schlegeli) is a type of possum known only from a handful of specimens collected in the Arfak Mountains. The reclusive ringtail (Pseudochirops coronatus) is known only from a few localities within the Arfak Mountains. The Vogelkop mountain rat (Rattus arfakiensis) is known only from specimens collected in the Arfak Nature Reserve in the 1930s. The grey-banded manakin (Lonchura vana) is a type of finch known for certain only from grasslands around Lake Anggi Gigi, although it may occur more widely in the Arfak Mountains. The Arfak forest dragon (Hypsilurus bruijnii) is confined to the Arfak Mountains. The Arfak frog (Litoria chloronota) is known only from two localities in the Arfak Mountains. The Arfak big-eyed tree frog (Nyctimystes montanus) is known only from a single specimen collected in 1878 from the Arfak Mountains.

New Guinea

The Hatam fanged frog (Xenorhina ophiodon) is known only from its original collection in 1878 from the Arfak Mountains. The Wakwam fanged frog (X. arfakiana) is known only from a single locality within the Arfak Mountains. The Wondiwoi Mountains The Wondiwoi Mountains (Pegunungan Wondiwoi in Indonesian) are located on the Wandammen Peninsula in north-western New Guinea (Papua Barat province, Indonesia). The Wondiwoi tree-kangaroo (Dendrolagus mayri) was until recently known only from a single specimen collected in 1928 and was feared extinct. In 2018, however, a tree-kangaroo fitting the description of the type specimen was photographed in a previously unexplored area of the Wondiwoi Mountains. The Wondiwoi frog (Litoria mareku) and Vera’s frog (L. verae) are each known only from a single locality within the Wondiwoi Mountains. The greater Mehely frog (Copiula major) and Obst’s Mehely frog (C. obsti) are each known only from a single locality within the Wondiwoi Mountains. The Wondiwoi Mawatta frog (Hylophorbus wondiwoi) is known only from a small area of the Wondiwoi Mountains.

Figure 6.1 Illustration of a Wondiwoi tree-kangaroo from 1936, based on the type specimen. (Credit: Rothschild & Dollman.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The noisy cross frog (Oreophryne clamata), unicolor cross frog (O. unicolor) and the hissing cross frog (O. sibilans) are each known only from a single locality within the Wondiwoi Mountains. The Wondiwoi rainforest frog (Cophixalus tridactylus) is known only from the eastern slopes of the Wondiwoi Mountains. The Wondiwoi callulops frog (Asterophrys pullifer) is known only from the eastern slopes of the Wondiwoi Mountains. The Wondiwoi wrinkled ground frog (Cornufer bimaculatus) is known only from a single locality within the Wondiwoi Mountains. The pygmy land frog (Austrochaperina minutissima) is known only from two localities within the Wondiwoi Mountains. The Wondiwoi narrow-mouthed frog (Choerophryne laurini) is known only from a single locality within the Wondiwoi Mountains. Specimens collected from the Maoke Mountains are thought to represent an as-yet undescribed species. The Maoke Mountains The Maoke or Snow Mountains (Pegunungan Maoke in Indonesian) are located in western and central New Guinea (Papua Barat and Papua provinces, Indonesia). Woolley’s moss-mouse (Pseudohydromys patriciae) is known only from a single locality within the Maoke Mountains (Papua province, Indonesia). Archbold’s robin (Petroica archboldi) is confined to the highest peaks of the Maoke Mountains. The Weyland Range is located in western New Guinea (Papua Barat province, Indonesia). The mouse bandicoot (Microperoryctes murina) is a type of terrestrial marsupial known only from three specimens collected in 1931 from the summit of Mount Sumuri, in the Weyland Range. The lesser small-toothed rat (Macruromys elegans) is only known from four specimens collected on Mount Kunupi in 1931. Stein’s mosaic-tailed rat (Paramelomys steini) is known only from six specimens collected from Mount Weyland. The Sudirman Range is located in west-central New Guinea (Papua province, Indonesia). Notable peaks include Puncak Trikora (known as Wilhelminatop or Mount Wilhelmina until 1963, whose glacier disappeared completely sometime between 1939 and 1962), and Mount Carstensz. The Tembagapura fanged frog (Xenorhina adisca) is known only from the Sudirman Range. The Went Range is located in west-central New Guinea (Papua province, Indonesia). The grey swamp frog (Papurana grisea) is known only from the Went Range. The Cyclops Mountains The Cyclops Mountains (Pegunungan Cyclops in Indonesian) are a small, coastal range located in north-western New

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Guinea, north of Lake Sentani (Papua province, Indonesia). They have been extensively deforested since World War II. Attenborough’s long-beaked echidna (Zaglossus attenboroughi) is known from a single specimen collected in the Cyclops Mountains in 1961. An expedition in 2007 found evidence of its existence (i.e. recent digging activity and burrows), although none were actually sighted. The Cyclops Mountains emo skink (Emoia cyclops) is known from a number of specimens collected in the Cyclops Mountains in 1938, but not reported since. Cheesman’s wrinkled ground frog (Cornufer cheesmanae) is known only from the foothills of the Cyclops Mountains, where it has not been reported in many years. The Star Mountains The Star Mountains (Pegunungan Bintang in Indonesian) are located in central New Guinea, where they straddle the border between Papua province, Indonesia, and western Papua New Guinea. The Majikthise frog (Litoria majikthise) is known only from a small area around the headwaters of the Ok Tedi River in the Star Mountains. The Ok Tedi frog (L. oktediensis) is confined to the southern slopes of the Star Mountains. The Star Mountains fanged frog (Xenorhina anorbis) is known only from two localities within the Star Mountains. The Star Mountains callulops frog (Callulops sagittatus) is known only from two localities within the Star Mountains. The twin cross frog (Oreophryne geminus) and the terrestrial cross frog (O. terrestris) are both known only from Dokfuma Meadow, south of Mount Capella in the Star Mountains. The Bewani Mountains The Bewani Mountains are located in north-western Papua New Guinea. Hill’s leaf-nosed bat (Hipposideros edwardshilli) is known only from a small area of the Bewani Mountains. The Menawa rainforest frog (Cophixalus bewaniensis) is known only from two specimens collected from Mount Menawa in the Bewani Mountains. The Torricelli Mountains The Torricelli Mountains are located in north-western coastal Papua New Guinea, between the Bewani Mountains to the west and the Prince Alexander Mountains to the east. The northern glider (Petaurus abidi) is a type of marsupial confined to Mount Somoro and adjacent areas in the Torricelli Mountains. Musser’s shrew mouse (Pseudohydromys musseri) is known only from a single specimen collected on Mount Somoro in 1972. The Torricelli land frog (Austrochaperina aquilonia) is known only from two specimens collected in the Torricelli ountains. The Prince Alexander Mountains The Prince Alexander Mountains are located in northern coastal Papua New Guinea.

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Ziegler’s water rat (Hydromys ziegleri) is a semi-aquatic species known only from the southern slopes of the Prince Alexander Mountains. The Hunstein Mountains The Hunstein Mountains are located in north-western Papua New Guinea. The pretty rainforest frog (Cophixalus pulchellus) is known only from Mount Hunstein. The Great Papuan Plateau The Great Papuan Plateau is a karst plateau located in westcentral Papua New Guinea. It is bordered by the upper stretches of the Kikori River on the east, the Strickland River on the west, the Karius Range to the north, and Mount Basavi to the south. Mount Bosavi is the collapsed cone of an extinct volcano on the Great Papuan Plateau. The Bosavi forest dragon (Hypsilurus ornatus) is a type of lizard known only from Mount Bosavi. The Karius Mountains The Karius Mountains are located in west-central Papua New Guinea. Mount Sisa is a heavily eroded, double-peaked stratovolcano. The white-bellied moss-mouse (Pseudohydromys sandrae) is known only from a single specimen collected on Mount Sisa. Allison’s narrow-mouthed frog (Choerophryne allisoni) is known only from a few specimens collected on Mount Sisa. The Crater Mountains The Crater Mountains are located in south-central Papua New Guinea. The Crater Mountains callulops frog (Callulops marmoratus) is known only from specimens collected in the vicinity of the Crater Mountain Biological Station. The Adelbert Mountains The Adelbert Mountains are an isolated coastal range located in north-eastern coastal Papua New Guinea. Gudrun’s narrow-mouthed frog (Choerophryne gudrunae) is known only from a single locality in the Adelbert Mountains. The small Papua frog (Barygenys parvula) is known only from two localities in the Adelbert Mountains. The Kratke Mountains The Kratke Mountains are located in eastern Papua New Guinea. They run south of the Finisterre Mountains on the other side of the Markham River and join the Bismarck Mountains to the east. The spotted worm-eating snake (Toxicocalamus spilolepidotus) is known only from two specimens collected in the Kratke Mountains in the 1960s. Archbold’s land frog (Austrochaperina archboldi) is known only from a type series collected in the Kratke Mountains.

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The Finisterre Mountains The Finisterre Mountains are located on the Huon Peninsula in north-eastern Papua New Guinea. The Finisterre bent-toed gecko (Cyrtodactylus minor) is known only from three specimens collected in the foothills at the northern edge of the Finisterre Mountains. The Finisterre frog (Litoria singadanae) is known from three specimens collected in 2005 in the eastern Finisterre Mountains.

The Cloudy Mountains The Cloudy Mountains are located near Milne Bay in far south-eastern Papua New Guinea. The Cloudy Mountains worm-eating snake (Toxicocalamus pachysomus) is known only from a single locality within the Cloudy Mountains. The Cloudy Mountains frog (Litoria rubrops) is known only from a few specimens collected in the Cloudy Mountains.

The Bartholomew Mountains The Bartholomew Mountains are located in south-eastern Papua New Guinea. The Bartholomew skink (Sphenomorphus annectans) is known only from the Bartholomew Mountains.

The lowland rainforests of New Guinea occur in three main subregions, located north and south of the Central Highlands and on the Vogelkop Peninsula, respectively. The dragon tube-nosed fruit bat (Nyctimene draconilla) is known from a handful of specimens collected from few disjunct localities on either side of the Central Highlands.

The Owen Stanley Mountains The Owen Stanley Mountains are located in south-eastern Papua New Guinea. German’s one-toothed moss-mouse (Pseudohydromys germani) is known only from the Owen Stanley Mountains. The Milne big-eyed tree frog (Nyctimystes persimilis) is known only from Mount Dayman and Mount Simpson in the Owen Stanley Mountains. The Dayman big-eyed tree frog (N. daymani) is known only from Mount Dayman. The Simpson rainforest frog (Cophixalus timidus) is confined to the northern slope of Mount Simpson where it is seriously threatened by frequent, deliberately set fires. The Myola rainforest frog (C. verecundus) is known only from a small area in the Owen Stanley Mountains. Dayman’s rainforest frog (C. daymani) and Zweifel’s rainforest frog (C. cryptotympanum) are both known only from Mount Dayman. The Victoria land frog (Austrochaperina brevipes) is known only from Mount Victoria and Myola Guest House in the Owen Stanley Mountains. It was last recorded in 1987. The Milne land frog (Sphenophryne crassa) is known only from Mount Dayman and Mount Simpson in the Owen Stanley Mountains. The Myola land frog (S. similis) is known only from a single locality in the Owen Stanley Mountains. The rosy-fingered land frog (S. rhododactyla) is found patchily throughout the Owen Stanley Mountains. The Tafa Papua frog (Barygenys cheesmanae) is known only from Mount Tafa. Sabin’s cross frog (Aphantophryne sabini) is known only from the region of the Myola Guest House on Mount Kenevi. The red-dotted narrow-mouthed frog (Choerophryne sanguinopicta) is confined to the northern slope of Mount Simpson, where it is threatened by frequent, deliberately set fires. The Nanneau Mountains The Nanneau Mountains are located in south-eastern coastal Papua New Guinea. The Nanneau long-beaked echidna (Zaglossus bartoni smeenki) is confined to the Nanneau Mountains.

Lowland Rainforests and Swamp Forests

Vogelkop Peninsula Lowland Rainforests and Swamp Forests The Vogelkop Peninsula lowland rainforests and swamp forests are located in north-western New Guinea (Papua Barat and Papua provinces, Indonesia). The Cenderawasih forest dragon (Hypsilurus nigrigularis) is an agamid lizard known only from a single locality on the southern coast of Cenderawasih Bay in Papua province, Indonesia. It is threatened by hunting and harvesting of its eggs. Aaron’s bent-toed gecko (Cyrtodactylus aaroni) is known only from three localities within the Vogelkop Peninsula (Papua Barat province, Indonesia). Brongersma’s skink (Lipinia venemai) is known only from two localities within the central Vogelkop Peninsula (Papua Barat province, Indonesia). The Manukwari blind snake (Ramphotyphlops similis) is known only from its original collection in 1934 from a small area on the northern coast of the Vogelkop Peninsula (Papua Barat province, Indonesia). The Umar frog (Litoria umarensis), Elke’s frog (L. elkeae) and the Nabire-Mapia frog (L. rara) are all known only from small areas of coastal Papua province, Indonesia. The black-throated cross frog (Oreophryne atrigularis) is known only from two disjunct localities in the region of the Vogelkop Peninsula (Papua Barat and Papua provinces, Indonesia). The golden swamp frog (Papurana aurata) is known only from a single locality in Papua province, Indonesia. The Bomberai rainforest frog (Cophixalus tetzlaffi) is known only from a single locality in the south-western Vogelkop Peninsula. Northern New Guinea Lowland Rainforests and Swamp Forests Rainforests as well as freshwater swamp forests are found across the northern lowlands of New Guinea between the Central Range to the south and the northern coast. They extend from the eastern shore of Cenderawasih Bay in Papua province, Indonesia east to northern Papua New Guinea.

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Several east–west mountain ranges rise from these lowlands, creating isolated pockets of montane rainforest. The Weikor skink (Lipinia albodorsalis) is known only from its original collection in 1932 from north-western Papua New Guinea. The Moaif frog (Litoria mystax) is known only from its original collection in 1906 from north-eastern coastal Papua province, Indonesia. The white-lipped frog (L. albolabris) is known only from its original collection in 1911 from northwestern coastal Papua New Guinea. Chris Dahl’s frog (L. chrisdahli) is known only from a single locality in northern coastal Papua New Guinea. The Simbang big-eyed tree frog (Nyctimystes obsoletus) and the Yuwong tree frog (N. dux) are both known only from a few localities on the Huon Peninsula in north-eastern Papua New Guinea. The Lae fanged frog (Xenorhina subcrocea) is known only from its original collection in 1977 from eastern coastal Papua New Guinea. All of the forest at this locality has since been destroyed, but suitable habitat remains nearby. Wolterstorff’s cross frog (Oreophryne wolterstorffi) is known only from a single specimen collected in 1901 from an imprecise locality in north-eastern Papua New Guinea.

The Mimika River skink (Sphenomorphus mimikanus) and Wollaston’s skink (S. wollastoni) are both known only from their original collection in 1914 from western New Guinea (Papua province, Indonesia). Fred Parker’s worm snake (Gerrhopilus fredparkeri) and McDowell’s worm snake (G. mcdowelli) are both known only from a small area near Port Morseby in south-eastern Papua New Guinea. The four-lined frog (Litoria quadrilineata) is known only from a small area of southern coastal New Guinea (Papua province, Indonesia). The Moroka cross frog (Oreophryne kampeni) and the Port Moresby cross frog (O. loriae) are both known only from historical specimens collected from a small area near Port Morseby in south-eastern Papua New Guinea. The Iarowari school frog (Mantophryne menziesi) is known only from a small area near Port Morseby in southeastern Papua New Guinea. The variegated narrow-mouthed frog (Choerophryne variegata) is known only from a single specimen collected in 1923 from an imprecise locality in south-western New Guinea (Papua province, Indonesia).

Southern New Guinea Lowland Rainforests and Swamp Forests In southern New Guinea, areas of rainforest and freshwater swamp forest lie between the Central Highlands and the southern coast. They extend from Papua province, Indonesia all the way to the eastern tip of Papua New Guinea, being interrupted only by coastal mangroves and a large pocket of savanna. The lowland tree-kangaroo (Dendrolagus spadix) is confined to the lowland rainforests of southern Papua New Guinea, where it is threatened by hunting and habitat destruction. The grey forest wallaby (Dorcopsis luctuosa) remains widespread within the lowland rainforests of southern New Guinea, but is everywhere threatened by hunting and habitat destruction. The giant bandicoot (Peroryctes broadbenti) is a rare type of terrestrial marsupial confined to coastal lowland areas of south-eastern Papua New Guinea. It is threatened by loss of habitat and hunting. The Papuan mastiff bat (Otomops papuensis) is known only from a handful of specimens collected from two disjunct localities in south-eastern Papua New Guinea. Thomas’ big-eared bat (Pharotis imogene) is known only from a small area near Port Morseby in south-eastern Papua New Guinea. It was thought to have been extinct since 1890 until a single specimen was discovered in 2012. The spangled owlet-nightjar (Aegotheles tatei) is known from a handful of records from the lowland rainforests of central-western and south-eastern Papua New Guinea. Scheepmaker’s crowned pigeon (Goura scheepmakeri) is confined to the lowland rainforests of southern Papua New Guinea, where it is threatened by loss of habitat and has been extirpated from many parts of its former range.

The Trans-Fly savanna and grasslands are a lowland ecoregion located on the south-central coast of New Guinea, on both the Indonesian and Papua New Guinea sides. With their monsoon and dry season climate these grasslands and woodlands are quite different from the tropical rainforest that covers much of the island, and resemble the landscape of northern Australia. They are renewed by regular fires at the end of each dry season. The bronze quoll (Dasyurus spartacus) is a type of nocturnal marsupial confined to savanna woodlands in south-central New Guinea, where it is threatened by hunting and, possibly, feral cats. Rand’s brush-tailed rabbit-rat (Conilurus penicillatus randi) is known only from two localities in south-western Papua New Guinea. The Fly River snake-eyed skink (Cryptoblepharus xenikos) is known only from a few specimens collected from two savanna localities in south-western Papua New Guinea.

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Lakes, Rivers, and Marshes New Guinea has several large river systems along with large areas of swamp forest and numerous isolated mountain lakes, all of which are notable for their extraordinary high species diversity. General threats include overfishing, sedimentation due to deforestation and pollution from mining activities. Salvadori’s teal (Salvadorina waigiuensis) occurs in fastflowing streams as well as lakes throughout the Central Highlands of New Guinea, but is everywhere threatened by hunting and loss of habitat. The pig-nosed turtle (Carettochelys insculpta) is a unique relict species inhabiting southern New Guinea as well as northern Australia (Northern Territory). It is exported in large

New Guinea

numbers from southern Papua province, Indonesia for the international live animal trade and locally consumed in Papua New Guinea, while in Australia it is threatened mainly by habitat destruction. Two species of giant softshell turtle (Pelochelys) are threatened by hunting and egg-harvesting. The northern New Guinea giant softshell turtle (P. signifera) remains relatively widespread in the northern lowlands of New Guinea. The striped New Guinea giant softshell turtle (P. bibroni) is confined to lowland rivers and estuaries across southern New Guinea. Pritchard’s snake-necked turtle (Chelodina pritchardi) is confined to a few localities within the Kemp Welch and Laloki river drainages of coastal south-eastern Papua New Guinea. Parker’s snake-necked turtle (C. parkeri) is confined to the swamps and rivers of south-central New Guinea. Both are threatened by hunting and collection for the international pet trade. The white-bellied snapping turtle (Elseya branderhorsti) is a little-known species from the rivers of southern New Guinea (Papua Barat province, Indonesia and Papua New Guinea). Adults are heavily hunted for their meat and the young are collected for use in ‘traditional medicine’. The New Guinea sailfin lizard (Hydrosaurus amboinensis), the world’s largest agamid lizard, is a little-known and ill-defined form that, as currently recognized, is found in New Guinea and on Ambon in the Moluccas. It is threatened by habitat destruction and overcollection for the international pet trade. The striped grunter (Hephaestus obtusifrons) is a perchlike fish confined to two disjunct, high-altitude areas of northcentral New Guinea, the first being the upper Sermowai River and its tributaries in Papua province, Indonesia, and the other the Bewani Mountains of north-western Papua New Guinea. It is threatened by loss of habitat and invasive species. The Popondetta blue-eye (Pseudomugil connieae) is confined to rainforest streams and creeks within a small area of south-eastern Papua New Guinea. The transparent blue-eye (P. pellucidus) is confined to the Aikwa (Iwaka) and Kopi rivers of western New Guinea (Papua province, Indonesia). Both are threatened by loss of habitat and pollution. The Kokoda glass perchlet (Tetracentrum caudovittatus) is known only from a small area of south-eastern Papua New Guinea, where it lives in pools in forest creeks. It is seriously threatened by habitat destruction and degradation. The Arfak rainbowfish (Melanotaenia arfakensis) is found patchily in rivers of the northern coastal area of the Vogelkop Peninsula in north-western New Guinea (Papua Barat province, Indonesia). The mountain rainbowfish (M. monticola) is confined to small, highland tributaries of the Purari and Kikori river drainages in central Papua New Guinea. Ogilby’s rainbowfish (M. ogilbyi) was long known only from a few specimens collected in 1907 from the Lorentz River in western New Guinea (Papua province, Indonesia). It was rediscovered in 1995 from disjunct areas too, but the species remains vulnerable to habitat destruction. The Bulolo rainbowfish (Chilatherina bulolo) is confined to fast-flowing hillstreams within the Markham and Ramu

river drainages of north-eastern Papua New Guinea. First collected in 1934, it was not recorded again until 1978. The spinach pipefish (Microphis spinachioides) is known only from the Sepik, May and Ramu rivers of Papua New Guinea, where it may be threatened by habitat degradation. The Ayamaru Lakes The Ayamaru Lakes (sometimes spelled Ajamaru) are a group of lakes found in a mountainous region of the west-central Vogelkop Peninsula in north-western New Guinea (Papua Barat province, Indonesia). They are notable for the deep holes in their bottoms that lead to underwater caverns. Two species of rainbowfish (Melanotaenia) endemic to the Ayamaru Lakes are threatened by predation by, and competition with, invasive fish species as well as by overcollection for the international aquarium trade. The Ayamaru Lakes rainbowfish (M. ajamaruensis) is known only from specimens collected in 1955, either from the lakes themselves or their tributary streams. It may now survive only within the Kaliwensi River. Boeseman’s rainbowfish (M. boesemani) is confined to Lake Ayamaru and its tributary streams, as well as to Lake Aitinjo. Hoese’s goby (Glossogobius hoesei) is confined to the Ayamaru Lakes and a few small tributary streams. Lake Ayamaru is the largest of the Ayamaru lakes. The Vogelkop blue-eye (Pseudomugil reticulatus) is known only from a handful of specimens collected from tributary streams flowing into Lake Ayamaru, where it is seriously threatened by invasive species. The Paniai Lakes The Paniai Lakes are located in western New Guinea (Papua province, Indonesia). The Paniai gudgeon (Oxyeleotris wisselensis) is a type of goby confined to the Paniai Lakes, where it is absent from Lake Paniai itself but present in lakes Tigi and Tage and their tributaries. Lake Sentani Lake Sentani (Danau Sentani in Indonesian) is a small, shallow, low-altitude lake located in northern coastal New Guinea (Papua province, Indonesia). It forms part of the Cyclops Strict Nature Reserve, although the growing human population along the shore threatens a number of endemic species. The Sentani rainbowfish (Chilatherina sentaniensis) was historically found throughout Lake Sentani and its tributaries, but now seems to be largely confined to a single creek. The red rainbowfish (Glossolepis incisus) is confined to Lake Sentani and its tributaries. The Sentani gudgeon (Oxyeleotris heterodon) is confined to Lake Sentani. Lake Kutubu Lake Kutubu is located in the highlands of central Papua New Guinea, east of the Kikori River into which it eventually drains.

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It is the second largest lake in New Guinea and home to a remarkable number of endemic species. Adamson’s grunter (Hephaestus adamsoni) is confined to Lake Kutubu. The Kutubu hardyhead (Craterocephalus lacustris) is confined to Lake Kutubu and its outlet, the Soro River. The Kutubu rainbowfish (Melanotaenia lacustris) is confined to Lake Kutubu. Five species of goby, the black mogurnda (Mogurnda furva), the variegated mogurnda (M. variegata), the Kutubu mogurnda (M. kutubuensis), the striped mogurnda (M. vitta) and the blotched mogurnda (M. spilota) are all confined to Lake Kutubu and its tributaries, where they are seriously threatened by activities related to oil and gas drilling, overfishing, and invasive species. The Kutubu tandan (Oloplotosus torobo) is a type of eeltail catfish confined to Lake Kutubu. The Mamberamo River The Mamberamo River (Sungai Mamberamo in Indonesian, formerly known as the Idenburg River) is located in northwestern New Guinea (Papua province, Indonesia). The largest in Indonesia by volume of discharge, and also the widest, it is formed by the confluence of two of its tributaries, the Tariku and Taritatu rivers. The Mamberamo water snake (Heurnia ventromaculata) is known only from a single specimen collected from the Mamberamo River in the 1920s. Allen’s river garfish (Zenarchopterus alleni) is known only from a single specimen collected from the Mamberamo River in 1982. The high-finned glassy perchlet (Parambassis altipinnis) is known only from a small number of specimens collected from a single locality within the Mamberamo River in 1920. Habitat in this area has been heavily impacted by logging, and the species may be extinct. The dwarf rainbowfish (Melanotaenia praecox) and Van Heurn’s rainbowfish (M. vanheurni) are both widespread within the Mamberamo River drainage, where they are nevertheless potentially threatened by loss of habitat and invasive species. Both are common in the international aquarium trade. Bleher’s rainbowfish (Chilatherina bleheri) is known from Lake Biru (Lake Holmes) and one other locality within the Mamberamo River drainage. The tiger goby (Pseudogobiopsis tigrellus) is known only from the Mamberamo River. The Fly River The Fly River is located in central-western and south-western Papua New Guinea. It rises in the Victor Emanuel Range (an arm of the Star Mountains) and crosses the south-western lowlands before flowing into the Gulf of Papua in a large delta. It is the largest river in the world without a single dam in its catchment.

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Paska’s blue-eye (Pseudomugil paskai) is known for certain only from the upper Fly River drainage. The Fly River rainbowfish (Melanotaenia sexlineata) is confined to the upper Fly River drainage. Taylor’s catfish (Neoarius taylori) is known only from a few localities within the upper Fly River drainage. The Ok Tedi River (formerly known as the Alice River) is located within the upper Fly River drainage of central-western Papua New Guinea. The open-pit Ok-Tedi Mine at its headwaters has caused considerable habitat degradation and pollution. The glass blue eye (Kiunga ballochi) is known only from a few small creeks along a 15–20 km stretch of the Ok Tedi Mine supply road between Kiunga and Tabubil. Most specimens have been collected from tributaries of the Ok Smak River, itself a tributary of the Ok Tedi River. The species is established in the aquarium trade but has not been seen in the wild for many years. The Ok Tedi rainbowfish (Melanotaenia oktediensis) is confined to the Ok Teki River and its tributaries. The Purari River The Purari River is located in the Central Highlands of central Papua New Guinea. The Pima hardyhead (Craterocephalus pimatuae) is known only from the junction of the Pima and Tua rivers, tributaries of the Purari River. The Pima River is a tributary of the Purari River. The Pima rainbowfish (Melanotaenia pimanensis) is confined to the Pima River. Miscellaneous Lakes, Rivers, and Marshes Lake Kurumoi (Danau Kurumoi in Indonesian) is located on the narrow isthmus at the base of the Vogelkop Peninsula in north-western New Guinea (Papua Barat province, Indonesia). The Kurumoi rainbowfish (Melanotaenia parva) is confined to this one small lake, where it is threatened by competition from introduced tilapia cichlids. Lake Yamur (Danau Yamur in Indonesian) is located on the narrow isthmus at the base of the Vogelkop Peninsula in north-western New Guinea (Papua Barat province, Indonesia). The Yamur grunter (Variichthys jamoerensis) is confined to this single small lake. Lake Wanam is a small lake located in north-eastern Papua New Guinea. The Wanam rainbowfish (Glossolepis wanamensis) has virtually disappeared from this lake due to competition from introduced fishes, although captive populations exist. Lake Tebera is located in south-central Papua New Guinea. The Tebera rainbowfish (Melanotaenia herbertaxelrodi) is confined to the Lake Tebera basin. The Yakati River is located on the narrow isthmus at the base of the Vogelkop Peninsula in north-western New Guinea (Papua Barat province, Indonesia). The Yakati rainbowfish (Melanotaenia angfa) is known only from two tributary streams within the Yakati River drainage.

New Guinea

The Siriwo River is located in western New Guinea (Papua province, Indonesia). Allen’s rainbowfish (Chilatherina alleni) is known only from the Aiborei River, part of the Siriwo drainage. The Sermowai River is located in northern New Guinea (north-western Papua province, Indonesia). The Corona rainbowfish (Melanotaenia corona) is known only from two specimens collected in 1911. The Tami River (Sungai Tami in Indonesian) is located in northern coastal New Guinea (north-eastern Papua province, Indonesia). The Tami rainbowfish (Glossolepis pseudoincisus) is confined to the Tami River. The Ramu River is located in north-eastern Papua New Guinea. The Ramu rainbowfish (Glossolepis ramuensis) is confined to the Ramu River. The Laloki River is located in south-eastern Papua New Guinea, near Port Moresby. The threespot grunter (Hephaestus trimaculatus) is confined to rocky pools within streams in the lower Laloki River drainage.

Coasts and Satellite Islands New Guinea is home to some of the largest areas of brackish water coastal mangroves in the world, which provide important habitat for both terrestrial and marine species. In addition, the island is surrounded by numerous smaller islands and island groups, many of them still relatively pristine. The Bramble Cay mosaic-tailed rat (Melomys rubicola) is known only from Bramble Cay, a tiny vegetated islet located some 50 km south of New Guinea in the Torres Strait. Last seen in 2007 and now considered to be extinct, it is possibly the first known mammalian extinction linked to human-induced climate change. Thomas’ woolly bat (Kerivoula agnella) is known only from Fergusson Island in the D’Entrecasteaux Islands, Tagula and Misima in the Louisiade Archipelago, and Woodlark Island. The mysterious tree monitor (Varanus telenesetes) was originally described based on a single specimen supposedly collected on Rossel Island, in the Louisiade Archipelago. Subsequent searches have failed to find any additional specimens, and native islanders have no knowledge of it. For these reasons it is now believed that the original collection data were in error, and that the species occurs (or occurred) on some other, as yet unknown, island east of New Guinea. The banded water snake (Djokoiskandarus annulata) is known only from a few localities along the south-central coast of New Guinea and its satellite islands. Watson’s coastal stream goby (Stenogobius watsoni) is known only from three islands in Milne Bay, south-eastern New Guinea, where it lives in freshwater creeks. Larson’s stiphodon (Stiphodon larson) is a type of freshwater goby known only from a small area of north-eastern Papua New Guinea and adjacent islands. Robert’s combtooth blenny (Omobranchus robertsi) is known only from brackish mangrove creeks in south-central

coastal Papua New Guinea, but may occur as well on the north-eastern portion of the Cape York Peninsula in Australia. The Schouten Islands The Schouten Islands (also known as the Biak Islands or Geelvink Islands) are located in Cenderawasih Bay, roughly 50 km off the north-western coast of New Guinea. They consist of the main islands of Biak, Supiori, and Numfor along with numerous smaller islands, most of which are still covered by rainforest. The Geelvink Bay flying fox (Pteropus pohlei) is confined to Numfor, Rani, and Yapen. The Schouten naked-backed fruit bat (Dobsonia emersa) is confined to Numfor, Biak, Supiori, and Owi. It is potentially threatened by hunting and disturbance of its roost sites. The Schouten scrubfowl (Megapodius geelvinkianus) is a chicken-like bird confined to Biak, Supiori, Numfor, Manim, Mios Korwar, and Mios Numin. The total population is small and declining due to a variety of threats. The black-winged lory (Eos cyanogenia) is confined to Biak, Supiori, Numfor, Manim, and Mios Num. It is threatened by loss of habitat and overcollection for the cagebird trade. Biak and Supiori are the largest of the Schouten Islands and are near enough to each other to be spanned by a bridge. Both have been heavily deforested, although most of Supiori is now protected by the Pulau Supiori Nature Reserve. Wilson’s spotted cuscus (Spilocuscus wilsoni) is known only from three specimens collected on Biak and Supiori. Boead’s giant naked-tailed rat (Uromys boeadii) is known only from a single specimen collected on Biak in 1963. Beccari’s scops owl (Otus beccarii) is confined to Biak and Supiori. Rosenberg’s lorikeet (Trichoglossus rosenbergii) is confined to Biak and Supiori, where it is uncommon. Brehm’s monarch (Symposiachrus brehmii) is confined to Biak and Supiori. Rosenberg’s hooded pitta (Pitta rosenbergii) is confined to Biak and Supiori. The Biak gerygone (Gerygone hypoxantha) is confined to Biak and Supiori. The Biak leaf warbler (Phylloscopus misoriensis) is confined to Biak. The Biak emerald monitor (Varanus kordensis) is confined to Biak. The Biak frog (Litoria biakensis) is known only from two localities on Biak. Yapen has still-extensive rainforests and a number of endemic species. Two protected areas cover fully one-third of the island. However, the human population is growing and much forest has been cleared. The Jobi frog (Litoria obtusirostris) is known only from its original collection on Yapen in 1875. Wünsches’ wrinkled ground frog (Cornufer wuenscheorum) is known only from Amoman Mountain on Yapen.

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The variable fanged frog (Xenorhina varia) and the Ambaiduru fanged frog (X. lanthanites) are both confined to Yapen. Three species of cross frog (Oreophryne) are endemic to Yapen. The Kontiunae cross frog (O. asplenicola) is known only from Mount Waira, while the Amoman cross frog (O. pseudasplenicola) is known only from Mount Amoman. The Waira cross frog (O. waira) is known only from Mount Waira and one other nearby locality. The Yapen Mehely frog (Copiula exspectata) is known only from a single locality on Yapen. Numfor is an oval-shaped island that served as a major airbase for both sides during World War II. It was formerly covered by pristine rainforest, which has since been heavily logged and degraded. The Numfor leaf warbler (Phylloscopus maforensis) is confined to the island, where it is considered uncommon. The Raja Ampat Islands Located off the north-western tip of the Vogelkop Peninsula, the Raja Ampat Islands (Kepulauan Raja Ampat in Indonesian, literally ‘Four Kings’) consist of over 1500 small islands, cays and shoals surrounding its four main islands. The Raja Ampat wrinkled ground frog (Cornufer batantae) is known only from Batana and Waigeo islands, but may occur on Yapen and Gag islands as well. Waigeo is the largest island in the Raja Ampat group and is dominated by moist lowland and mangrove forest. The naturalist Alfred Russel Wallace spent some time here during the 1850s while on his scientific exploration journey. The Waigeo spotted cuscus (Spilocuscus papuensis) is possibly endemic to Waigeo, although it may be found on Batanta as well. While still fairly common, it is subject to hunting by locals. The Waigeo brush-turkey (Aepypodius bruijnii) was long known only from a small number of specimens collected no later than 1938. In spite of many dedicated expeditions it was not found again until 2002, when a single individual was sighted on Mount Nok near Majalibit Bay. Subsequent searches revealed a number of incubation mounds in a relatively small area. In 2007 another individual was observed on Mount Danai, representing the first time the species had been photographed in the wild. It is thought than less than 1000 individuals still survive in remote hilly areas. The golden-speckled tree monitor (Varanus boehmei) is known only from two museum specimens, both collected from the wildlife trade. Salawati is a large island separated from mainland New Guinea by a narrow strait. The Salawati bent-toed gecko (Cyrtodactylus irianjayaensis) is known only from lowland forests on Salawati. The Salawati blind snake (Ramphotyphlops supranasalis) is known only from two specimens collected during the 1930s. Batanta is located north of Salawati. The blue-speckled tree monitor (Varanus macraei) is known only from Batanta and two small offshore islets. It is

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threatened by loss of habitat and collection for the international pet trade. Kofiau is a relatively small island consisting of raised coral limestone with some volcanic hills, covered in low forest. It has been selectively logged, and as yet has no protected areas. The Kofiau paradise kingfisher (Tanysiptera ellioti) is confined to Kofiau. The Kofiau black-backed monarch (Symposiachrus julianae) is confined to Kofiau. Misool is the southernmost of the Raja Ampat Islands. The yellow tree monitor (Varanus reisingeri) is confined to this one small island, where it is vulnerable to exploitation by the international pet trade. Gag Island is one of the smaller and isolated of the Raja Ampat Islands. Niken’s rat (Rattus nikenii) is seemingly confined to this one small island, where it might potentially be threatened by proposed nickel mining operations. The Trobriand Islands The Trobriand Islands are an archipelago off the eastern coast of New Guinea. They consist of four main islands and a number of smaller coral atolls. The group is an important tropical rainforest region in need of protection. The Trobriand bronzeback tree snake (Dendrelaphis papuensis) is known only from historical specimens with an unclear distribution within the Trobriand Islands. Kiriwina is a small, low-lying, heavily populated island. David’s spiny bandicoot (Echymipera davidi) is known only from Kiriwina. The Woodlark Islands Located east of New Guinea, the Woodlark Islands consist of the eponymous main island and its smaller satellites, among them Mudua, Nusam, Nubara, and the Marshall Bennett group to the south-east. Owing to their relative isolation they are home to several endemic species. A plan in the 1990s to log most of the forest on Woodlark Island and replace it with palm oil plantations was fortunately scrapped thanks to opposition by local residents. The Woodlark cuscus (Phalanger lullulae) is known from Woodlark and Alcester islands (having possibly been introduced on the latter) and may still be present on Madua. The Woodlark bent-toed gecko (Cyrtodactylus murua) is confined to Woodlark Island. The Woodlark worm-eating snake (Toxicocalamus longissimus) is known only from a few specimens collected on Woodlark Island. The D’Entrecasteaux Islands The volcanic D’Entrecasteaux Islands consist of the three main islands of Goodenough, Fergusson and Normanby, along with numerous smaller islets and reefs, situated near the eastern tip of New Guinea.

The Bismarck Archipelago

The D’Entrecasteaux tree mouse (Pogonomys fergussoniensis) is confined to Fergusson, Goodenough and Normanby islands, where it is threatened by deforestation. Goldie’s bird-of-paradise (Paradisaea decora) is confined to Ferguson and Normanby islands. The D’Entrecasteaux keelback (Tropidonophis dolasii) is a type of snake confined to Goodenough and Fergusson islands. The D’Entrecasteaux cross frog (Oreophryne insulana) is confined to Goodenough and Fergusson islands. Goodenough Island is known locally as Nidula. The black forest wallaby (Dorcopsis atrata) is confined to montane forest on Goodenough Island, where it is threatened by habitat destruction and hunting. The loud big-eyed tree frog (Nyctimystes avocalis) is known only from a few localities on Goodenough Island. The Goodenough Mehely frog (Copiula minor) is known only from Goodenough Island. Fergusson Island is known locally as Maratau. Tate’s striped possum (Dactylopsila tatei) is confined to the mountains of western Fergusson Island. The black-naped pheasant pigeon (Otidiphaps insularis) is a rare species confined to Fergusson Island. The Louisiade Archipelago The Louisiade Archipelago is located 200 km south-east of New Guinea. It consists of a string of 10 larger volcanic islands, most fringed by coral reefs, along with 90 smaller coral islands. The Louisiade butcherbird (Cracticus louisiadensis) is confined to Tagula and the smaller islands of Junet (Panatinani), Panawina, and Sabara. The white-chinned myzomela (Myzomela albigula) is a type of honeyeater known only from the smaller islands of the Louisiade Archipelago. The Louisiade land frog (Austrochaperina yelaensis) is known only from Rossel and Tagula. Tagula (also known as Vanatinai or Sudest) is the largest island in the archipelago, with a forested mountain range running through the centre. Over half of its forests have already been logged. The Tagula white-eye (Zosterops meeki) is a type of passerine bird confined to higher-elevation forest on the island. The Tagula honeyeater (Meliphaga vicina) is confined to Tagula. Hill’s frog (Litoria hilli) is confined to Tagula. The Tagula rainforest frog (Cophixalus tagulensis) is known only from three specimens and has not been reported since 1956. Rossel Island (also known as Yela) is the easternmost island in the Louisiade Archipelago. It is relatively large and mountainous with extensive forests, mangroves, and a fringing coral reef. The Rossel mosaic-tailed rat (Melomys arcium) is known only from a small number of specimens collected in 1956 and 1960.

The Rossel cicadabird (Edolisoma rostratum) is known only from specimens collected in 1898, along with a reported sighting of one individual in 2014. Meek’s pitta (Erythropitta meeki) is a type of passerine bird scientifically known only from a single specimen collected in 1898. It is familiar to local people, however, who report that it is restricted to higher elevations. The Rossel worm snake (Gerrhopilus hades) is known only from Rossel Island. The Rossel school frog (Mantophryne louisiadensis) is confined to Rossel Island. Misima (also known as St Aignan) is mountainous and densely forested. There has been extensive mining activity on the island. Richards’ snake-eyed skink (Cryptoblepharus richardsi) is known only from Misima. The Misima worm-eating snake (Toxicocalamus misimae) is known only from three specimens collected in the 1960s. The Misima rainforest frog (Cophixalus misimae) is confined to Misima. The Paidaido Islands The Paidaido Islands are located off the coast of north-western New Guinea (Papua province, Indonesia). Emma’s giant naked-tailed rat (Uromys emmae) is known only from a single specimen collection from Owi Island in 1946. It is most likely extinct.

The Bismarck Archipelago The Bismarck Archipelago is a group of mainly volcanic islands off the north-eastern coast of New Guinea. It includes the large islands of New Britain and New Ireland along with a number of smaller islands and island groups. Many of these rainforest-covered atolls are still uninhabited. The New Britain water rat (Hydromys neobritannicus) is known only from three specimens collected from New Britain during the 1930s; there are also unconfirmed reports from Umboi. Gilliard’s flying fox (Pteropus gilliardorum) is known only from two specimens collected on New Britain and a third from New Ireland. The Bismarck woolly bat (Kerivoula myrella) occurs on New Britain, Duke of York, Umboi and Manus islands, where it is believed to be uncommon. The slaty-backed goshawk (Accipiter luteoschistaceus) is known only from a handful of specimens collected from New Britain and Umboi. The Bismarck sparrowhawk (A. brachyurus) is found in the montane forests of New Ireland and (at least historically) New Britain. The Bismarck bronzewing (Henicophaps foersteri) is a rare type of dove confined to New Britain and Umboi. The Bismarck kingfisher (Ceyx websteri) is confined to the banks of small, slow-moving rivers on New Britain, New Ireland, Umboi, New Hanover, and Lihir.

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The Bismarck spiny skink (Tribolonotus annectens) is confined to montane forests on New Britain and Umboi.

New Britain New Britain is the largest island in the Bismarck Archipelago. Largely formed by volcanic processes, there are several active volcanoes on the island and steep cliffs form some sections of the coastline; in others the mountains are further inland, while the coastal areas are flat and bordered by coral reefs. Most of the terrain was historically covered with tropical rainforest, although the latter have been largely destroyed in recent years, largely to clear land for oil palm plantations. Several large as well as many small rivers are fed by New Britain’s high rainfall. The black honey buzzard (Henicopernis infuscatus) is a large raptor confined to New Britain and the offshore island of Lolobau. The golden masked owl (Tyto aurantia) was long known only from a few specimens and field sightings, although in recent years the species has been recorded fairly regularly, even in and around oil palm plantations, suggesting that it has some tolerance of degraded habitats. The New Britain hawk-owl (Ninox odiosa) is confined to New Britain, where it remains relatively common and widespread. The New Britain goshawk (Accipiter princeps) is known only from four historical specimens and a few recent records. The blue-eyed cockatoo (Cacatua ophthalmica) is confined to New Britain, where it remains relatively common and shows some tolerance to degraded habitat. The New Britain thicketbird (Megalurulus grosvenori) is a type of warbler known only from two specimens collected in the Whiteman Range of central New Britain in 1959. Gilliard’s wrinkled ground frog (Cornufer gilliardi) and Zweifel’s wrinkled ground frog (C. nexipus) are both confined to a few localities on New Britain, where they are threatened by loss of habitat. The New Britain land frog (Austrochaperina novaebritanniae) is known only from a few localities in north-eastern New Britain. The New Britain coastal stream goby (Stenogobius alleni) is known only from 10 specimens collected from a small freshwater stream in north-eastern New Britain.

New Ireland New Ireland lies north-east of New Britain. Originally covered by dense lowland and montane rainforest, today much has been lost or degraded, and what remains has been littlestudied. The demonic tube-nosed fruit bat (Nyctimene masalai) appears to be confined to New Ireland. The New Ireland mangrove monitor (Varanus douarrha) was long known only from a single specimen collected in 1823. It was reportedly rediscovered in 2017. The Tabar Islands The Tabar Islands are located north-east of New Ireland.

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The Tabar pitta (Erythropitta splendida) is a type of passerine bird confined to the Tabar Islands, where it is threatened by forest loss. Dyaul Island Dyaul Island is located off the southern coast of north-western New Ireland. The Dyaul monarch (Symposiachrus ateralbus) is a type of passerine bird confined to Dyaul. The Dyaul flycatcher (Myiagra cervinicolor) is confined to Dyaul.

The Admiralty Islands The Admiralty Islands are a group of 18 islands within the Bismarck Archipelago. The Admiralty monarch (Symposiachrus infelix) is a type of passerine bird divided into two subspecies endemic to the Admiralty Islands. Coultas’ monarch (S. i. coultasi) is confined to the islands of Rambutyo and Tong. The Admiralty scaly-toed gecko (Lepidodactylus pulcher) is known only from a few islands and has rarely been collected. Surveys since 1990 have failed to locate it. Manus Manus is the largest of the Admiralty Islands and, unlike the other islands in its group, still retains much of its original rainforest cover. The Manus mosaic-tailed rat (Melomys matambuai) is a type of arboreal rodent known only from two specimens. The Manus masked owl (Tyto novaehollandiae manusi) is known from two specimens. Recent surveys have failed to locate it. The superb pitta (Pitta superba) is a type of passerine bird endemic to Manus, where it is known to be rare and declining. The Manus fantail (Rhipidura semirubra) was historically common on the island that bears its name, but there have been no records there since 1934. It is now known only from a few small neighbouring islands. The Manus monarch (Symposiachrus infelix infelix) is confined to Manus. The Manus bumblebee gecko (Nactus kunan) is known only from two specimens collected in 2012, but may occur more widely on the island. The Admiralty five-striped emo skink (Emoia mivarti) is confined to Manus and perhaps a few nearby islands. The Admiralty spiny skink (Tribolonotus brongersmai) is known only from its type locality in the mountains of Manus.

The St Matthias Islands The St Matthias Islands are a small group of around 10 islands located at the northern end of the Bismarck Archipelago. Mussau Mussau is the largest of the St Matthias Islands.

The Solomon Islands

The Mussau fantail (Rhipidura matthiae) is a type of passerine bird confined to Mussau, where the population is believed to be small. The Mussau triller (Lalage conjuncta) is a type of passerine bird known only from a single specimen and a few field observations from the hill forests of Mussau. The Mussau flycatcher (Myiagra hebetior) is largely confined to old-growth forests on Mussau, where it is threatened by logging.

The Solomon Islands The Solomons consist of six major islands and over 900 smaller ones lying to the east of Papua New Guinea and north-west of Vanuatu. They have distinct affinities with nearby New Guinea in their fauna and in the dense tropical forests covering their mountain slopes. Several of the islands have been seriously impacted by human activities and introduced species. Poncelet’s naked-tailed rat (Solomys ponceleti) and the rugged naked-tailed rat (S. salebrosus) are known from Bougainville and Choiseul, with additional fossil remains recorded from Buka. Both are threatened by habitat destruction and hunting. Andersen’s monkey-faced fruit bat (Pteralopex anceps) is known only from Bougainville and Choiseul. Thomas’ monkey-faced fruit bat (P. atrata) is now most likely confined to Guadalcanal, although a single skull has also been recorded from New Georgia (where it is now almost certainly extirpated). Flannery’s monkey-faced fruit bat (P. flanneryi) is known from Bougainville (including the satellite islet of Puruata), Choiseul, Buka and Santa Isabel (including the satellite islet of Barora Fa). All are seriously threatened by habitat destruction and hunting. The flower-faced bat (Anthops ornatus) is a rare species known only from a handful of specimens collected from Bougainville, Choiseul, Ngella, Santa Isabel, and Guadalcanal. The fierce leaf-nosed bat (Hipposideros dinops) is found on Bougainville, Guadalcanal, Malaita, New Georgia, San Jorge and Santa Isabel, but is everywhere uncommon. Sanford’s sea eagle (Haliaeetus sanfordi) is found throughout the Solomon Islands but is everywhere uncommon, particularly on the larger islands, and appears to be declining. The fearful owl (Nesasio solomonensis) is a rare species confined to old-growth forests on Bougainville, Choiseul, Santa Isabel and possibly Buka. The imitator goshawk (Accipiter imitator) is known only from a handful of specimens and infrequent sightings on Bougainville, Choiseul and Santa Isabel. The chestnut-bellied imperial pigeon (Ducula brenchleyi) is known from Guadalcanal, Malaita and Makira in the southern Solomons (including, at least historically, the satellite islands of Ulawa, Ugi, and Three Sisters, although it may now be extirpated from there). An expedition in 1953 failed to record the species on Guadalcanal and there are few recent records, while on Malaita it is known only from a single

specimen along with a further two sightings in 1990. On Makira it is reported to be somewhat more common, at least in some areas. Hunting and habitat destruction are the major threats. The Solomons nightjar (Eurostopodus nigripennis) is a rare species that has historically been reported in coastal areas throughout the northern and central Solomon Islands, but now appears to be largely confined to Tetepare, nearby Hele Bar, and Nabonibao. The white-eyed starling (Aplonis brunneicapillus) is a rare species found patchily on Bougainville, Choiseul, Rendova, and Guadalcanal. It may possibly occur on other islands as well. The black-faced pitta (Pitta anerythra) is a type of passerine bird confined to Bougainville, Choiseul and Santa Isabel, where it was at one time fairly common. The species is now notably rare due to logging of its lowland forest habitat. The giant spiny skink (Tribolonotus ponceleti) is confined to lowland areas of Shortland, Santa Isabel, and Gatokae islands, where it is said to be very rare. There is some doubt as to whether the species also occurs on Bougainville. The Solomons black-banded krait (Loveridgelaps elapoides) is a rare type of snake from the western and central Solomon Islands. The small-headed blind snake (Ramphotyphlops mansuetus) is known only from two specimens collected from Bougainville and Makira. The arboreal blind snake (R. angusticeps) is known only from a few specimens collected from Choiseul, Guadalcanal, Malaita, Makira, and Malaupaina in the Olu Malau Islands. The Solomons palm frog (Palmatorappia solomonis) is known from several large islands in the Solomons, including Bougainville and Buka, but is everywhere threatened by habitat destruction.

Bougainville Bougainville is the largest island in the Solomon Islands archipelago. Along with the nearby island of Buka, it is a single landmass separated by a deep 300-m wide strait. There are several active, dormant or inactive volcanoes that rise to 2400 m. Introduced cats and black rats are common throughout the island. The Bougainville moustached kingfisher (Actenoides bougainvillei) is known only from a few specimens taken prior to 1938. There were no more reports until the 1980s, when calls of this species were heard on the edge of a lowland swamp forest and a pair was observed at a nest-hole. More recently, a freshly killed bird was shown to birdwatchers by local people. The Bougainville thicketbird (Megalurulus llaneae) is only definitely known from the Crown Prince Range, although it may occur more widely within the island’s montane forests. The Bougainville bush-warbler (Cettia haddeni) is a poorly known species confined to montane areas. The Kunua skink (Sphenomorphus transversus) is known only from a single specimen collected in 1971. Taylor’s skink

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(S. taylori) and the fragmented skink (S. fragosus) are both confined to Bougainville. The Bougainville coral snake (Parapistocalamus hedigeri) is a very rare species from eastern and southern Bougainville. The Kunua blind snake (Acutotyphlops kunuaensis) is a little-known fossorial species confined to lowland forests. The Lake Loloru wrinkled ground frog (Cornufer gigas) is known only from its original collection in 1970 from a single locality in southern Bougainville. The Aresi wrinkled ground frog (C. macrops) is known only from a small area of northern Bougainville. Parker’s wrinkled ground frog (C. parkeri) is divided into two subspecies. The nominate form (C. p. parkeri) is confined to northernmost Bougainville, while the Buka wrinkled ground frog (C. p. bukanensis) is confined to the satellite island of Buka.

Guadalcanal Guadalcanal (known locally as Isatabu) is mountainous and still largely covered in dense tropical rainforest. The porcine giant naked-tailed rat (Uromys porculus) and the emperor giant naked-tailed rat (U. imperator) are both known only from specimens collected in the 1880s. The king rat (U. rex) is similarly known only from a few specimens. The Makarakomburu monkey-faced fruit bat (Pteralopex pulchra) is known from a single specimen collected on the southern slopes of Mount Makarakomburu. The Guadalcanal thrush (Zoothera turipavae) is known only from a few specimens. The Guadalcanal moustached kingfisher (Actenoides excelsus) is confined to the mountains of Guadalcanal. The Lunga River scaly-toed gecko (Lepidodactylus shebae) is known only from a single specimen collected in 1944. The yellow-eyed scaly-toed gecko (L. flaviocularis) is known only from two specimens collected in 1978 and 1990 near the summit of Mount Austen. Beck’s blind snake (Ramphotyphlops becki) is confined to Guadalcanal. The Malukuna wrinkled ground frog (Cornufer malukuna) is known only from its original description in 1968.

Malaita Malaita is a thin, mountainous island that has seen much of its once-pristine forests converted into subsistence gardens. The Malaita hawk-owl (Ninox malaitae) is known only from a few specimens. The Malaita fantail (Rhipidura malaitae) is a type of passerine bird endemic to mountainous areas, where it is rarely seen.

Santa Isabel Santa Isabel is the longest of the Solomon Islands and the third largest in terms of surface area. The Isabel naked-tailed rat (Solomys sapientis) is now confined to Santa Isabel Island, although it may have once occurred on Malaita and possibly Makira as well.

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Makira Makira (also known as San Cristóbal Island) is located east of Guadalcanal and south of Malaita. Introduced rats, pigs, cats, and dogs are widespread, and most of the lowland forest has been logged. Surrounding islets have fared somewhat better. The Ugi naked-tailed rat (Solomys salamonis) is known only from a single specimen collected on the islet of Ugi. It may still exist on Makira itself. The Makira flying fox (Pteropus cognatus) is known only from a few specimens collected from Makira and the islet of Uki Ni Masi. A report during the early 1990s from Olu Malau (Three Sisters) Island likely represents vagrants. It is known to be hunted by locals for food. The Makira leaf-nosed bat (Hipposideros demissus) is confined to Makira, where it still appears to be fairly common. The Makira hawk-owl (Ninox roseoaxillaris) is known from a few records on Makira and possibly the islets of Ugi and Santa Catalina as well. The Makira moorhen (Pareudiastes silvestris) is a flightless and almost tailless species that was long known only from a single specimen collected within the central mountains in 1929. It was not observed again until 1953, and was last seen by hunters in 1974. A survey during the 1990s failed to find any trace of it, although more recently there have been reports of birds having been caught by dogs. The thick-billed ground dove (Pampusana salamonis) is known only from two specimens, the first taken from Makira in 1882 and the second from the islet of Ramos in 1927. Searches since then have failed to find any others and the species is now believed to be extinct, a victim of introduced predators.

Choiseul Choiseul is a large island located south-east of Bougainville. The Choiseul crested pigeon (Microgoura meeki) was a unique representative of its genus believed to have been endemic to the island. Introduced dogs and cats were likely responsible for the extermination of this species, which was last recorded in 1904.

The New Georgia Islands The New Georgia Islands are located north-west of Guadalcanal. They are comprised of the main island of New Georgia along with numerous satellite islands. The New Georgia monkey-faced fruit bat (Pteralopex taki) is confined to New Georgia and the islet of Vangunu. It is thought to have been extirpated from the island of Kolombangara during the mid-1970s. Kolombangara The island of Kolombangara is an almost perfectly round, dormant stratovolcano that reaches a height of 1770 m. The Kolombangara leaf warbler (Phylloscopus amoenus) is confined to open slopes on the inner caldera of Kolombangara, where the total population is thought to be between 1000 and 2000.

Vanuatu (New Hebrides)

and extensive surveys in the 1980s and 1990s failed to find any trace. The Nendö shrikebill (Clytorhynchus sanctaecrucis) is a type of songbird known from two specimens collected in 1927, two pairs seen in 2004, and two more pairs reported in 2014. Other surveys have failed to find the species. Vanikoro Although usually referred to as a single island, Vanikoro is technically a cluster of small islands surrounded by a coral reef. Much of the original forest cover was logged in the midtwentieth century. The Vanikoro flying fox (Pteropus tuberculatus) is confined to Vanikoro, where it is known only from a few specimens all of which were collected before 1930. Subsequent surveys have failed to find it. The Vanikoro monarch (Mayrornis schistaceus) is a type of passerine bird confined to Vanikoro and its small satellite, Buma.

Rennell Island

Figure 6.2 Choiseul crested pigeon, 1904. (Credit: John Gerrard Keulemans.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

Ghizo Ghizo is a tiny islet whose remaining forest is very fragmented and covers less than 1 km2 in total. The Ghizo white-eye (Zosterops luteirostris) is a type of passerine bird confined to Ghizo. Ranongga Ranongga is a small island located west of Ghizo. The Ranongga white-eye (Zosterops splendidus) is confined to Ranongga, where it is threatened by habitat destruction.

The Santa Cruz Islands The Santa Cruz Islands are a particularly isolated group lying approximately 400 km to the south-east of the main Solomon Islands chain, and north of Vanuatu. The Tomotu Neo flying fox (Pteropus nitendiensis) is confined to the islands of Nendö and Tömotu Neo in the Santa Cruz Islands. Nendö (Santa Cruz Island) Nendö is the largest of the Santa Cruz Islands. The Nendö tube-nosed fruit bat (Nyctimene sanctacrucis) is known only from a single specimen collected in the late nineteenth century. It was last reported on the island in 1907,

Isolated some 180 km south of the Solomons, Rennell Island (known locally as Mugaba) is the second largest upraised coral atoll in the world. It appears to have been left more or less undamaged by European explorers and traders, and remains largely unmodified. Apparently, the Spaniards did not bring in any domestic animals, and even rats are said to be lacking. Therefore, this island still possesses a rich bird life, with more than half of the species found nowhere else. The Rennell flying fox (Pteropus rennelli) is only known from a few specimens. It is presumably vulnerable to stochastic events such as cyclones. Lake Te-Nggano Lake Te-Nggano is the largest lake in the South Pacific. The Rennell sea krait (Laticauda crockeri) is a type of venomous snake confined to Lake Te-Nggano.

Ontong Java Atoll Ontong Java Atoll is remote and consists of some 50 tiny vegetated islands that have largely been converted into coconut plantations, and seems likely to be submerged by rising sea levels in the future. The Ontong Java flying fox (Pteropus howensis) is confined to Ontong Java, where it has not been collected since 1945.

Vanuatu (New Hebrides) The Vanuatu Islands (formerly known as the New Hebrides) are located about 1750 km east of northern Australia. They comprise some 40 volcanic islands, many of which are still covered by dense forests. The Vanuatu flying fox (Pteropus anetianus) is still widespread in the lowland forests of Vanuatu, but has been extirpated from many areas due to hunting.

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The white-necked petrel (Pterodroma cervicalis) is divided into two subspecies. The Vanuatu white-necked petrel (P. c. magnificens) is only known to breed on Vanua Lava and Tanna in Vanuatu. The Vanuatu scrubfowl (Megapodius layardi) is a chickenlike bird endemic to Vanuatu, where it is generally uncommon and threatened by hunting, egg collection and feral dogs. The Vanuatu imperial pigeon (Ducula bakeri) is confined to a few of the larger northern islands in Vanuatu, where it is threatened by habitat destruction and hunting. The Erronan emo skink (Emoia erronan) is confined to the islands of Futuna and Aniwa. The Vanuatu stiphodon (Stiphodon astilbos) is a type of freshwater goby known only from coastal streams on Santo, Pentecost and Efate islands.

Espiritu Santo Espiritu Santo is the largest of the Vanuatu Islands. The Espiritu Santo mountain starling (Aplonis santovestris) is confined to the cloud forests of Espiritu Santo’s highest mountains. Bulel’s scaly-toed gecko (Lepidodactylus buleli) is known only from the western coast of Espiritu Santo, but may range more widely. Kalfatak’s stiphodon (Stiphodon kalfatak) is a type of freshwater goby known only from eight specimens collected from the Namatia River. The Espiritu Santo goby (Schismatogobius vanuatuensis) is confined to streams on Espiritu Santo.

Tanna Tanna is located in southern Vanuatu. The Tanna ground dove (Pampusana ferrugineus) was recorded on Tanna Island in 1774, but has never been seen again and was probably exterminated by the natives.

Figure 6.3 Tanna ground dove painted by Johann Georg Adam Forster in 1774. (Credit: Johann Georg Adam Forster.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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Aneityum Aneityum (also known as Anatom) is the southernmost island in Vanuatu. The Aneityum emo skink (Emoia aneityumensis) is confined to this one small island, where it appears to be relatively common.

Erromango Erromango is located in southern Vanuatu. The Erromango saw-tailed gecko (Perochirus guentheri) is known only from four specimens collected from Erromango.

The Banks Islands The Banks Islands are located in northern Vanuatu. They are comprised of Gaua, Vanua Lava, Ureparapara, and a number of smaller islands. The Banks flying fox (Pteropus fundatus) is confined to lowland areas of Vanua Lava and Mota, where it is threatened by hunting.

The Fiji Islands The Fiji Islands are located in southern Melanesia, and consist of two major islands surrounded by an archipelago of hundreds of smaller ones as well as islets, volcanic rocks, and atolls. Many of the larger islands are mountainous and covered by forests. The Fijis, as elsewhere, suffered from the early European practice of introducing animals indiscriminately. Both the mongoose – released to control the black rats brought to the Fijis by the Polynesians – and the sambar deer (Rusa unicolor) have been most harmful, the former killing groundnesting birds, the latter destroying vegetation. At least 15 species of birds have also been introduced, which compete with local species. The bar-winged rail (Hypotaenidia poeciloptera) was an almost flightless species that was long known only from a dozen specimens collected on Viti Levu and Ovalau during the nineteenth century. It was reported from Taveuni in 1971 and from Viti Levu in 1973, but there have been no more reports since then and the species is now thought to be extinct, a victim of introduced cats and mongooses. The red-throated lorikeet (Charmosyna amabilis) historically occurred on Viti Levu, Vanua Levu, Taveuni and Ovalau. Always regarded as rare, the species was decimated by introduced black rats and loss of habitat. There were unconfirmed reports from the 1980s and 1990s from Vanua Levu, Ovalau, and Taveuni, but no actual specimens, photos or observations exist from the latter two islands since 1965. The species now appears to be confined to the mountains of Viti Levu, and perhaps Vanua Levu. The long-legged thicketbird (Megalurulus rufus) is a large, thinly built warbler with a long tail and legs. Four specimens were collected on Viti Levu between 1890 and 1894, after which the species was not reported again (apart from a few unconfirmed sightings) until 1974, when a fifth specimen was

The Fiji Islands Figure 6.4 Illustration of a bar-winged rail from 1867. (Credit: Unknown.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

found on Vanua Levu. In 2003 a small population was discovered in the Wabu Forest Reserve on Viti Levu, and subsequently others in a few areas of montane forest. The total population is thought be between 70 and 400. The Fijian crested iguana (Brachylophus vitiensis) was historically widespread on Viti Levu and Vanua Levu, but was extirpated there due to mongoose predation and the conversion of most of their dry forest habitat into sugarcane plantations and other development. By the latter part of the twentieth century it was confined to a few smaller satellite islands in the north-western Fiji archipelago (Yadua Taba, Yadua, Macuata, Yaquaga, Devuilau, Waya, Malolo Levu, Qalito, Monu, and Monuriki). Since then, with the exception of those on Yadua Taba and possibly Macuata, all other populations have become barely detectable and perhaps functionally extinct due to habitat degradation by goats, predation by feral cats, and fires. The Fijian banded iguana (B. bulabula) is historically known from the wetter islands of Vita Levu (where it is now present only in a few remote parts of the highlands), Vanua Levu (now extirpated), Ovalau, Kadavu, Mali, and Cikobia, and may have once inhabited several others. It too is seriously threatened by mongoose, black rat and feral cat predation, goat herding, and loss of habitat. The Fijian scaly-toed gecko (Lepidodactylus manni) is confined to highland areas of Viti Levu, Ovalau, and the Kadavu Islands. The Viti copper-headed emo skink (Emoia parkeri) is known only from six islands within the Fiji Islands, where it is threatened by loss of habitat and mongoose predation.

The Fijian wrinkled ground frog (Cornufer vitianus) historically occurred widely on Fiji but is now restricted to the mongoose-free islands of Ovalau, Gau, Taveuni, and Viwa. Lever’s goby (Redigobius leveri) is known only from a few shallow freshwater creeks and rivers on Viti Levu, Vanua Levu and Taveuni.

Viti Levu Viti Levu is the largest and most populous of the Fijian Islands. Its rather rugged terrain is divided into roughly equal halves by a north–south mountain range. The centre of the island is forested and features Mount Tomanivi, Fiji’s highest peak. The pink-billed parrotfinch (Erythrura kleinschmidti) is confined to Viti Levu, where it is rare and patchily distributed. The Viti Levu mountain emo skink (Emoia campbelli) is known only from a single locality in the mountains of Viti Levu. It has not been recorded since the 1980s. The Viti Levu snake (Ogmodon vitianus) is confined to the south-eastern part of the island. The Viti Levu dartfish (Parioglossus triquetrus) is known from a small area of mangrove swamps and creeks in southern Viti Levu.

Vanua Levu Vanua Levu (formerly known as Sandalwood Island) is the second largest island in Fiji, and is located some 65 km to the north of Viti Levu. The Natewa silktail (Lamprolia klinesmithi) is a type of passerine bird confined to the Natewa Peninsula in eastern Vanua Levu, which continues to be extensively logged.

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The Vanua Levu mountain emo skink (Emoia mokosariniveikau) is known only from a few localities. The Lekutu goby (Redigobius lekutu) is a freshwater fish confined to two river systems in the north of the island.

The Kadavu Islands The Kadavu Islands are an archipelago located south of Viti Levu. Dominated by Kadavu, the fourth largest island in Fiji, it also includes Ono, Drauni, Galoa, and a number of islets in the Great Astrolabe Reef. The crimson shining-parrot (Prosopeia splendens) is confined to the islands of Kadavu and Ono. Reports of breeding on other islands are unconfirmed, but are likely to originate from escaped cage birds.

Taveuni Taveuni is a massive, cigar-shaped shield volcano located east of Vanua Levu. The Fijian monkey-faced fruit bat (Mirimiri acrodonta) is only positively known from the summit of Des Voeux Peak on this island, although it may also be present at high elevations on Vanua Levu. The Taveuni blind snake (Ramphotyphlops aluensis) is known only from a single locality on Taveuni.

Rotuma Rotuma is a shield volcano and considered an Important Bird Area as defined by BirdLife International. The Rotuma myzomela (Myzomela chermesina) is a type of honeyeater confined to Rotuma and a few offshore islets. The Rotuma scaly-toed gecko (Lepidodactylus gardineri) is highly specialized within its already restricted distribution.

Gau Island Gau is a large island whose forested interior is considered an Important Bird Area as defined by BirdLife International. The Fijian petrel (Pseudobulweria macgillivrayi) was long known only from a single fledgling male collected on Gau Island in 1855. In recent years, however, there have been a number of reports of grounded birds on the island, while at sea the only unequivocal sighting was in 2009. The total population is unknown, but may number less than 100. The Gau banded iguana (Brachylophus gau) is entirely confined to Gau, where it mainly inhabits upland forests as well as coastal forest patches. It is threatened by rat and feral cat predation, degradation of habitat by goats, and fires.

The Lau Islands The Lau Islands are a chain of about 60 islands and islets located in eastern Fiji. The Lau banded iguana (Brachylophus fasciatus) is native to the Lau Islands, where it is was at one time widespread but is now confined to around 11 islands. An additional population in the Tonga Islands was introduced by humans during historic times, most likely for use as a food source. It is threatened

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by loss of habitat and introduced species, in particular rats and feral cats. Oni-i-Lau Ono-i-Lau is actually a group of six islands within a common barrier reef system in the Lau Islands. It consists of four central volcanic islands and three clusters of limestone islets. The Ono-i-Lau ground skink (Leiolopisma alazon) was probably widespread on Ono-i-Lau prior to the introduction of cats and pigs, but is now restricted to three small islets. It has only been recorded once since its original description in the 1980s.

New Caledonia New Caledonia (Nouvelle-Calédonie in French) is an archipelago located in the south-western Pacific, 1210 km east of Australia. It includes the main island of Grande Terre as well as the smaller Loyalty Islands, Chesterfield Islands, the Bélep Archipelago, the Isle of Pines, and a few remote islets. All are fragments of the ancient supercontinent Gondwana, separated from other landmasses for tens of millions of years, and feature a variety of habitats from coastal scrub to mountain ranges. Eucalypt forests grow in the drier parts, while more humid areas have produced dense tropical rainforests. Introduced cats, dogs, pigs, rats and other species have had a devastating effect, particularly upon birds. The ornate flying fox (Pteropus ornatus) is found on the main island of Grande Terre as well as on Lifou and Maré in the Loyalty Islands, where it is threatened by habitat destruction and local hunting. Gould’s petrel (Pterodroma leucoptera) is a type of small seabird divided into two subspecies, both of which are confined to a few breeding colonies but which range widely across the Indo-Pacific at other times. The New Caledonian Gould’s petrel (P. l. caledonica) breeds on steep forested valleys of the central mountain chain, with an additional small colony on Raivavae in French Polynesia. It is threatened mainly by introduced pigs, which dig up nesting burrows, and black rats which prey upon the eggs and nestlings. Adults are sometimes killed at night when they fly into lights in Nouméa. The rough-snouted giant gecko (Rhacodactylus trachyrhynchus) is confined to few widely scattered populations on Grande Terre and to the Isle of Pines and surrounding islets. The New Caledonian mossy gecko (Mniarogekko chahoua) is known from a few widely scattered populations on Grande Terre, the Isle of Pines, and Ile Art in the Bélep Islands. The eyelash crested gecko (Correlophus ciliatus) is confined to two disjunct forest fragments on Grande Terre (Provence Sud) and on the Isle of Pines and surrounding islets. Long thought extinct until its rediscovery in 1994, it is threatened mainly by introduced fire ants. Aubrey’s whiptailed skink (Tropidoscincus aubrianus) is found patchily on Grande Terre and on the Isle of Pines.

New Caledonia

The Néhoué River forest skink (Kanakysaurus viviparus) is known only from a few localities in northern Grande Terre (Provence Nord) and from Ile Art and Ile Pott in the Bélep Islands, where it is threatened by loss of habitat and invasive species.

Grande Terre Grande Terre is the largest and principal island of New Caledonia and one of the largest islands in the Pacific Ocean. Nearly 400 km in length and 50–70 km wide in most places, it features a range of habitats from tropical moist and dry forests to maquis shrubland and dry grassland. The New Caledonian flying fox (Pteropus vetulus) is confined to Grande Terre, where it is threatened by hunting and disruption of its roost sites. The New Caledonian long-tailed fruit bat (Notopteris neocaledonica) is known for certain only from two cave-roosting colonies in northern Grande Terre (Provence Nord). The New Caledonian wattled bat (Chalinolobus neocaledonicus) is presently known only from three roof-nesting colonies on Grande Terre. The fairy tern (Sternula nereis) is divided into three colony-nesting subspecies. The New Caledonian fairy tern (S. n. exsul) breeds only on Grande Terre and a few offshore islands where it was historically abundant, but by the twentyfirst century had been reduced to around 100–200 pairs due to human disturbance and other factors. The kagu (Rhynochetos jubatus) is a beautiful, heron-like bird formerly widespread on Grande Terre. Predation by humans and hunting dogs as well as habitat destruction have long confined it to a few areas of dense montane forest, where the total population is thought to be under 2000. Most are found within the Parc des Grandes Fougères and Parc Provincial Rivière Bleue (Provence Sud). The New Caledonian wood rail (Gallirallus lafresnayanus) is a large, cryptic species known only from 17 specimens collected between 1860 and 1890, and likely exterminated by introduced cats, rats, and pigs. However, unconfirmed reports from the 1960s and 1984 suggest that it may still survive in higher montane forests. The New Caledonian owlet-nightjar (Aegotheles savesi) was long known only from a single specimen collected in 1880 near Nouméa. A second specimen dated from 1915 has also been discovered in an Italian museum, and there have been a handful of other isolated records since then. It seems likely that the species occurs in low numbers on remote forest massifs. The horned parakeet (Eunymphicus cornutus) was historically found throughout Grande Terre during the nineteenth century, but is today confined to the Panié Massif in northeastern Grande Terre (Provence Nord). The New Caledonian lorikeet (Charmosyna diadema) is known for certain only from two specimens collected on Grande Terre in 1859 and another in 1913. There are unconfirmed reports from the 1880s to the 1920s, in the 1950s, and finally in 1976.

The crow honeyeater (Gymnomyza aubryana) is a relatively large and spectacular bird historically found throughout Grande Terre, but now largely confined to a few small populations in the south-east. It is threatened mainly by habitat destruction due to logging and fires. The marbled gecko (Oedodera marmorata) is confined to the region of the Dôme de Tiébaghi Massif in far northwestern Grande Terre (Provence Nord). The large-scaled chameleon gecko (Eurydactylodes symmetricus) is confined to areas of forest and maquis shrubland in southern Grande Terre (Provence Sud). Sarasins’ crested gecko (Correlophus sarasinorum) is confined to a few forest remnants in southern Grande Terre (Province Sud). The Goro Plateau gecko (Bavayia goroensis) is confined to a few localities in southernmost Grande Terre (Provence Sud), where it is threatened by habitat destruction and introduced species. The sclerophyll gecko (B. exsuccida) is found patchily in north-western and central coastal Grande Terre (Provence Nord). The zebra forest skink (Kanakysaurus zebratus) is known only from two disjunct localities along the central-western coast of Grande Terre (Provence Nord), where it is threatened by loss of habitat and invasive species. The white-lipped skink (Lioscincus steindachneri) is known only from seven localities within three disjunct areas of central and north-eastern Grande Terre (Provence Nord). The New Caledonian leopard skink (Lacertoides pardalis) is known only from four localities in southern Grande Terre (Provence Sud), where it inhabits rocky outcrops within maquis shrubland and forest margins. It is threatened by invasive species and wildfires. The northern pale-hipped skink (Celatiscincus similis) is known only from four localities representing two widely separated subpopulations in northern Grande Terre (Provence Nord). Chazeau’s litter skink (Caledoniscincus chazeaui) is confined to a few isolated localities in north-eastern Grande Terre (Provence Nord). The earless dwarf skink (Nannoscincus mariei) is confined to southern Grande Terre (Provence Sud). Slevin’s dwarf skink (N. slevini) is known only from a few localities within a small area of Grand Terre. The gracile dwarf skink (N. gracilis) occurs in a few widely scattered populations throughout Grande Terre. All are threatened by loss of habitat and introduced species (particularly fire ants). The gracile burrowing skink (Graciliscincus shonae) is confined to a few localities in southern Grande Terre (Provence Sud). The orange-bellied burrowing skink (Simiscincus aurantiacus) is known only from a few localities in southern Grande Terre (Provence Sud). The Chaîne Centrale The Chaîne Centrale mountain range runs the length of the island and features five peaks over 1500 m. The highest point is Mount Panié, although Mount Humboldt is nearly as high.

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The ornate gecko (Bavayia ornata) is known only from the Panié and Tchingou massifs in northern Grande Terre (Provence Nord). The Coula earth skink (Geoscincus haraldmeieri) is known only from two specimens collected in the 1970s from central Grande Terre. The locality has since been intensively cultivated and no forest remains, suggesting that the species may be extinct. The Maruia skink (Phasmasaurus maruia) is confined to five massifs on the central-west coast of Grande Terre (Provence Nord). The red-tailed shiny skink (Sigaloseps ruficauda) is known only from the summits of Mount Mou, Mount Ouin and Mount Humboldt in southern Grande Terre (Provence Sud). Salier’s litter skink (Caledoniscincus orestes) occurs in three disjunct subpopulations in north-eastern, north-western and south-central Grande Terre, where it is threatened by habitat destruction and introduced species. The garrulous dwarf skink (Nannoscincus garrulus) is confined to forest fragments on Pic Ningua and Mount Çidoa in south-central Grande Terre (Provence Sud). The Poum Massif is located in far north-western Grande Terre (Provence Nord). The Poum striped gecko (Dierogekko poumensis) is confined to the Poum Massif, where it is seriously threatened by mining operations that are intended to remove the entire top of the mountain. The Panié Massif is located is located in north-eastern Grande Terre (Provence Nord). The bold-striped gecko (Dierogekko validiclavis) is known only from Mount Mandjélia and Mount Panié. The Mandjélia litter skink (Caledoniscincus terma) is confined to Mount Mandjélia. The Tnâno Massif is located is located in north-eastern Grande Terre (Provence Nord). The Tnâno dwarf skink (Nannoscincus exos) is confined to two disjunct localities on the Tnâno Massif, where it is seriously threatened by wildfires and introduced species. The Ouazangou-Taom Massif is located in north-western Grande Terre (Provence Nord). The Taom striped gecko (Dierogekko thomaswhitei) is known only from Mount Taom. The Taom marble-throated skink (Marmorosphax taom) is confined to the summit of Mount Taom. The Kaala Massif is located in north-western Grande Terre (Provence Nord). The Kaala striped gecko (Dierogekko kaalaensis) is known only from the Kaala Massif. The Kaala marble-throated skink (Marmorosphax kaala) is known only from the summit of the Kaala Massif. The Koniambo Massif is located in north-western Grande Terre (Provence Nord). The Koniambo striped gecko (Dierogekko koniambo) is confined to the region of the Koniambo Massif.

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The Dôme de Tiébaghi is located is located in northwestern Grande Terre (Provence Nord). The Dôme de Tiébaghi striped gecko (Dierogekko nehoueensis) is confined to the Dôme de Tiébaghi and adjacent lowlands, where it is threatened by mining activities. The Kopéto-Paéoua Massif is located in central-west coastal Grande Terre (Provence Nord). The Kopéto-Paéoua skink (Lioscincus vivae) is confined to the Kopéto-Paéoua Massif. The Kopéto-Paéoua dwarf skink (Nannoscincus manautei) is confined to the summit of the Kopéto-Paéoua Massif. The Boulinda Massif is located in central-west coastal Grande Terre (Provence Nord). The Boulinda marble-throated skink (Marmorosphax boulinda) is known only from a single locality near the summit of the Boulinda Massif. Mount Aoupinié is located in central Grande Terre (Provence Nord). Rankin’s dwarf skink (Nannoscincus rankini) is confined to the summit of Mount Aoupinié. Mount Ouin is located in south-central Grande Terre (Provence Sud). The Ouin marble-throated skink (Marmorosphax montana) is confined to two localities near the summit of Mount Ouin. Mount Koghis is located in south-western Grande Terre (Provence Sud). The New Caledonian long-eared bat (Nyctophilus nebulosus) is known only from Mount Koghis. Lowland Tropical Moist Forests Most of Grande Terre’s lowland moist forests have been lost and only sporadic fragments now remain, primarily in protected areas. The key striped gecko (Dierogekko inexpectatus) is confined to low-elevation forest fragments within a small area of the Poum Massif in far north-western Grande Terre (Provence Nord), where it is seriously threatened by loss of habitat and invasive fire ants. Greer’s tree skink (Epibator greeri) is known only from a single specimen collected in 1979 from an unknown locality in north-western Grande Terre (Provence Nord). The cryptic litter skink (Caledoniscincus cryptos) is known only from a single specimen collected from south-central Grande Terre (Provence Sud). Renevier’s litter skink (C. renevieri) is known only from a small area of east-central Grande Terre (Provence Nord). The Koumac litter skink (C. auratus) is found patchily along the north-western coast of Grande Terre (Provence Nord). All are threatened by loss of habitat and invasive species. Greer’s dwarf skink (Nannoscincus greeri) is known only from three localities on the north-eastern coast of Grand Terre (Provence Nord). The moist dwarf skink (N. humectus) is confined to two localities in north-central Grande Terre (Provence Nord). Both are threatened by loss of habitat and invasive species.

Balance for the Papua-Melanesian Realm

Lowland Tropical Dry Forests Fragments of lowland dry sclerophyll and mesophyll forest are found along the western coast of Grande Terre. The western chameleon gecko (Eurydactylodes occidentalis) is confined to two forest remnants in central-western coastal Grande Terre (Provence Sud), where it is threatened by loss of habitat, invasive species and collection for the international pet trade. The Pindai dwarf skink (Nannoscincus hanchisteus) is confined to small, remnant pockets of closed sclerophyll forest on the Pindai Peninsula of central-western Grande Terre (Provence Nord). Tropical Dry Grassland Areas of tropical dry grassland occur on the western and northern coasts of Grande Terre. The New Caledonian buttonquail (Turnix novaecaledoniae) is known only from two specimens collected prior to 1912 and some fossil material found in caves. It is now thought to be extinct. The New Caledonian nightjar (Eurostopodus exul) is known from a single specimen collected in 1939. It is most likely extinct. Lakes, Rivers, and Marshes Grande Terre is covered with rivers and streams originating from the island’s mountains, which are home to a number of unusual endemic fishes. The New Caledonian gallinule (Porphyrio kukwiedei) was a turkey-sized marshland species known only from subfossil remains, although it is believed to have survived well into the nineteenth century. The New Caledonian galaxias (Galaxias neocaledonicus) is a type of freshwater fish confined to two small artificial lakes in south-eastern Grande Terre (Lac en Huit and Grand Lac), where it is threatened by introduced species. The New Caledonian silverside (Bleheratherina pierucciae) is known only from a dozen specimens collected from two rivers in southern Grande Terre, and from a third locality in the north. Sarasin’s goby (Sicyopterus sarasini) was historically widespread in southern Grande Terre, but is currently known only from four rivers in Provence Sud. It is threatened by habitat degradation due to nickel-mining activities. The sooty goby (Schismatogobius fuligimentus) is known only from a few streams in southern Grande Terre (Provence Sud), where it is threatened by nickel-mining activities. The Grande Terre loach goby (Protogobius attiti) is confined to a handful of rivers in southern Grande Terre (Provence Sud). The Dumbéa River is located in south-western Grande Terre (Provence Sud). The Dumbéa River pipefish (Microphis cruentus) is known only from a few specimens collected from the Dumbéa River.

The Isle of Pines Located to the south of Grande Terre, the Isle of Pines (L’Îledes-Pins in French) is a former penal colony that today is a major tourist hot-spot. Bocourt’s terrific skink (Phoboscincus bocourti), the largest skink in the world, was long thought to be extinct as it had not been recorded since the collection of a single specimen in 1876. It was rediscovered in 1993 on a small uninhabited offshore islet, and may persist on others as well. The southern pale-hipped skink (Celatiscincus euryotis) is known only from two localities on the Isle of Pines.

The Loyalty Islands The Loyalty Islands (Îles Loyauté in French) lie about 100 km north-east of New Caledonia. The archipelago consists of six inhabited coral islands as well as several smaller uninhabited islands and islets. The Loyalty Islands bent-winged bat (Miniopterus robustior) is known only from the islands of Lifou and Maré. The strand gecko (Bavayia crassicollis) is known only from the islands of Lifou and Maré. The Loyalty Islands emo skink (Emoia loyaltiensis) is known only from the islands of Lifou and Maré. Willey’s blind snake (Ramphotyphlops willeyi) is known only from three specimens collected from the islands of Lifou and Maré. It was last recorded in 1939. Lifou Lifou (Île de Lifou in French) is the largest and most heavily populated of the Loyalty Islands. Two subspecies of ground-nesting island thrush (Turdus poliocephalus) have become extinct from the Loyalty Islands. The Lifou thrush (T. p. pritzbueri) could not be found during an expedition to the island in 1939. Maré Maré (Île de Maré in French) is the second largest of the Loyalty Islands. The Maré thrush (Turdus poliocephalus mareensis) could not be found during an expedition to the island in 1939, and is presumed extinct. Ouvéa Ouvéa (Île de Ouvéa in French) is comprised of Ouvéa Island, the smaller Mouli Island and Faiava Island, and several islets surrounding the three. The Ouvéa parakeet (Eunymphicus uvaeensis) is confined to Ouvéa, where it is threatened by invasive species, loss of habitat and illegal collection for the international pet trade.

Balance for the Papua-Melanesian Realm The first people to arrive in Melanesia likely came from southern Asia tens of thousands of years ago. For some of these migrants the endpoint in their journey was the ancient

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subcontinent of Sahul, a single land mass comprising both what is now Australia and New Guinea which, at the time, were united by a land bridge due to lowered sea levels. When sea levels begin to rise once more the Australian and island aboriginal populations become isolated from one another. The aboriginal people of New Guinea developed one of the earliestknown examples of irrigated agriculture. Prior to the midtwentieth century European maps still showed the highlands of New Guinea as ‘terra incognito’, but when first flown over by aircraft numerous Stone Age settlements were discovered. The first European contact with New Guinea was by the Portuguese explorer Jorge de Menezes in 1526–27, who landed first on the island of Biak before proceeding on along the northern coast of the Bird’s Head Peninsula. In 1568 the Spanish sailor Álvaro de Mendaña reached the Solomon Islands. In 1606 the Portuguese explorer Pedro Fernandes de Queirós discovered Espiritu Santo, the largest island in what is now Vanuatu. In 1616 the Dutch explorers Jacob Le Maire and Willem Schouten discovered several islands in the Bismarck Archipelago (including New Hanover and New Ireland), and in 1644 Dutch explorer Abel Tasman discovered Fiji and New Britain. In 1774 the British explorer James Cook first sighted the island of Grande Terre and named it ‘New Caledonia’. Eventually, this name was applied to it as well as the surrounding islands. European colonization efforts began during the nineteenth century. In 1828 the Netherlands formally claimed

the western half of the island of New Guinea, and in 1883, following a short-lived French annexation of New Ireland, the British colony of Queensland seized south-eastern New Guinea. Germany claimed north-eastern New Guinea the following year. In 1905 the British government transferred some administrative responsibility over south-eastern New Guinea to Australia (who renamed the territory ‘Papua’). During the First World War Australian forces seized German New Guinea. After a brief Japanese occupation during World War II, New Guinea reverted to colonial status before ultimately achieving independence. Throughout this long period human impact on the environment had been minimal on New Guinea itself, although many of the smaller islands were devastated. In recent historical time (i.e. since A . D . 1500), the PapuaMelanesian Realm has lost at least 11 species/2 subspecies of vertebrates. Among the extinct forms 5 species are mammals and 6 species/2 subspecies are birds. Five other species are possibly extinct. In addition, there are 564 species/11 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 104 species/6 subspecies are mammals, 103 species/5 subspecies are birds, 139 species are reptiles, 160 species are amphibians, and 58 species are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

5 species

1 species

~ species

104 species

~ subspecies

~ subspecies

~ subspecies

6 subspecies

5 taxa

1 taxon

~ taxa

110 taxa

6 species

2 species

~ species

103 species

2 subspecies

~ subspecies

~ subspecies

5 subspecies

8 taxa

2 taxa

~ taxa

108 taxa

~ species

1 species

~ species

139 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

1 taxon

~ taxa

139 taxa

~ species

~ species

~ species

160 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

~ taxa

~ taxa

160 taxa

~ species

1 species

~ species

58 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

1 taxon

~ taxa

58 taxa

11 species

5 species

~ species

564 species

2 subspecies

~ subspecies

~ subspecies

11 subspecies

13 taxa

5 taxa

~ taxa

575 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

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Chapter

7

The Australian Realm

The Australian Zoogeographic Realm comprises the island continent of Australia and nearby islands, the largest of which is Tasmania. There has long been much confusion over the terms ‘Australian’ and ‘Australasian’, and a number of competing definitions as to what exactly should be included within them. From a purely geological standpoint Australia, New Zealand and New Caledonia are all fragments of the ancient supercontinent Gondwana. These three landmasses have been separated from other continents, and from each other, for tens of millions of years. Australia, New Guinea, and Tasmania, for their part, are separated from one another by shallow continental shelves and were linked together when the sea level was lower during the ice ages. They therefore share a similar fauna, which includes marsupial and monotreme mammals and ratite birds (New Zealand also has ratite birds, the kiwis along with the now-extinct moas), along with certain common floristic elements. From a biogeographic perspective, however, the fauna and flora of Australia (together with Tasmania), New Guinea and New Zealand, while sharing the aforementioned similarities, are all highly unique in their own ways. They are accordingly treated here as separate. Australia itself seems to have been isolated by water barriers longer and more effectively than any other continent, except perhaps for Antarctica. This explains why it is so rich in unique animals of great scientific interest. Surrounded by the Indian and Pacific oceans, its great size gives it a wide variety of landscapes, with tropical rainforests in the north-east, mountain ranges in the east and south-west, and dry desert and semi-desert in the centre. It is mostly flat with few large rivers, and has therefore few geographical barriers to the spread of plants and animals. But there are climatic boundaries. Thus, low precipitation has had a tremendous influence. Practically the whole continent is arid, one-third of it having an annual rainfall not exceeding 25 cm and the average precipitation throughout Australia just 38 cm, with high evaporation. Only about one-third of the continent can be cultivated, chiefly in the coastal fringe that receives sufficient rainfall to grow crops or artificial pastures. Twenty-five per cent of Australia is desert and 47 per cent arid or semi-arid grassland. Despite this dry climate, the vegetation varies greatly, ranging from tropical in the north to temperate in the south. Australia’s relatively few woods and forests, which are unfortunately concentrated along the eastern coast (the most heavily populated area), are dominated by eucalypts.

They grow in open or closed communities with thin or dense undergrowth of shrubs and herbs in wet or dry habitats. The resulting deforestation has had serious repercussions on the fauna, because many species were restricted to this habitat and others took refuge there when they were persecuted in the plains. In terms of its animal life, Australia is a world unto itself. Its small size as a continent, isolated location, and the length of time it has been separated from other continents have led its flora and fauna to follow other lines of evolution than those that can be traced elsewhere. In particular, the vertebrate fauna of this region represents several ancient elements that do not exist elsewhere. They came from Asia a long time ago, were geographically isolated, and survived after their Asian ancestors died out. They then evolved and radiated into many different species, which adapted to local habitats. The great specialization and differentiation of the marsupials (pouched mammals) is an outstanding example of such evolution and adaptation. They have filled habitats that on other continents are occupied by a large number of placental mammals of various other classes: primates, rodents, carnivores, and ungulates. This predominance of pouched animals among the mammals of Australia will be realized when it is noted that before the colonization of the continent by Europeans there were only two other orders of mammals there: rodents and bats (if man himself and the dingo he introduced are excepted). The success of pouched animals in Australia must be seen in the light of the fact that they had no competition from other orders of mammals, and that they were peculiarly able to adjust themselves to the rather difficult conditions of life in Australia. It is wrong, therefore, to regard – as is often done – pouched animals as failures because they have not evolved into the higher forms found on other continents. On the contrary, marsupials have shown a degree of adaptability that not even the most specialized mammals on other continents can equal. Perhaps the most famous animal resident of Australia is the koala (Phascolarctos cinereus). This species is entirely dependent for food on a few species of Eucalyptus. Before the arrival of Europeans, it was plentiful throughout eastern Australia from northern Queensland to the south-eastern corner of mainland South Australia. It was some time before the Europeans discovered that the fur of the koala was

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The Australian Realm

valuable; then a reckless persecution was initiated. The koala was a peculiarly helpless prey: sitting or climbing slowly among the light foliage of the crowns of the eucalypt trees, a myriad was shot or trapped. The koala was also vulnerable in the sense that it was too attached to the eucalypts and could not move to safer habitats. Moreover, it is a slow-breeding animal and the young were killed or captured with their mothers. The slaughter brought the koala near extinction. In 1920 only 500 koalas reportedly remained in Victoria, and the total Australian export was two million skins. Three years later, 10,000 licensed trappers in Queensland took half a million koalas, only half the number taken in 1921. By 1930 the koala had become extremely rare in New South Wales as a result of exploitation and disease. This extermination of such a harmless and unique animal is one of the more shameful episodes in the destruction of nature in Australia. Fortunately, public opinion finally reacted strongly to the slaughter, and the authorities proclaimed complete protection for the koala and prohibited the export of its skins. The koala was saved at the very last moment, and has made a remarkable recovery in the decades since. However, it remains threatened by loss of habitat and stochastic events such as droughts and bushfires. It currently occurs in suitable areas in north-eastern, central and south-eastern Queensland, with patchy populations in western areas, eastern New South Wales including the coastal strip and highlands of the Great Dividing Range and the western plains, Victoria, and south-eastern South Australia. It has also been introduced to at least a dozen islands, including Magnetic Island in Queensland, French and Philip islands in Victoria, and Kangaroo Island in South Australia. The crescent nail-tail wallaby (Onychogalea lunata) was once widespread in semi-arid south-western Australia, where it was described as abundant in the western Australian wheatbelt in the early 1900s. The species was probably wiped out by a combination of introduced predators, habitat degradation and perhaps disease. The last individual to be collected alive was caught in a dingo trap on the Nullarbor Plain in the late 1920s, which was later sent to the Taronga Zoo in Sydney. The last known specimen was killed in 1956, although reports indicate that the species may have hung on in some restricted desert areas. In any case it is now certainly extinct. The bridled nailtail wallaby (O. fraenata) was also historically found over a wide area extending through the inland regions of eastern Australia, from south-western South Australia to as far north as the base of the Cape York Peninsula. It was said to have been common at the time of European settlement, but declined catastrophically during the nineteenth and early twentieth centuries, probably due to introduced predators. By 1937 it was thought to be extinct. In 1973, however, an isolated population was discovered near the town of Dingo in central Queensland, on a property now protected as Taunton National Park. Reintroduced populations have been established from there to three other protected areas in Queensland and New South Wales.

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The Parma wallaby (Macropus parma), the smallest member of its genus, was first described by British naturalist John Gould in northern New South Wales around 1840. A shy, cryptic species, it was never commonly encountered and indeed was thought to have been extinct by the end of the nineteenth century. In 1965, however, workers on Kawai Island, New Zealand unexpectedly discovered a substantial population. The animals had apparently been imported and released there about 1870, together with three other species of Australia wallabies. These were captured and sent to institutions around the world for purposes of captive breeding, in the hope that they could eventually be reintroduced into their native habitat. Another major discovery took place in 1967, when it was found that the species still existed in forests near Gosford, New South Wales. Before long other populations were discovered in forests of the Great Dividing Range as far north as the border with Queensland. Although not common, it is no longer considered to be seriously threatened. Several species of rock wallaby (Petrogale) are threatened by habitat destruction and predation by foxes, dogs, and feral cats. The Mareeba rock wallaby (P. mareeba) is confined to a small area west and south-west of Cairns in north-eastern Queensland, where it lives in a handful of discontinuous colonies. Godman’s rock wallaby (P. godmani) lives in a series of discontinuous colonies within a small area north of Cairns in north-eastern Queensland. The Cape York rock wallaby (P. coenensis) is found patchily in the eastern part of the Cape York Peninsula of far northern Queensland. The Proserpine rock wallaby (P. persephone) is confined to a small area of central-eastern coastal Queensland, mainly within protected areas. A translocated population has also been established on Hayman Island from captive-bred stock. The purple-necked rock wallaby (P. purpureicollis) is confined to north-western Queensland, where it lives in a series of discontinuous colonies. The pygmy rock wallaby (P. concinna) is found patchily in northern Australia (north-eastern Western Australia including four offshore islands, and northern parts of Northern Territory). Burbidge’s rock wallaby (P. burbidgei) is confined to a small coastal area of the Kimberley region in north-eastern Western Australia, including a few offshore islands. Subfossil material indicates that it formerly had a wider range. The brush-tailed rock wallaby (P. penicillata) historically had a wide range across eastern Australia, but is now confined to scattered populations in eastern New South Wales and southeastern Queensland. Introduced populations occur in Hawaii and (formerly) New Zealand. The black-flanked rock wallaby (P. lateralis) is generally divided into three subspecies. The nominate form (P. l. lateralis) was historically widespread through most of central and north-central Australia wherever suitable rocky habitat occurred through most, along with a few islands off the western and southern coast. Owing to predation by introduced European red foxes (Vulpes vulpes) and, to a lesser extent, feral dogs and cats it has been extirpated over most of its former range and now survives only in a few widely scattered areas of Northern Territory, Western Australia, and

The Australian Realm

South Australia. The yellow-footed rock wallaby (P. xanthopus) was described as common in parts of South Australia, New South Wales, and Queensland in the 1920s, but declined markedly as a result of hunting for its very attractive and valuable pelt. Fragmented populations of the southern yellow-footed rock wallaby (P. x. xanthopus) persist in South Australia within the Gawler, Flinders, and Olary ranges, and in the Gap and Coturaundee ranges in New South Wales. The northern yellowfooted rock wallaby (P. x. celeris) has a restricted distribution in the rocky ranges of central-western Queensland. Hare-wallabies of the genus Lagorchestes are a group of small macropods that have been much reduced since the nineteenth century by habitat destruction and introduced predators. The eastern hare-wallaby (L. leporides) was a little-known species that occurred in central New South Wales, north-western Victoria and eastern South Australia. Subfossil material showed that it formerly occurred in southern Queensland as well. It was reported to be common in the 1850s on the plains around the junction of the Murray and Darling rivers, but declined rapidly after that owing to the alteration of its grassland habitat. It was last recorded in 1890. The rufous hare-wallaby (L. hirsutus) was formerly widely distributed in central and western Australia, where it was divided into two named subspecies and a third undescribed one, now extinct, from the central deserts. The nominate form (L. h. hirsutus), from south-western Australia, is also now extinct. The Shark Bay rufous hare-wallaby (L. h. bernieri) has long been restricted to two natural populations in the Bernier and Dorre islands of Shark Bay, Western Australia. It has also been introduced on nearby Trimouille Island and, more recently, to a few protected areas on the mainland (Western Australia, Northern Territory, and New South Wales). The banded hare-wallaby (Lagostrophus fasciatus) formerly occurred in south-western Australia, the Nullarbor Plain, southern South Australia, and western Victoria, but was exterminated on the mainland at the beginning of the twentieth century and henceforth confined to Bernier and Dorre Islands. It has since been successfully introduced to Faure Island in Shark Bay. The short-tailed scrub wallaby or quokka (Setonix brachyurus) is a cat-sized, nocturnal species that was historically widespread and abundant throughout south-western Australia including Rottnest, Bald, and Breaksea islands. Populations are now severely fragmented due to a drying climate and predation by introduced red foxes. Bettongs (Bettongia) are rabbit-sized jumping marsupials sometimes known as rat kangaroos. The burrowing bettong (B. lesueur), the only burrowing species of the kangaroo family and once distributed almost throughout Australia, is today restricted to the Bernier, Dorre, and Barrow islands off Western Australia. This tremendous decrease was caused by the spread of pasture lands, disturbance by man and cattle, competition for water, hunting, predation by dogs and foxes, and so on.

The northern hairy-nosed wombat (Lasiorhinus krefftii) formerly ranged from central Queensland to at least southcentral New South Wales, although fossil material has been found as far south as Victoria. Today it is entirely confined to a small area of open eucalypt woodland in Epping Forest National Park, central Queensland. In 2008 the total population was just 115 animals, up from 30–40 in the early 1980s. In 2000 dingoes killed 15–20, after which a 20-km long fence was built, encompassing the entire population. The northern quoll (Dasyurus hallucatus) is a small carnivorous marsupial that historically occurred across northern Australia from Pilbara to south-eastern Queensland, including offshore islands. Owing to habitat destruction and introduced species it is now confined to a few disjunct areas. The eastern quoll (D. viverrinus) was historically widespread across mainland south-eastern Australia including New South Wales, Victoria, and eastern South Australia, but became extinct there by the mid-1960s due to introduced predators and human persecution. It was thereafter entirely restricted to Tasmania and Bruny Island, although starting in 2016 there have been limited reintroductions to fenced-in protected areas in New South Wales and near Canberra. The tiger quoll (D. maculatus) is divided into two subspecies. The southern tiger quoll (D. m. maculatus) was historically found throughout the wet forests of south-eastern Queensland, eastern New South Wales, Victoria, South Australia, and Tasmania (including some of the Bass Strait Islands). It has been extirpated in many areas due to habitat destruction, introduced predators and human activities. The northern tiger quoll (D. m. gracilis) is confined to a small area of north-eastern Queensland, where it is also threatened mainly by loss of habitat and introduced predators. The numbat (Myrmecobius fasciatus) is a small insectivorous marsupial historically widespread across western, central, and southern Australia. The deliberate introduction of European red fox (Vulpes vulpes) during the nineteenth century resulted in its almost complete eradication. The nominate form (M. f. fasciatus) had, by the 1970s, been reducted to two small areas near Perth where the total population was thought to be less than 1000. It has since been reintroduced to other protected areas in Western and South Australia. The rusty numbat (M. f. rufus) has been extinct since at least the 1960s. The desert bandicoot (Perameles eremiana) was at one time widespread in the Great Sandy, Gibson, and Tanami Deserts, as well as in the Central Ranges region. It appears to be another victim of introduced predators, the last specimen having been collected from Well 33 on the Canning Stock Route in 1943. Aboriginals of the Western Desert indicated that it disappeared sometime in the mid-twentieth century, with some reporting having eaten it near Lake Mackay as recently as the late 1960s. The eastern barred bandicoot (P. gunnii) has been extirpated from its historical range on the mainland of Australia, but survives in two small introduced populations in Victoria and on the island of Tasmania. The western barred bandicoot (P. bougainville) formerly occurred over vast areas of southern Australia from Western Australia

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to central New South Wales, but by the late twentieth century was confined to Bernier, Dorre, and Faure islands in Shark Bay. It is currently being reintroduced to selected mainland areas where predators such as red fox are the subject of control programmes. The golden bandicoot (Isoodon auratus) is divided into three subspecies that, together, were historically widespread in western, central, and northern Australia. Owing to introduced predators it is now almost entirely extirpated from the mainland, surviving only on isolated islands. The nominate form (I. a. auratus) is confined to a small area of north-eastern coastal Western Australia and from Augustus and Uwins islands. The Arnhem Land golden bandicoot (I. a. arnhemensis) is found in an undefined area of coastal Northern Territory. Two species of pig-footed bandicoot (Chaeropus) historically inhabited much of Australia, where they were reportedly rare even before the arrival of Europeans. Habitat changes, rather than introduced species, are thought to have brought about their decline. The southern pig-footed bandicoot (C. ecaudatus) is believed to have been extinct since the midtwentieth century after vanishing from its last refuge in southern Australia by 1945. The northern pig-footed bandicoot (C. yirratji) from central Australia was last collected near Alice Springs in 1901. Aboriginal records, however, indicate populations surviving in the Gibson and Great Sandy deserts into the 1950s. It too is now considered to be extinct. The scaly-tailed possum (Wyulda squamicaudata) is an arboreal species confined to the Kimberley region of coastal north-western Australia and adjacent islands. Long known only from four specimens, two of which were collected in 1954, it was rediscovered in 2010. In more recent years populations have been found in a number of localities. The greater glider (Petauroides volans) is a large, arboreal marsupial that remains widespread across eastern Australia from northern Queensland to southern Victoria, but is everywhere under threat by habitat destruction. In 2020 the species was taxonomically split into three rather ill-defined forms. The broad-faced potoroo (Potorous platyops) is a littleknown, small terrestrial marsupial that, to judge from subfossil remains, was once widespread in the semi-arid coastal regions of South Australia and Western Australia, possibly as far north as North West Cape and as far east as Kangaroo Island. It was first collected in 1839, at which time it was already rare, and few other live specimens were ever recorded. The last record is from 1875, when five were sold to the National Museum in Victoria. It is thought to have been driven extinct by introduced cats and bush fires. A surviving species, the long-footed potoroo (P. longipes), is confined to three disjunct areas of south-eastern Australia on the New South Wales–Victoria border, where it is threatened mainly by predation from foxes, dingoes, and feral dogs. The sandhill dunnart (Sminthopsis psammophila) is a mouse-like marsupial that was long known only from a single specimen collected near Lake Amadeus, Northern Territory in 1896. In recent years it has been recorded from a small number

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of widely separated areas within the Great Victoria Desert and on the Eyre Peninsula in southern Australia. Two of Australia’s largest rodents are now extinct. The Capricorn rabbit-rat (Conilurus capricornensis) is known only from fossil and subfossil material collected from four localities in Queensland. It is believed to have disappeared after the arrival of Europeans to north-eastern Australia, and may have survived up until very recently. The white-footed rabbit-rat (C. albipes) historically had a wide range in areas of eucalyptus woodland from south-eastern South Australia and Victoria to New South Wales and eastern Queensland. Last recorded in 1860–62 (although with possible sightings as late as the early 1940s), it appears to have been exterminated by a combination of settlement, domestic cats, and introduced foxes. A surviving species, the brush-tailed rabbit-rat (C. penicillatus), is divided into two subspecies found in Australia and New Guinea. The Australian brush-tailed rabbit-rat (C. p. penicillatus) historically occurred throughout much of the monsoonal northern areas, but has disappeared from most of its former range due to habitat destruction and degradation, and most likely feral cat predation as well. The only recent records are from the Cobourg Peninsula in Northern Territory, the Mitchell Plateau and Prince Regent National Park in Western Australia, and from Bathurst, Melville, and Centre Island and Groote Eylandt. The greater stick-nest rat (Leporillus conditor) became extinct on mainland Australia in the 1930s, surviving only on a few offshore islands along the southern coast. It has since been translocated to other predator-free islands as well as to a fenced-off area at Roxby Downs in South Australia. The broad-toothed rat (Mastacomys fuscus) occurs at present only in small relict colonies in the Kosciuszko National Park in New South Wales and parts of southern Victoria and Tasmania. The black-footed tree rat (Mesembriomys gouldii) was historically found discontinuously in northern Australia, specifically in a small area of north-eastern coastal Western Australia, the northern coast of Northern Territory including Melville Islands, and north-eastern Queensland. There have been no records from the Kimberley region since 1982 despite considerable survey efforts. Threats include loss of habitat and predation by feral cats. The Carpentarian rock rat (Zyzomys palatalis) is known only from a few gorges and associated rocky areas in northcentral coastal Australia (north-eastern Northern Territory). The Arnhem Land rock rat (Z. maini) is confined to the Arnhem Land Plateau and nearby rocky outcrops in northcentral Northern Territory. Both are threatened by loss of habitat and predation from feral cats. Members of the genus Pseudomys are small nocturnal rodents similar in appearance to the common house mouse (Mus musculus). The blue-grey mouse (P. glaucus) is known only from three historical specimens, two collected in southeastern Queensland and a third from north-eastern New South Wales. It is presumed to be extinct, a victim of habitat

The Australian Realm

destruction and predation by introduced foxes and feral cats. Field’s mouse (P. fieldi) was once found throughout the western two-thirds of Australia, but was extripated by introduced species during the late nineteenth century. Long reduced to a few coastal sand dunes on Bernier Island, in 1999 it was successfully translocated to North West Island in the Montebello Islands, and to Faure Island in Shark Bay in 2003. The Hastings River mouse (P. oralis) is patchily distributed in south-eastern Queensland and north-eastern New South Wales. The smoky mouse (P. fumeus) is confined to a few disjunct areas of southeastern New South Wales and Victoria. Hopping mice (Notomys) are similar to North American kangaroo rats (Dipodomys), and as such are an interesting example of parallel evolution. The big-eared western hopping mouse (N. macrotis) is known only from a single specimen collected in 1843 near the Moore River, Western Australia. It appears to have been restricted to the western margin of the wheatbelt, where it may have survived well into the twentieth century. The Darling Downs hopping mouse (N. mordax) is known only from a single skull collected in 1922 from southern Queensland. The robust hopping mouse (N. robustus) is known only from skulls taken from old owl roosts within the Flinders and Davenport ranges of South Australia. Never collected alive, the species is now thought to be extinct. All three are thought to have been exterminated by introduced predators such as foxes and feral cats. The grey-headed flying fox (Pteropus poliocephalus) is found across eastern coastal Australia from south-eastern Queensland, through eastern New South Wales to southern Victoria. There has been contraction of the northern extent of the range in recent years due to loss of foraging and roosting habitat, with a corresponding increase in the number of permanent colonies in the south. The ghost bat (Macroderma gigas) was historically found across northern and central Australia but now has a patchy distribution within the Pilbara and Kimberley regions of Western Australia, the Top End of Northern Territory, northern and eastern Queensland, and possibly parts of New Guinea. It is threatened mainly by loss of its roosting sites. Troughton’s pouched bat (Saccolaimus mixtus) is known only from a small number of specimens collected from the Cape York Peninsula in far north-eastern Australia and fron southern New Guinea. The eastern free-tailed bat (Micronomus norfolkensis) is largely confined to the eastern coastal plains and ranges of New South Wales and south-eastern Queensland, where it is threatened by loss of habitat. The original description from Norfolk Island appears to be erroneous, as the species has never been found there since. The Australian sarus crane (Antigone antigone gilliae) is a non-migratory subspecies confined to grassland and wetland areas of northern and north-eastern Australia (Northern Territory and Queensland). The Cape Barren goose (Cereopsis novaehollandiae) is a large grazing bird found in the coastal areas and islands of

southern Australia. During the mid-twentieth century it was intensively hunted in some areas and reduced to a low of perhaps 4000 in all, although numbers have since recovered. The Recherche Cape Barren goose (C. n. grisea) breeds only on the islands of the Recherche Archipelago, where the population has remained more or less stable at around 1000. Loss of suitable grazing areas is now the main threat to this subspecies, but for the moment it appears to be safe. The Australasian bittern (Botaurus poiciloptilus) is a large, cryptic, heron-like bird found very disjunctly in wetland areas of south-eastern and south-western Australia (including Tasmania), New Zealand, and New Caledonia. The overall population is very small and threatened by loss of habitat. The Australian painted snipe (Rostratula australis) is a type of migratory wading bird found sporadically in wetland areas across Australia. It is threatened by water drainage and diversion. The malleefowl (Leipoa ocellata) is a stocky, grounddwelling bird that was formerly widespread in semi-arid mallee scrub across southern Australia. As a species it is dependent for both food and reproduction on more or less untouched open forests, and is thought to have lost about half its historical range during the twentieth century alone. The male builds large pyramidal nesting mounds of decomposing leaves and grass which he then covers with sand, gravel, and debris. The eggs are deposited within these mounds, but foxes and dogs dig them out, destroying many clutches. Drought, grazing by sheep and bush fires have also had a serious impact. It is now largely reduced to three main population pockets. The grey falcon (Falco hypoleucos) is a widely but sparsely distributed species from the arid and semi-arid areas of Australia. It is threatened by habitat destruction and degradation. The buff-breasted buttonquail (Turnix olivii) is known only from a few localities within the Cape York Peninsula of north-eastern Australia (north-eastern Queensland). The night parrot (Pezoporus occidentalis) is one of the world’s most mysterious and elusive birds. A smallnocturnal species first described by John Gould in 1861, it inhabits the remote arid and semi-arid inland regions of Western Australia, Northern Territory, South Australia, and Queensland. Although it is possible that it persists throughout much of this area, sightings have always been scanty and anecdotal. There were no confirmed reports at all between 1912 and 1979, leading to speculation that it was extinct. Sightings since have been extremely rare and total population size remain unknown, although it is likely less than 250. The swift parrot (Lathamus discolor) is confined to southeastern Australia, where it breeds in a few areas of Tasmania during the summer and migrates north to south-eastern South Australia, Victoria, New South Wales, and south-eastern Queensland in winter. It is threatened mainly by nest predation in Tasmania by introduced suger gliders (Petaurus breviceps) and by loss of habitat. Coxen’s fig parrot (Cyclopsitta coxeni) is a small species known only from four small subpopulations in eastern

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Australia (south-eastern Queensland and extreme northeastern New South Wales). It appears to have historically declined due to the widepread clearance of rainforest. Its small range coupled with a paucity of recent sightings suggests a total population of a few hundred at most. The orange-bellied parakeet (Neophema chrysogaster) is a small species that was formerly found over a wide area of southern Australia. Habitat destruction and commercial trapping had already reduced it by the mid-twentieth century to just 50–70, but the total rose to around 150 by 2005. Unfortunately, the population has suffered a drastic decline since then for reasons that are unclear (although most likely due to disease and drought). The birds are currently known to breed only at a single site in south-western Tasmania, from where they migrate to coastal south-eastern South Australia and Victoria for the winter. A captive breeding programme has been established and currently numbers 300–350. The short-billed black cockatoo (Zanda latirostris) is found in woodlands, shrublands, and heathlands in southwestern Australia (Western Australia), where it is threatened by loss of habitat. The regent honeyeater (Anthochaera phrygia) was formerly common throughout eastern and south-eastern Australia but is now found patchily in south-eastern Queensland, eastern New South, Wales and Victoria. It is threatened mainly by loss of habitat. The painted honeyeater (Grantiella picta) is found widely but patchily across eastern and northern Australia, where it is threatened by loss of habitat. The helmeted honeyeater (Lichenostomus melanops cassidix) has suffered greatly from habitat destruction, and is now confined to a small area in south-eastern Australia near Melbourne. The rufous scrub-bird (Atrichornis rufescens) is divided into two subspecies. The northern rufous scrub-bird (A. r. rufescens) and the southern rufous scrub-bird (A. r. ferrieri) are both confined to a few small areas of montane forest in coastal south-eastern Queensland and north-eastern New South Wales, but historically ranged further into the surrounding lowlands. By the mid-twentieth century both had been on the brink of extinction due to loss of habitat, and while their status is generally improved populations remain small and continue to fluctuate. The eastern bristlebird (Dasyornis brachypterus) is a type of ground-dwelling, semi-flightless passerine bird divided into two subspecies. The southern eastern bristlebird (D. b. brachypterus) is confined to a few localities in coastal New South Wales and eastern Victoria, where in 2011 the total population was thought to be less than 50. The northern eastern bristlebird (D. b. monoides) is known from a few isolated localities in south-eastern Queensland and northeastern New South Wales. Both are mainly threatened by bushfires. The even-scaled earless dragon (Tympanocryptis uniformis) is known only from a single specimen collected in

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1948 from an imprecise locality in northern Australia (‘near Darwin’). Numerous targeted searches have so far failed to rediscover the species. The Corangamite water skink (Eulamprus tympanum marnieae) is a semi-aquatic subspecies confined to a few scattered localities in south-eastern Australia (southern Victoria). The broad-headed snake (Hoplocephalus bungaroides) is a venomous species confined to a few areas of sandstone outcropping near Sydney in New South Wales. Several semi-aquatic frogs of the genus Litoria are seriously threatened by loss of habitat, introduced predatory fish, and possibly chytridiomycosis. The Nyakala frog (L. nyakalensis), Day’s frog (L. dayi), Liem’s frog (L. rheocola), and the waterfall frog (L. nannotis) are all confined to a small area of coastal northeastern Queensland centered on Cairns. The green-thighed frog (L. brevipalmata) is found in coastal areas of south-eastern Queensland and north-eastern New South Wales. The yellowspotted frog (L. castanea), as currently defined, appears to be confined to two disjunct localities in north-eastern and southeastern New South Wales. Spencer’s frog (L. spenceri) is found patchily in south-eastern New South Wales and north-eastern Victoria. The green and golden frog (L. aurea) is found in coastal New South Wales and eastern Victoria, with introduced populations on North Island, New Zealand, New Caledonia, and Vanuatu. The growling frog (L. raniformis) remains widespread in south-eastern South Australia, southern New South Wales, Victoria and parts of Tasmania, with additional introduced populations throughout much of New Zealand, but has undergone a massive population decline. The Alexandria frog (Uperoleia orientalis) is known only from a few specimens collected in 1940 from a swamp in north-central Northern Territory. The Jabiru frog (U. arenicola) is known only from a small area in north-western Northern Territory, on the western edge of the Arhhem Land escarpment. The sunset frog (Spicospina flammocaerulea) is confined to a few isolated peat swamps in the extreme south-western corner of Western Australia. The yellow-bellied frog (Geocrinia vitellina) and the whitebellied frog (G. alba) are each confined to a small area of wetlands in extreme south-western Western Australia. The red-crowned brood frog (Pseudophryne australis) is confined to a small area of sandstone escarpments near Sydney, New South Wales. The silver-eyed barred frog (Mixophyes balbus) is a semiaquatic species from the eastern slopes of the Great Divide of south-eastern Australia (New South Wales and Victoria). It has been extripated from many areas where it was formerly common. Fleay’s barred frog (M. fleayi) and the giant barred frog (M. iteratus) are both confined to a small area of eastern Australia (south-eastern Queensland and north-eastern New South Wales), where they are threatened by loss of habitat. The sharpsnout torrent frog (Taudactylus acutirostris), the Kroombit Tops torrent frog (T. pleione), and the Eungella torrent frog (T. eungellensis) are all confined to rainforest

Mountains and Highlands

fragments in central-eastern coastal Queensland, where they are seriously threatened by loss of habitat, introduced species, and possibly chytridiomycosis.

Mountains and Highlands Australia consists mainly of lowland deserts and savannas with the notable exception of the eastern coast, where the Great Dividing Range predominates. For long stretches these mountains plunge straight into the Pacific Ocean, but here and there they slope gently down, leaving room for a lowland coastal strip sometimes 160 km wide. This lowland belt in Queensland, New South Wales, and Victoria contains a high proportion of the entire population of Australia, and is often the only part of the continent a casual visitor sees. Indeed, often Australia is identified with this well-known narrow strip of land and the extensive grasslands and deserts, while the mountains are forgotten. Yet these alpine regions are invariably quite close to the largest towns and cities and are among the continent’s most characteristic and splendid natural features. From tropical Queensland to the subtropical belts of New South Wales and Victoria they form a massive barrier of granite, sandstone, and basalt between the Pacific in the east and the vast plains in the west. It is also a boundary between climates and vegetation. From the east humid winds blow and from the west desert winds, while in the southern part, the Australian Alps, hurricanes often in conjunction with cyclones from Antarctica storm over the range. These climatic factors have given the Great Dividing Range a rich variety of natural environments, many of them not found elsewhere on the continent. Other, smaller highland areas are scattered all over the continent, from the Hamersley and Wunaamin Miliwundi ranges in Western Australia and the MacDonnell Range in the deserts of the Northern Territory, to the Musgrave and Flinders ranges in South Australia. Taken together, as in other parts of the world, they often serve as a last refuge for threatened species.

The Great Dividing Range The Great Dividing Range (also known as the Eastern Highlands) is Australia’s most substantial mountain chain and the third longest land-based range in the world. It stretches more than 3500 km from Dauan Island off the north-eastern tip of Queensland, running the entire length of the eastern coastline through New South Wales before finally turning west and fading into the central plain of western Victoria. It varies in width from about 160 to 300 km. In terms of vegetation the northern areas are dominated by tropical and subtropical rainforest, being replaced in the east and south-east by temperate broadleaf and mixed forests dominated by Eucalyptus. A few areas of temperate rainforest are also to be found in eastern New South Wales. The Mount Claro rock wallaby (Petrogale sharmani) is confined to the Seaview and Coane ranges in coastal northeastern Australia (north-eastern Queensland). The northern bettong (Bettongia tropica) is a type of ‘rat kangaroo’ confined to three localities in far north-eastern

Queensland, specifically the Lamb Range, the Mount Carbine Tableland, and the Coane Range. It may also survive on the Mount Windsor Tableland. The silver-headed antechinus (Antechinus argentus) is a mouse-like marsupial known only from Kroombit Tops National Park in south-eastern Queensland. Several semi-aquatic frogs of the genus Litoria endemic to the Great Dividing Range are seriously threatened by loss of habitat, introduced predatory fish, and possibly chytridiomycosis. The armoured frog (L. lorica) is known only from four montane localities in coastal north-eastern Queensland. Last seen in 1991 and thought to be extinct, a small population was rediscovered in 2008. The peppered frog (L. piperata) was historically known from five streams draining the east of the Northern Tablelands in north-eastern New South Wales. Assuming that it still survives at all, the total population is thought to be less than 50. Davies’ frog (L. daviesae) is confined to a few localities on the eastern escarpment of the Great Dividing Range in north-eastern New South Wales. The New England frog (L. subglandulosa) is confined to the eastern escarpment of the Great Dividing Range in north-eastern New South Wales and south-eastern Queensland. The Booroolong frog (L. booroolongensis) historically ranged throughout the Great Dividing Range in New South Wales, but has now disappeared from most areas. The southern gastric-brooding frog (Rheobatrachus silus) was restricted to the Blackall and Conondale ranges of coastal eastern Australia (south-eastern Queensland). It has not been recorded since 1981, and is now considered to be extinct. The magnificent brood frog (Pseudophryne covacevichae) is confined to a small area of north-eastern Queensland centered on the Glen Gorden Volcanics. Pengilley’s brood frog (P. pengilleyi) is confined to two small areas near Canberra in south-eastern New South Wales. Both are threatened by habitat destruction. Several frogs of the genus Philoria are threatened by habitat degradation. The red and yellow mountain frog (P. kundagungan) and Loveridge’s mountain frog (P. loveridgei) are both confined to small areas in south-eastern Queensland and north-eastern New South Wales. The sphagnum mountain frog (P. sphagnicolus) and Pugh’s mountain frog (P. pughi) are both confined to a small area of north-eastern New South Wales. The Richmond Range mountain frog (P. richmondensis) is known for certain only from three localities within the Richmond and Yabbra ranges in north-eastern New South Wales. Several species of rainforest frog (Cophixalus) are threatened by habitat degradation and wildfires. The elegant rainforest frog (C. concinnus) is confined to Thornton Peak in coastal north-eastern Queensland. McDonald’s rainforest frog (C. mcdonaldi) is confined to Bowling Green Bay National Park on Mount Elliott, south-east of Townsville in coastal north-eastern Queensland. The Black Mountain rainforest frog (C. saxatilis) is confined to Black Mountain National Park south of Cooktown in coastal north-eastern

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Queensland, where it lives in caverns formed by large boulders. The Woowoonooran rainforest frog (C. neglectus) is confined to Woowoonooran National Park between Cairns and Innisfail in coastal north-eastern Queensland. The tapping rainforest frog (C. aenigma) is known from the Mount Carbine Tableland, Thornton Uplands, Finnigan Uplands, and Bakers Blue Mountain in north-eastern Queensland. The Mount Glorious torrent frog (Taudactylus diurnus) historically occurred in disjunctive subpopulations in the Blackall, Conondale, and D’Aguilar ranges of coastal eastern Australia (south-eastern Queensland). In the early 1970s it was considered to be relatively common, but it declined quickly and dramatically over the next few years and has not been oberved in the wild since 1979. It is now considered to be extinct, although the reason remains a mystery. The tinkling torrent frog (T. rheophilus) is confined to five mountaintops in coastal north-eastern Australia (north-eastern Queensland). The species has undergone a significant population decline, most likely due to chytridiomycosis. The giant burrowing frog (Heleioporus australiacus) is confined to the lower eastern slopes of the Great Dividing Range in New South Wales and Victoria.

The McIlwraith Range The McIlwraith Range is a rugged granite plateau located on the Cape York Peninsula in far north-eastern Australia (northeastern Queensland). The long-necked northern leaf-tailed gecko (Orraya occultus) is confined to the McIlwraith Range. The McIlwraith rainforest frog (Cophixalus peninsularis) is known only from the McIlwraith Range.

The Melville Range The Melville Range is located on the Cape York Peninsula in far north-eastern Australia (northern Queensland). The Melville leaf-tailed gecko (Saltuarius eximius) is known only from a few specimens collected from a single locality within the Melville Range. The Melville shadeskink (Saproscincus saltus) is confined to the highest ridge within the Melville Range. The Melville rainforest frog (Cophixalus petrophilus) is confined to two disjunct boulder fields a few kilometres apart within the Melville Range.

The Clarke Range The Clarke Range is located in coastal north-eastern Australia (north-eastern Queensland). The northern gastric-brooding frog (Rheobatrachus vitellinus) was discovered in 1984 in the rainforests of the Clarke Range. It seems to have disappeared within a year, never to be recorded again despite extensive efforts to locate it.

The Mount Carbine Tableland The Mount Carbine Tableland is a plateau located in northeastern Australia (north-eastern Queensland).

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The Carbine rainforest frog (Cophixalus monticola) is confined to a small area of the Mount Carbine Tableland.

The Paluma Range The Paluma Range is located in coastal north-eastern Australia (north-eastern Queensland). It is protected within Paluma Range National Park. The Gulbaru leaf-tailed gecko (Phyllurus gulbaru) is confined to a few isolated populations in Patterson Gorge, at the extreme southern end of the Paluma Range.

The Blue Mountains The Blue Mountains are located in south-eastern Australia (eastern New South Wales). The Blue Mountains water skink (Eulamprus leuraensis) is a semi-aquatic species confined to a small area so-called ‘hanging swamps’ west of Sydney. It is threatened by mining operations and bushfires.

The Australian Alps The Australian Alps are located in south-eastern Australia (eastern Victoria, south-eastern New South Wales, and the Australian Capital Territory). The mountain pygmy possum (Burramys parvus) was long known only from fossil material and thought to be extinct until 1966, when a living specimen was discovered on Mount Hotham in the Victorian Alps (the southern part of the Australian Alps). In 1969 a colony was found near the treeline in Kosciuszko National Park, New South Wales. Since then, two more isolated populations have been discovered, one between Mount Bogong and Mount Higginbotham, and the other at Mount Buller, both in Victoria. The Snowy Mountains are located in south-eastern New South Wales. They are the highest mountains in Australia, with Mount Kosciusko the highest peak. The corroboree brood frog (Pseudophryne corroboree) was considered to be relatively common within its historically small distribution as recently as the 1970s, but has since suffered massive declines. It is now confined to a few fragmented populations totalling less than 200 within Kosciuszko National Park.

The Central Highlands The Central Highlands are located in south-eastern Australia (east-central Victoria). Leadbeater’s possum (Gymnobelideus leadbeateri) is a highly specialized non-gliding species almost entirely confined to small pockets of alpine ash, mountain ash, and snow gum forests north-east of Melbourne, with an additional isolated subpopulation occupying a low-elevation swamp forest south-west of the main range. Historically common within the areas it inhabited, the species is nevertheless elusive. It had not been discovered until 1867, and was long known only from five specimens, the last of which was collected in 1909. From then on it was feared extinct, a victim

Mediterranean Forests, Woodlands, and Scrub

of habitat destruction and the devastating Black Friday bushfires of 1939. However, in April 1961 one was spotted in the forests near Cambarville, and an individual was captured soon after. Additional searches resulted in the location of a colony near Marysville, and others were found in the years that followed. The combination of regrowth and the many large, dead trees (for nesting and shelter) from the 1939 fires allowed the population to expand to an estimated 7500 by the early 1980s. The population then began to decline sharply due to a habitat bottleneck. The Black Saturday bushfires of 2009 destroyed 43 per cent of the species’ remaining habitat and halved the wild population to just 1500.

The Baw Baw Plateau The Baw Baw Plateau is located in south-eastern Australia (south-eastern Victoria). It is protected within Baw Baw National Park. Frost’s mountain frog (Philoria frosti) is confined to alpine heathland on the Baw Baw Plateau, where it has declined considerably in recent years due to infectious chytridiomycosis fungus. A captive breeding programme has recently been undertaken.

The Macdonnell Ranges The MacDonnell Ranges are a 644-km long series of parallel ridges running to the east and west of Alice Springs in central Australia (south-central Northern Territory). The central rock rat (Zyzomys pedunculatus) has exhibited a very marked decline in both range and abundance since the European settlement of Australia, and is now confined to a few localities within the western MacDonnell Ranges. The main threats are introduced predators and habitat degradation.

Lowland Tropical and Subtropical Rainforests Deforestation in north-eastern Australia has been extensive, and most tropical and subtropical rainforest is now associated with the Great Dividing Range. Surviving patches of distinctive lowland rainforest are still to be found, however. Of these, the largest and most important is the Daintree Rainforest on the north-eastern coast of Queensland. The mahogany glider (Petaurus gracilis) is a cryptic and elusive species that had been lost to science for over a hundred years until its rediscovery in 1989. It is restricted to small areas of coastal lowland rainforest in north-eastern Queensland. The Kuranda frog (Ranoidea myola) is confined to a very small area of north-eastern coastal Queensland north of Cairns. The Cape Melville frog (R. andiirrmalin) is confined to a few localities in north-eastern Queensland. Both are threatened by loss of habitat and chytridiomycosis. Zweifel’s rainforest frog (Cophixalus zweifeli) is confined to a small area within Cape Melville National Park in northeastern coastal Queensland, where it lives in boulder fields among and adjacent to rainforest.

Mediterranean Forests, Woodlands, and Scrub Mediterranean forests, woodlands, and scrub are to be found in southern Australia (south-western and southern Western Australia, southern Southern Australia, northern Victoria, and south-western and central New South Wales). The brush-tailed bettong or woylie (Bettongia penicillata) was once a common species within eastern Australia, where it still exists locally in New South Wales and Victoria. The nominate form (B. p. penicillata) is now extinct, while the south-western subspecies (B. p. ogilbyi) survives only in a small population in south-western Western Australia. The freckled marsupial mouse or dibbler (Parantechinus apicalis) was widespread during the nineteenth century, but declined thereafter owing to the predations of feral foxes and cats. It was thought to have become extinct in the 1930s, until rediscovered at Cheyne Beach on the southern coast of Western Australia in 1967. Since then other populations have been found in the Torndirrup, Waychinicup, and Fitzgerald River national parks and the Arpenteur Nature Reserve. In 1985 it was discovered as well on Boullanger and Whitlock islands in Jurien Bay. The species has also been introduced to Escape Island in Jurien Bay and to other protected areas on the mainland. The sooty dunnart (Sminthopsis fuliginosus) was a mouselike marsupial known only from specimens collected during the 1840s from south-western Australia, and now most likely extinct. Gould’s mouse (Pseudomys gouldii) is known from living specimens as well as subfossil remains collected in southwestern Western Australia, eastern South Australia, and New South Wales. Last collected in the 1850s, it was likely driven extinct by feral cats and habitat degradation. In 2021, however, a genetic study appeared to indicate that the species was actually conspecific with Field’s mouse (P. fieldi), previously discussed, which survives on a few small islands off the coast of Western Australia. I have retained both species in this volume. The western ground parrot (Pezoporus flaviventris) was formerly widespread on coastal heaths and offshore islands, but is today found only in some swampy areas of the southern coastlands of Western Australia. Here again altered habitat, brush fires, intensive hunting (because its flesh was much esteemed), predation from foxes and domestic cats were more than the species could withstand. It is now one of the world’s rarest birds. The noisy scrub-bird (Atrichornis clamosus) is a small passerine bird that lives in densely vegetated, eucalyptdominated gullies on the southern coast of Western Australia. Destruction of its habitat led to such a decline in its numbers that it was considered extinct as early as 1889. In 1961, however, a population of about fifty birds was rediscovered in the Mount Gardner area, about 40 km east of Albany. After a well-publicized battle with land developers a small protected area was ultimately established for the species there, and as a result of intensive conservation work over the

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succeeding decades both the range and numbers have increased, but fires remain a threat. The black-throated whipbird (Psophodes nigrogularis) is divided into two subspecies. The western heath black-throated whipbird (P. n. nigrogularis) is confined to a small patch of scrubland in coastal south-western Australia (Western Australia), having disappeared from large parts of former range due to land clearance. The western mallee blackthroated whipbird (P. n. oberon) survives in a few isolated populations in coastal south-western Australia (Western Australia). The white-bellied whipbird (P. leucogaster) is also divided into two subspecies. The nominate form (P. l. leucogaster) occurs in scattered populations across north-western Victoria and southern South Australia. The rufous bristlebird (Dasyornis broadbenti) is a thrushlike, largely terrestrial bird that, as a species, is still relatively widespread in south-eastern coastal Australia. One subspecies is now extinct and will be discussed below. Of the remaining two, the Coorong rufous bristlebird (D. b. broadbenti) occurs in near-coastal habitats from Port Fairy, Victoria to the mouth of the Murry River in south-eastern South Australia. The Otways rufous bristlebird (D. b. caryochrous) was formerly thought to be largely confined to the coast between Peterborough and Point Addis in western Victoria, but is now known to occur extensively within the Otway Range.

Subtropical Mediterranean Forests and Woodlands Areas of subtropical Eucalyptus forest were formerly found throughout south-western and southern Western Australia. Sadly, their destruction has been pronounced and more than half of the state’s forests are under mining tenements. The western ringtail possum (Pseudocheirus occidentalis) already had a limited distribution in south-western Western Australia when if first became known in the 1880s. Considered to be already near extinction by the early 1960s, by the 1980s it was largely confined to the coastal strip between Bunbury and Albany, where it occurs in habitat fragments mainly on private lands. Efforts at translocation to protected areas have met with little success. Baudin’s black cockatoo (Zanda baudinii) is confined to heavily forested areas of south-western Western Australia, where it is threatened by loss of habitat.

Mallee Scrub Mallee scrub is a semi-arid region located in southern Australia (Western Australia, South Australia, New South Wales, and Victoria). The vegetation is dominated by mallee eucalypts, which are rarely over 6 m in height, while the understorey consists of hummock grasses and shrubs. Most of it has been destroyed since the arrival of Europeans, and today the largest remaining areas are to be found on the fringes of the Great Victoria Desert and in the Murray–Darling drainage. The black-eared miner (Manorina melanotis) is a type of honeyeater historically found throughout the mallee scrub

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region. Its range and numbers declined steadily over the twentienth century due to loss of habitat and interbreeding with the related yellow-throated miner (M. flavigula), and the species is now confined to two small, disjunct areas of south-eastern South Australia and a third in north-western Victoria. The mallee emuwren (Stipiturus mallee) has a severely fragmented distribution in mallee scrub regions of southeastern South Australia and north-western Victoria. The red-lored whistler (Pachycephala rufogularis) is a type of passerine bird confined to mallee areas of eastern South Australia and north-western Victoria, with isolated outlying populations in New South Wales. Large parts of its historically much wider range have been lost to habitat conversion, although in recent years the greater threat has been bushfires.

Coastal Heathlands The coastal heathlands of southern and south-western Australia are characterized by dense, low shrubs with scattered, twisted trees. Gilbert’s potoroo (Potorous gilbertii) is a small, nocturnal macropod currently thought to be Australia’s most endangered marsupial. First discovered in 1840, it was even then intensively hunted by Aborigines who could kill a great number in just a few hours. However, it was predation by introduced red foxes and feral cats that has been the greatest threat. Thought to have gone extinct in the 1970s until its unexpected rediscovery in 1994 in the Two People’s Bay Nature Reserve, Western Australia, it has since been successfully introduced to Bald Island Nature Reserve as well as to Waychinicup National Park. The total population in 2012 was estimated at 100, but this was approximately halved in 2015, mostly due to bushfires. The western bristlebird (Dasyornis longirostris) is endemic to the coastal heathlands of south-western Western Australia, where it was at one time thought to be extinct until a single bird was collected in 1945. The nominate subpecies (D. l. longirostris) was subsequently found in a handful of scattered localities, mainly within protected areas. The total population of around 1000 continues to be threatened by wildfires. The western rufous bristlebird (D. b. litoralis) was confined to a small stretch of south-western coastal Western Australia, where it was last reliably recorded in 1908 (although unconfirmed reports continued up until 1940). It is thought to have been driven to extinction by the deliberate burning of its habitat to create pastureland.

Savannas and Grasslands Along with deserts and subdeserts, the scrub and grasslands, ranging from steppe-like plains to true savannas, occupy the largest part of Australia. In a belt of varying width the scrub and grasslands almost entirely surround the deserts of the interior, but they are also found along some of the coasts, particularly in the northern parts of the continent. The broadest expanses of grassland are in eastern Australia west

Savannas and Grasslands

of the Great Dividing Range and east of the desert. There are two main types of scrub: the mulga scrub and the mallee scrub, the former dominated by mulga (Acacia aneura), while the latter is formed mostly of dwarf eucalypts, some shrubs, and sclerophyllous grass. Many former savannas have been converted into cultivated fields and grazing areas for livestock. Some grasslands still show their original vegetation, dominated by feather grass (Stipa), wallaby grass (Danthonia), and kangaroo grass (Themeda australis). The latter, notably nutritious, was once of great importance to Australia’s rich animal world, apparently playing the same role as its relative T. triandra does for many species of antelopes on the savannas of Africa. It was to such rich grassland that a multitude of grazing marsupials as well as many birds had adapted. The product of millions of years of natural selection, they roamed the plains freely and in harmony with the vegetation. Suddenly, with the arrival of the Europeans, the whole system was altered and disaster followed for many living things. Numerous smaller marsupials disappeared forever, destroyed not only by introduced predators and competition with sheep but also directly by human hunters. In many areas nutritious plants disappeared and were replaced by less-palatable species like spinifex. The plains wanderer (Pedionomus torquatus) is a quail-like ground bird historically found on the grasslands of northeastern South Australia, west-central Queensland, northcentral Victoria, and southern New South Wales. It has declined dramatically due to hunting pressure, drought, and introduced foxes. In 2015 the total population was estimated at less than 1000.

Tropical and Subtropical Savannas and Grasslands Tropical and subtropical savannas, grasslands, and shrublands, often insterspersed with areas of open or closed dry forest, are to be found across much of northern Australia (northern Western Australia, northern and eastern Northern Territory, northern and eastern Queensland, and parts of north-eastern New South Wales). The Kakadu pebble mouse (Pseudomys calabyi) is known only from a small area of Northern Territory, including Kakadu National Park and Litchfield National Park. The Arnhem leaf-nosed bat (Hipposideros inornatus) is known only from four roost sites within Kadadu National Park in north-central coastal Australia (Northern Territory), none of which are currently known to be occupied owing to human disturbance. The paradise parrot (Psephotellus pulcherrimus) is only known with certainty from areas of open savanna woodland and shrubby grassland in south-eastern Queensland, although it may perhaps have ranged a little more widely. Locally common although generally rare during the nineteenth century, it declined rapidly due to excessive hunting for commercial purposes, perhaps combined with other factors such as introduced cats (the species nested exclusively in termite

Figure 7.1 A live paradise parrot photographed by the entrance to its nest, Burnett River, Queensland, 1922. (Credit: C. H. H. Jerrard.)

mounds). It was thought to have become extinct as a result of a drought in 1902, until rediscovered in 1918. The last confirmed observation was in 1928. A few credible reports continued into the 1930s and 1940s, and it was widely believed, although without much evidence, to have survived in small numbers up to the 1970s. There was a further, doubtful report made in 1990, and nothing since. A related species, the golden-shouldered parrot (P. chrysopterygius), survives in the central and southern Cape York Peninsula of Queensland. It has been eliminated from most of its historic range, at first due mainly to illegal trapping for use as a cage bird. In more recent years loss of habitat has become the primary threat. The total population is estimated at around 3750 in two main subpopulations, and declining. The partridge pigeon (Geophaps smithii) is divided into two subspecies, both of which are declining due to habitat degradation. The Top End partridge pigeon (G. s. smithii) is now confined to northern coastal Northern Territory and the Tiwi Islands, but was formerly more widespread. The Kimberley partridge pigeon (G. s. blaauwi) is found in remote areas of extreme north-eastern Western Australia, where there are few recent records. The black grasswren (Amytornis housei) is very rare and has been seen only a few times in the Kimberleys of northeastern Western Australia, where it is confined to hummock grasslands in especially rugged sandstone environments. The white-throated grasswren (A. woodwardi) is confined to a small area of northern coastal Northern Territory in and around the Arnhem Land sandstone massif, where it is threatened by habitat destruction and degradation. The Carpentarian grasswren (A. dorotheae) is known from a few

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localities within the hills of north-western Queensland and north-eastern Northern Territory. Weigel’s toad (Notaden weigeli) is known only from the Kimberley region of north-western Australia (north-eastern Western Australia). The marbled frog (Uperoleia marmorata) is known only from a single specimen collected in 1841 from an undefined locality in northern Australia (north-eastern Western Australia). The Darwin frog (U. daviesae) is known only from a small area located south of Darwin in northern Australia (Northern Territory).

Tropical and Subtropical Wooded Savannas Areas of both open and closed tropical and subtropical dry forest, interspersed with savanna, are to be found in northern Australia (Western Australia, Northern Territory, and Queensland), including offshore islands. The northern brush-tailed phascogale (Phascogale pirata) is a small, arboreal, carnivorous marsupial confined to a small area of northern Australia’s Top End (Northern Territory), including Melville Island. The species formerly occurred as well in north-western Western Australia, but there have been no recent records from there. The fawn marsupial mouse (Antechinus bellus) is a type of small carnivorous marsupial found patchily in northern Australia (Northern Territory), including Melville Island. It is threatened by habitat destruction and predation by feral cats. The Crystal Creek two-lined dragon (Diporiphora convergens) is a type of agamid lizard known only from a single specimen collected from Crystal Creek, in the Kimberley region of north-western Australia (north-eastern Western Australia).

Allan’s lerista (Lerista allanae) and the Mount Cooper striped lerista (L. vittata) are fossorial lizards each known only from a small area of eastern coastal Queensland. Both are threatened by habitat destruction and degradation.

Temperate Savannas and Grasslands Temperate savannas, grasslands, and shrublands, often insterspersed with areas of open or closed dry forest, are found in eastern and south-eastern Australia (south-central Queensland, central and south-western New South Wales, and northern Victoria). The toolache wallaby (Macropus greyi) historically lived within open country in near-coastal areas of extreme southwestern Victoria to the upper south-east of South Australia. In the early nineteenth century this extraordinarily beautiful species was still common within its restricted range, but was becoming quite rare by the 1870s owing to loss of habitat, predation by introduced foxes, and in particular hunting. It is reported that hunters had become eager to kill the last survivors for their pelts or as trophies. By 1910 it had been reduced to a few scattered populations and in 1924 only one small group was known to survive, on Konetta Station about halfway between the towns of Robe and Penola in South Australia. An attempt was made to translocate some of the animals to a sanctuary on Kangaroo Island, but this failed. The last known individual, a doe that had been rescued from dogs, died in captivity in Robe in 1939. The grassland earless dragon (Tympanocryptis pinguicolla) is a type of small lizard known only from two disjunct areas of treeless grassland in south-eastern Australia (south-eastern New South Wales and Australian Capital Territory).

Figure 7.2 Toolache wallaby (Macropus greyi). (Credit: F.R.S., Mammals of Australia, Vol. II Plate 19, London, 1863, © John Gould.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

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Deserts and Shrublands

A population at least formerly occurred as well in southcentral Victoria, but has not been recorded there since 1969 despite extensive survey work and has most likely been extirpated. It is worth noting that a recent revision seems to indicate that the two populations outside of Victoria actually represent different taxa; if so, this species may well be extinct. The pygmy blue-tongued skink (Tiliqua adelaidensis) is confined to a small area of south-eastern South Australia, where it is was long believed extinct until its rediscovery in 1992. The striped legless lizard (Delma impar) is found patchily within remnant habitat in southern New South Wales, Victoria and the Australia Capital Territory. Sloane’s froglet (Crinia sloanei) is confined to southeastern Australia (central New South Wales and north-central Victoria), where it is associated with temporary pools within grassland areas.

Deserts and Shrublands Deserts and dry shrublands are to be found across central and southern Western Australia, central and southern Northern Territory, most of Southern Australia except for the coastal south, north-western New South Wales, and south-western Queensland. Named deserts cover roughly 18 per cent of the Australian mainland. However, about 35 per cent of the continent receives so little rain that it, too, is effectively desert. Vast plains of gravel and stone with a minimum of vegetation, as well as semi-arid deserts with sparse tussocks of sclerophyllous grass, chiefly spinifex (Triodia), occupy the lowlands of the interior central-west. After the sporadic rains, some areas explode into life but it is, in general, of short duration. Of all Australian habitats it is the deserts that have been the least modified by man, but this does not mean that desert animals are not endangered. Many desert vertebrates have become extinct or are extremely rare. The Lake Mackay hare-wallaby (Lagorchestes asomatus) is known only from a single specimen collected in 1932 between Mount Farewell and Lake Mackay in central-western Northern Territory. It became extinct sometime during the midtwentieth century. The lesser bilby or yallara (Macrotis leucura) was a type of medium-sized marsupial that inhabited the central deserts of Western Australia, Northern Territory, South Australia, and south-western Queensland. The last specimen was collected near Koonchera Dune in north-eastern South Australia in 1931, although a skull of unknown age was found south-east of Alice Springs, near the edge of the Simpson Desert, in 1967. Aboriginal records indicate that populations may have survived into the 1960s. Another species, the greater bilby (M. lagotis), formerly occurred over 70 per cent of the arid and semi-arid mainland of Australia south of 18°S latitude. It disappeared from the Flinders Ranges in the 1920s, from northern South Australia in the 1930s, and from New South Wales and southern South Australia by the 1950s at the latest.

It declined dramatically in south-western Australia in the 1920 and 1930s, although it may have persisted there until the 1980s. Today it occurs patchily in north-western Australia, with wild subpopulations now restricted predominantly to the Tanami Desert (Northern Territory), and in the Gibson, Little Sandy, and Great Sandy deserts and parts of the Pilbara (Western Australia). There is an additional, isolated pocket in central Australia (south-western Queensland). There is a large captive population, and the species has been reintroduced in recent years into suitably predator-free protected areas on the mainland as well as on Thistle Island, South Australia. The desert rat-kangaroo (Caloprymnus campestris) originally occurred in north-eastern South Australia and adjacent south-eastern Queensland. The last confirmed record came in 1935 from near Ooroowilanie, east of Lake Eyre. There were unconfirmed sightings between 1957 and 2011, but the species is generally considered to be extinct. The Nullarbor dwarf bettong (Bettongia pusilla) is known only from subfossil material originating from caves within the Nullarbor Plain in south-western South Australia, but is believed to have survived into historic times. The desert bettong (B. anhydra) is known from a single modern specimen (a damaged skull) collected in 1933 from a freshly dead animal east of Lake Mackay within the Tanami Desert, in centraleastern Western Australia. Subfossil material is known from Stegamite Cave in south-eastern Western Australia. As these localities are widely separated the species is presumed to have once had a relatively large distribution. The brush-tailed marsupial rat or kowari (Dasyuroides byrnei) is a small carnivorous marsupial found in the stony desert areas of the Lake Eyre basin. Two subspecies are recognized. The northern brush-tailed marsupial rat (D. b. byrnei) is found in south-western Queensland, having been extirpated from adjacent Northern Territory. The southern brush-tailed marsupial rat (D. b. pallidior) is confined to a small area of north-eastern South Australia. Both have suffered declines in areas where livestock grazing is intense, in particular near waterholes used by stock, and are additionally threatened by feral cat predation.

Figure 7.3 Desert rat-kangaroo (Caloprymnus campestris). (Credit: Hedley Herbert Findlayson.)

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The lesser stick-nest rat (Leporillus apicalis) historically occurred in great numbers across central Australia, with many early explorers commenting on the abundance and large size of stick-nests. By the early twentieth century, however, it was already considered rare, due mainly to predation by feral cats. There have been no confirmed reports since 1933 and the species is probably extinct, although it is possible that it may persist in small numbers in remote areas. The plains mouse (Pseudomys australis) is a small rodent formerly widespread within the arid and semi-arid regions of Australia, extending to coastal or near-coastal South Australia, the Nullarbor Plain area of south-eastern Western Australia, western New South Wales, central and south-western Queensland, and probably north-western Victoria. Habitat degradation due to grazing, introduced predators and drought have all contributed to a significant decline. The species has not been recorded from New South Wales since 1936, or from Western Australia since 1969. In Queensland it has only been reported once since 1936, when remains were discovered in owl pellets in Diamantina National Park, and in far northeastern South Australia the only specimen reported since 1969 was in 1998 in the Simpson Desert. All told it now appears to be largely confined to the western gibber plains of the Lake Eyre Basin in south-central Australia, where populations are highly fragmented. The short-tailed hopping mouse (Notomys amplus) was, to judge by subfossil material, at one time found across northeastern South Australia, south-eastern Northern Territory, and parts of Western Australia. Only two complete specimens were ever collected, however, both from a small area around Charlotte Waters, near Alice Springs in Northern Territory. Last recorded in 1896, it is thought to have been driven extinct by habitat destruction and introduced predators. The longtailed hopping mouse (N. longicaudatus) is known only from a handful of specimens, the first of which were collected by English naturalist and explorer John Gilbert in 1843 in Northern Territory and Western Australia. Others were later collected in north-western New South Wales, indicating that the species historically had a wide distribution within the arid interior of Australia. It was last recorded in 1901–02 and presumed to have become extinct within a few decades thereafter, although a skull fragment found in an owl pellet in 1977 indicates that it possibly survived much longer in remote areas. Old records indicate that a surviving species, the dusky hopping mouse (N. fuscus), also occupied a large area of central Australia at one time, including parts of Queensland, New South Wales, Victoria, Northern Territory, and South Australia. It now appears to be confined to a small number of localities on the South Australia–Queensland border area, and perhaps in some remote areas of Northern Territory. The Eyrean grasswren (Amytornis goyderi) is an uncommon and cryptic species endemic to the arid regions of Central Australia. It had not been observed since its discovery in 1874 when a pair was at last found in 1931, near Lake Eyre. During the 1950s it was considered extinct until again

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rediscovered in 1962. Two more were collected in Queensland in 1976, and a follow-up expedition revealed a small population. In recent years it has come to be reliably known from the dune fields of the Simpson and Strzelecki deserts, where it favours areas of sandhill canegrass. Despite its small population and highly restricted habitat it is not thought to be presently at risk for extinction. The great desert skink (Liopholis kintorei) is found patchily across a vast area of west-central Australia (north-eastern Western Australia, north-western South Australia, and southwestern Northern Territory). It is threatened mainly by loss of habitat and introduced predators.

Isolated Caves, Springs, and Pools Australia has a considerable number of cave systems throughout the continent, many of which remain undiscovered or unexplored. The cave-dwelling frog (Litoria cavernicola) is known only from caves and rocky areas in the sandstone gorges flanking the Mitchell Plateau, in the Kimberly region of north-western Australia (northern Western Australia). The blind gudgeon (Milyeringa veritas) and the blind cave eel (Ophisternon candidum) are confined to subterranean waters in the caves of the Cape Range, north-western Western Australia.

The Great Artesian Basin The Great Artesian Basin is the largest and deepest in the world. Located in north-eastern Australia, it underlies most of Queensland, south-eastern Northern Territory, northeastern South Australia, and northern New South Wales. In addition to providing the only source of freshwater for much of the inland parts of the continent, its above-ground, mostly seasonal manifestations are vitally important for wildlife, in particular waterfowl.

The Dalhousie Springs The Dalhousie Springs are a group of over 60 natural artesian springs located on the western fringe of the Simpson Desert in northern South Australia. The water, which is highly mineralized, ranges in temperature from 38 to 43°C. Drilling has considerably reduced the flow rates over the past century. The Dalhousie hardyhead (Craterocephalus dalhousiensis) and Glover’s hardyhead (C. gloveri) are both confined to the Dalhousie Springs. The Dalhousie mogurnda (Mogurnda thermophila) is confined to the Dalhousie Springs. The Dalhousie goby (Chlamydogobius gloveri) is confined to the Dalhousie Springs.

The Elizabeth Springs The Elizabeth Springs are located in western Queensland. The Elizabeth Springs goby (Chlamydogobius micropterus) is confined to the marshy pools of Elizabeth Springs.

Lakes, Rivers, and Marshes

The Edgbaston Springs The Edgbaston Spings are a complex of small, shallow springs located on a former sheep and cattle property in centralwestern Queensland. They and the surrounding area are now protected by the Edgbaston Reserve. The Edgbaston goby (Chlamydogobius squamigenus) is confined to the Edgbaston Springs. The red-finned blue-eye (Scaturiginichthys vermeilipinnis) is confined to the Edgbaston Springs.

Lakes, Rivers, and Marshes As a consequence of its very dry climate Australia has a poorly developed system of rivers, and indeed has the fewest number and the least run-off of any continent after Antarctica. Moreover, relatively few of its watercourses are permanent, with lakes and marshes varying in size in relation to the periods of rain and drought. Occasionally even large rivers may dry up entirely and their beds remain waterless for several years. A large proportion of rivers fade out before they even reach the sea, and erosion has led to the silting-up of waterholes and billabongs along inland rivers. Such conditions make it difficult for aquatic animals to survive. However, many species, particularly fishes, have solved the problem by developing an ability to aestivate for prolonged periods, as for instance the Queensland lungfish (Neoceratodus forsteri), a ‘living fossil’ which has managed to exist for millions of years. Thus, these special climatic challenges combined with the long isolation of Australia have made its freshwater fishes unique in the same evolutionary sense as the eucalypts and marsupials. Many of them are now threatened, not only by man-made changes in their freshwater habitats but also by the introduction of numerous exotic fish species, which prey on or compete with the Australian ones. The platypus (Ornithorhynchus anatinus) is a unique egglaying mammal that lives in long burrows and forages for food on the bottom of rivers and streams with its sensitive, ducklike bill. During the nineteenth century it was extensively hunted for pelts, which was used in the making of rugs, and for a time seriously threatened. Long protected by law it is now once again widespread in eastern Australia from Queensland to Victoria, as well as on Tasmania and King Island. An additional, introduced population occurs on Kangaroo Island. The Australian freshwater crocodile (Crocodylus johnsoni) is a relatively small crocodilian from northern Australia that, for much of the twentieth century, was in danger of extinction due to overhunting for its skin. With better protection it has made a comeback, only to see its numbers start to decline once again due to ingestion of the poisonous, invasive cane toad (Rhinella marina). It is not currently considered to be threatened with extinction. The western swamp turtle (Pseudemydura umbrina) is perhaps Australia’s rarest reptile. First described in 1839 from a specimen collected on the Swan Coastal Plain in Western Australia, it was not seen again for more than a century.

Intensive clearing and drainage of the swamps north and south of Perth had already destroyed much of the available habitat of the species, and it had long been considered extinct when another individual was at last recorded in 1953. Fortunately, two small reserves were quickly established in the swamps where it had been found. Subsequent investigation revealed a population of around 300 turtles. Mitchell’s water monitor (Varanus mitchelli) is confined to wetland areas throughout the northern parts of Western Australia, Northern Territory, and possibly far north-western Queensland. It is not known to occur on any offshore islands. Mertens’ water monitor (V. mertensii) is found in coastal and inland waters across much of northern Australia, from the Kimberley region of Western Australia, across the Top End of Northern Territory to the western side of the Cape York Peninsula in northern Queensland. Both are seriously threatened by invasive cane toads (Rhinella marina), through poisoning after eating them. The ornamental snake (Denisonia maculata) is a small, venomous species confined to ephemeral wetland areas of central-eastern Queensland. The Magela hardyhead (Craterocephalus marianae) is confined to a small area of northern-flowing creeks and rivers on the Kakadu escarpment of Northern Territory. The eastern freshwater cod (Maccullochella ikei) is a large, predatory fish that was historically fairly widely distributed within the Clarence, Richmond, and possibly Brisbane rivers of coastal north-eastern New South Wales and south-eastern Queensland. By the 1930s, naturally occuring populations were probably near to extirpation within the Richmond and Brisbane rivers due to overfishing and habitat destruction, and by the 1970s the species was entirely confined to a small number of isolated tributaries within the Clarence River. Since then, legal protection and restocking with hatchery-bred individuals has resulted in a general increase in both abundance and distribution within the latter drainage, along with the reintroduction of several fragmented populations within the Richmond River. The variegated pygmy perch (Nannoperca variegata) is known from the Ewens Ponds and Deep Creek, south-eastern South Australia, as well as from a few tributaries of the Glenelg River in south-western Victoria. The Yarra pygmy perch (N. obscura) is found patchily in the coastal drainages of southeastern Australia (south-eastern South Australia and Victoria). The Oxleyan pygmy perch (N. oxleyana) is confined to coastal eastern Australia (south-eastern Queensland and north-eastern New South Wales), where it lives in dune lakes, ponds, creeks, and swamps. All are threatened by habitat destruction and degradation and from introduced fish species. The Cairns rainbowfish (Cairnsichthys rhombosomoides) is confined to fragmented populations inhabiting shallow, fastflowing streams in north-eastern Queensland. The Eacham rainbowfish (Melanotaenia eachamensis) was long thought to have been endemic to Lake Eacham in northeastern coastal Queensland, where it was extirpated by introduced predatory fish. It has since been rediscovered in the

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private collections of aquarists, and subsequently found to have a wider distribution within the Barron and Johnstone river systems. The dwarf galaxias (Galaxiella pusilla) is confined to freshwater coastal wetlands in south-eastern Australia, specifically southern Victoria and Tasmania. The honey blue-eye (Pseudomugil mellis) is confined to two disjunct areas in coastal central and south-eastern Queensland. The Australian brook lamprey (Mordacia praecox) is confined to a few isolated coastal rivers in southern Queensland, New South Wales, and possibly eastern Victoria.

Lake Eyre Lake Eyre is located in northern South Australia. A shallow, endorheic lake; on the rare occasions when it fills during the rainy season it is the largest in Australia. It contains the lowest natural point in Australia.

Cooper Creek Cooper Creek is part of the Lake Eyre drainage system. The Cooper Creek turtle (Emydura macqauarii emmotti) is confined to the Cooper Creek drainage. The Cooper Creek catfish (Neosiluroides cooperensis) is confined to the Cooper Creek drainage.

The Fitzroy River The Fitzroy River is located in north-eastern Australia (southcentral and south-eastern Queensland). The Fitzroy River turtle (Rheodytes leukops) is confined to the Fitzroy River. The leathery grunter (Scortum hillii) is a perch-like fish found patchily in streams and pools within the Fitzroy River drainage, where it is threatened by loss of habitat and pollution from mining operations.

The Mary River The Mary River is located in south-eastern Queensland. The Mary River turtle (Elusor macrurus) is confined to the Mary River drainage. The Mary River cod (Maccullochella mariensis) is a large, predatory fish confined to the Mary River drainage, where it is threatened by habitat destruction and overfishing.

The Murray–Darling River Drainage Located in central-eastern and south-eastern Australia, the Murray River is Australia’s longest. It rises on the western side of the Australian Alps and meanders some 2500 km through South Australia, Victoria, New South Wales, and southern Queensland. When combined with its major tributary the Darling River the total course extends to 3750 km. Bell’s sawshelled turtle (Elseya bellii) is confined to the upper reaches of the Namoi, Gwydir, Macdonald, and Severn rivers in northern New South Wales and southern Queensland.

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The trout cod (Maccullochella macquariensis) is a large, predatory species that was historically widespread and common within the Murray–Darling drainage, but has been much reduced by overfishing, habitat degradation and invasive trout species. Only a single wild, naturally occuring population remains, although there are number of self-sustaining introduced ones. The Macquarie perch (Macquaria australasica) is still found widely but patchily within the Murray–Darling drainage, but remains threatened by overfishing, siltation and dam construction, and disease. The Murray hardyhead (Craterocephalus fluviatilis) was historically widespread and abundant within the middle and lower Murray River drainage, but has undergone a considerable decline due to habitat destruction and degradation and by introduced fish species. It is now confined to a few scattered localities. The flathead galaxias (Galaxias rostratus) is currently known only from the southern Murray–Darling drainage in Victoria, where it is found patchily.

The Goulburn River The Goulburn River is a perennial river located in eastcentral Victoria. The barred galaxias (Galaxias fuscus) is confined to the Goulburn River, where it is seriously threatened by habitat degradation and introduced fish species.

Miscellaneous Lakes, Rivers, and Marshes The Burdekin River is located in north-eastern coastal Queensland. The small-headed grunter (Scortum parviceps) is a perch-like species known only from the upper Burdekin River. The Manning River is located in north-eastern New South Wales. The Manning River sawshelled turtle (Elseya purvisi) is confined to the Manning River drainage. The Bellinger River is located in north-eastern New South Wales. The Bellinger River sawshelled turtle (Elseya georgesi) is confined to the Bellinger River drainage.

Coasts and Satellite Islands The primary threat to Australian coastal areas is development coupled with industrial pollution and the potential for oil spills. It is tragic that the eastern coast of Australia and the offshore islands, the richest in plants and animals, have been the most affected, although in recent years all islands and coasts have become vulnerable to exploitation and degradation. The Australian sea lion (Neophoca cinerea) is confined to south-western coastal Australia, from The Pages (an island group just east of Kangaroo Island) to Houtman Abrolhos on the western coast of Western Australia. Aborigines hunted the seals for subsistence purposes for thousands of years, and early European colonists also relied on them for food and other products. But it was not until sealers began to harvest

Coasts and Satellite Islands

them on a large scale during the seventeenth and eighteenth centuries that populations began to be seriously affected, eventually extirpating the species from areas around the Bass Strait and Tasmania. They are now protected by a variety of laws and no longer hunted, but have never quite recovered fully in numbers or reoccupied all of their former range. The Barrow Island golden bandicoot (Isoodon auratus barrowensis) is confined to Augustus, Barrow, and Middle islands off the north-western coast of Australia (Western Australia). Butler’s dunnart (Sminthopsis butleri) is a mouse-like marsupial known only from a small area of coastal northern Australia (north-eastern Western Australia) and from Bathurst and Melville islands (Northern Territory). The water mouse (Xeromys myoides) is a little-known species found disjunctly in coastal regions of southern and eastern Australia (Northern Territory and Queensland) and southern New Guinea. It is threatened by loss of habitat. The long-eared mouse (Pseudomys auritus), which lived along the edges of dry salt-water lagoons in South Australia, seems to have been exterminated more than 150 years ago. The New Holland mouse (P. novaehollandiae) was long known only from four specimens, the last of which was collected prior to 1887. Long considered extinct, in 1967 a fifth individual was found in the Ku-ring-gai Chase National Park north of Sydney. Other fragmented populations have since been found across south-eastern Australia in New South Wales, Queensland, Victoria, and Tasmania. It is threatened by the development of its coastal dune habitat. The northern hopping mouse (Notomys aquilo) is known only from coastal Northern Territory (where the species has not been recorded in decades) and the island of Groote Eylandt. The Providence petrel (Pterodroma solandri) is a small seabird that is widespread and common throughout much of the Tasman Sea and adjacent areas of the Pacific, but nests only on two mountaintops on Lord Howe Island and, marginally, on Phillip Island, a tiny islet off Norfolk Island. It historically bred on Norfolk Island as well, but was extirpated there by massive overharvesting for food during the late eighteenth century. The Australian Gould’s petrel (P. leucoptera leucoptera) ranges widely across both the Pacific and Southern oceans, but breeds only on a few small islands off the coast of New South Wales, primarily Cabbage Tree Island. The hooded plover (Thinornis cucullatus) is a type of shorebird divided into two subspecies. The eastern hooded plover (T. c. cucullatus) is found in southern and south-eastern Australia (New South Wales, South Australia, Tasmania, and a few nearby islands). The western hooded plover (T. c. tregellasi) occurs in south-western Australia (Western Australia). Both are threatened by the crushing and trampling of its nests and eggs by off-road vehicles and people as well as habitat destruction and degradation. The Australian fairy tern (Sternula nereis nereis) breeds only in a few widely scattered colonies in western and southern coastal Australia, which are vulnerable to human disturbance

Guenther’s southern gecko (Christinus guentheri) is confined to Lord Howe Island, Norfolk Island, and a few smaller islands. The Lord Howe skink (Oligosoma lichenigera) is confined to Lord Howe Island, Norfolk Island, and a few smaller islands. The Hermite worm lizard (Aprasia rostrata) is confined to the Montebellow Islands and Barrow Island off north-western Western Australia, and to a small area of the adjacent coastal mainland. The Cooloola frog (Litoria cooloolensis) is known only from wetland areas on Fraser Island and North Stradbroke Island, off the coast of south-eastern Queensland. Freycinet’s frog (L. freycineti) and the Olongburra frog (L. olongburensis) are both found patchily in coastal eastern Australia (southeastern Queensland and parts of New South Wales, including offshore islands), where they are threatened by loss of habitat. The wallum froglet (Crinia tinnula) is found patchily in coastal dunes and swampy areas from Litabella National Park in south-eastern Queensland to central-eastern New South Wales, as well as on a few offshore islands (Fraser, Bribie, Moreton, and North Stradbroke).

Tasmania Tasmania is a large island located 240 km south of southeastern Australia, directly in the pathway of the notorious ‘Roaring Forties’ wind that encircles the southern part of the globe. It is very mountainous, with the central highlands covering most of the central-west and moors occurring above the timber line. Much is still densely forested, with the western areas containing some of the largest remaining temperate rainforests in the Southern Hemisphere. Connected with the continent in geologically recent time (probably about 10,000 years ago during the last world-wide lowering of sea level), it has harboured some animals after their disappearance from continental Australia, and vice versa. The island is thought to have been occupied by aboriginals for 30,000 years prior to their being wiped out by British colonizers. The AngloTasmanians were no more merciful to animal life although, to their credit, they also protected certain species that faced extinction on the Australian mainland, among them the eastern quoll (Dasyurus viverrinus), platypus (Ornithorhynchus anatinus), swamp antechinus (Antechinus minimus), southern brown bandicoot (Isoodon obesulus), and Tasmanian pademelon (Thylogale billardierii). More recently Tasmania became the founding place of the first environmental political party in the world, and more than 40 per cent of the island has been set aside as protected areas. One of the most regrettable instances of the disappearance of an animal on the Australian mainland was that of the Tasmanian tiger or thylacine (Thylacinus cynocephalus), the largest of the marsupial carnivores. Europeans did not have a hand in the retreat of this predator from continental Australia, but they were certainly responsible for exterminating it in Tasmania. It has often been argued that the mainland thylacine

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succumbed during the Pleistocene as a result of competition from the dingo. This seems unlikely because there was sufficient prey for both species, and the dingo does not seem to have been numerous. It also fails to explain the species’ disappearance from New Guinea. In Tasmania, however, man alone was responsible for the tremendous decrease of the thylacine. When it was still fairly common in many areas it was accused of damaging livestock, and the authorities declared war against it. Bounties were paid beginning in 1888, and up to 1914 at least 2268 of the animals were reported killed, although this official figure is much below the true total. About 1910 a sharp decline, thought to have been caused by disease, set in. Of course, destruction of habitats also played its part and the result was the almost complete extinction of the species. The last known individual died in the Hobart Zoo in 1936, the same year that it finally received official protection under Tasmanian law. Since then there have been numerous reports of its continued existence, not just in Tasmania but on the Australian mainland as well. It is claimed that one was killed in 1961, and in 1966 footprints and even fresh hair from a lair were reported to have been found in north-western Tasmania. Yet, despite several organized searches there has been no irrefutable evidence of its survival, and it must be considered extinct. The Tasmanian devil (Sarcophilus harrisii) became, with the extinction of the Tasmanian tiger, the largest carnivorous marsupial in the world. It too historically inhabited the Australian mainland but was extirpated there most likely owing to competition with dingoes, and was afterwards confined to Tasmania. There it was heavily hunted both for its fur and for its perceived attacks upon livestock, but managed to hold on in remote areas as well as on tiny Maria Island off the eastern coast. Unfortunately, since the 1990s the population has crashed owing to an invariably fatal infectous cancer (devil facial tumour disease) that now threatens the species’ survival

in the wild. Collisions with motor vehicles also take a considerable toll in some areas, particularly when the animals are eating roadkill. In 2020 the species was successfully reintroduced to a protected area in New South Wales. It is the first time that it has lived on the Australian mainland in over 3000 years. The Tasmanian emu (Dromaius novaehollandiae diemenensis) was delibately exterminated by European settlers sometime before 1850. Interestingly, the latter would repeat the process when the mainland emu was introduced later that century, as they considered the birds to be a pest. The forty-spotted pardalote (Pardalotus quadragintus) is a type of wren that has long been one of Australia’s rarest birds. It is only reliably found in a few isolated colonies in southeastern Tasmania and offshore islands (most notably, Maria Island and Bruny Island).

Lakes, Rivers, and Marshes With its rugged topography, Tasmania has a great number of rivers and estuaries and consequently a highly endemic fish fauna. Unfortunately, its lakes are among the most polluted in the world. Several small, tubular-shaped fish of the genus Galaxias endemic to Tasmania are threatened by loss of habitat due to dam construction and by introduced species. The Pedder galaxias (G. pedderensis) was historically found in Lake Pedder, Lake Maria, and their tributaries and surrounding swamps in southern Tasmania. The species declined rapidly from the late 1970s due to the flooding of Lake Pedder by a hydroelectric project, and by the introduction of predatory brown trout throughout its range. It has not been recorded within its native range after 1996, and was for a time confined to a single, remote highland lake in south-western Tasmania (Lake Oberon), where it had been previously introduced as a safeguard. A second translocated population has since been Figure 7.4 The last-known Tasmanian tiger, ‘Benjamin’, photographed at the Beaumaris Zoo, Hobart, during the 1930s. (Credit: Hobart Zoo.)

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Coasts and Satellite Islands

established. The small Pedder galaxias (G. parvus) is found in swampy areas and suitable streams surrounding Lake Pedder in southern Tasmania, along with a few streams draining to Lake Gordon and within the Huon River catchment. It was extirpated from much of its former range by the flooding of Lake Pedder in the early 1970s. The saddled galaxias (G. tanycephalus) occurs in four lakes and a connecting stream in east-central Tasmania. The Swan galaxias (G. fontanus) occurs in a few fragmented localities within the trout-free headwaters of the Swan and Macquarie rivers in north-eastern Tasmania. The species has also been successfully introduced into other predator-free streams. The Arthurs Lake paragalaxias (Paragalaxias mesotes) occurs in Arthurs Lake, Lake Woods, and a small area of Lake River in the Central Highlands of Tasmania. It is threatened by fluctuating water levels and by trout predation. Great Lake is located on Tasmania’s Central Plateau. The Great Lake paragalaxias (Paragalaxias dissimilis) is confined to Great Lake and two artificial impoundments downstream from it (Shannon Lagoon and Penstock Lagoon). The Derwent River is located in central and southeastern Tasmania. The Clarence galaxias (Galaxias johnstoni) is confined to a few isolated localities within the upper Derwent River drainage, where it is threatened by introduced fish species.

Pedra Branca Rock Pedra Branca is a windswept rocky islet located off the southern coast of Tasmania. The Pedra Branca skink (Carinascincus palfreymani) is confined to Pedra Branca Rock, where it is dependent upon seabird colonies. The total population is thought to be between 300 and 500.

Kangaroo Island Located south-west of Adelaide, Kangaroo Island is Australia’s third largest. Once occupied by Aborigines, the native population disappeared after the land became an island following rising sea levels several thousand years ago. It was subsequently resettled by Europeans from the early nineteenth century onwards. Although fully half of the island has been cleared of vegetation, about a quarter of it has been set aside to protect the remnants. In addition to the native species koala (Phascolarctos cinereus), platypus (Ornithorhynchus anatinus) and other threatened species have been introduced. The island was devastated by the massive bushfires of 2019–20. The Kangaroo Island dunnart (Sminthopsis aitkeni) is a small, carnivorous marsupial known only from Flinders Chase National Park on the western end of the island. The Kangaroo Island emu (Dromaius baudinianus) was last collected in 1802. It is thought to have been systematically exterminated by hunting pressure prior to the arrival of permanent settlers in 1836.

The Kangaroo Island glossy black cockatoo (Calyptorhynchus lathami halmaturinus) is an endemic subspecies with a total population estimated as low as 100. The Kangaroo Island whipbird (Psophodes leucogaster lashmari) is confined to Kangaroo Island, where its population is stable at around 2000.

Flinders Island Flinders Island is located in Bass Strait and is nearer to Tasmania than to continental Australia. About a third of the island is mountainous, and the coastal areas are dominated by scrubby areas and sandy deposits that often take the shape of dunes. Higher elevations still feature eucalyptus forest. The Flinders Island wombat (Vombatus ursinus ursinus) formerly occurred on all the islands in the Bass Strait, but has since disappeared as a result of being hunted for food as well as changes of habitat. It is now confined to Flinders Island, where it was said to be very common during the early nineteenth century.

King Island King Island is located in the Bass Strait about halfway between Tasmania and the mainland state of Victoria. The King Island emu (Dromaius minor) was historically endemic to King Island. Although numerous bones have been found the only existing skin was collected by Nicolas Baudin in 1802, shortly before the species became extinct. It is thought that hunting by sealers for food was the cause.

Lord Howe Island Located in the Tasman Sea between Australia and New Zealand, Lord Howe Island is a roughly crescent-shaped volcanic remnant. The island has undergone a tremendous environmental depletion since Europeans came there. In addition to those species and subspecies wiped out by man and his domestic animals, many others were killed between 1918 and 1928 by rats that came ashore from a wrecked ship. The Lord Howe long-eared bat (Nyctophilus howensis) is known only from a single incomplete skull found in 1972 and not found despite extensive surveys since. However, local island people continue to report seeing it. The Lord Howe hawk-owl (Ninox novaeseelandiae albaria) appears to have gone extinct through predation by, or competition with, the Tasmanian masked owls (Tyto novaehollandiae castanops) which were introduced in the 1920s in a failed attempt to control the rat population. The Lord Howe white swamphen (Porphyrio albus) was not definitely recorded after 1790, and was extinct due to hunting by the time the island was settled in 1834. The Lord Howe woodhen (Hypotaenidia sylvestris) is a flightless endemic rail that was once highly threatened by human persecution and predation by pigs and rats. By the 1970s the total population was down to less than 30.

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Figure 7.5 Illustration (probably based on a live specimen) of a Lord Howe white swamphen. (Credit: Arthur Phillip.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

However, between 1978 and 1984 feral animals were removed and captive-raised birds were successfully reintroduced into the wild. The species is now considered to be fairly safe. The Lord Howe red-fronted parakeet (Cyanoramphus subflavescens) was last reported in 1869. The Lord Howe white-throated pigeon (Columba vitiensis godmanae) has been extinct since the mid-nineteenth century. The Lord Howe thrush (Turdus poliocephalus vinitinctus) became extinct in 1918. The Tasman starling (Aplonis fusca) historically occurred on both Lord Howe and Norfolk Islands, where it was divided into two subspecies. The Lord Howe starling (A. f. hullianus) became extinct in 1918, most likely due to the arrival of black rats on the island. The Lord Howe white-eye (Zosterops strenuus) was a type of passerine bird last recorded in 1908, and not found during a survey in 1928. The Lord Howe gerygone (Gerygone insularis) was a wrenlike bird last recorded in 1928, with none being found during a survey of the island in 1936. The Lord Howe grey fantail (Rhipidura fuliginosa cervina) was a type of passerine bird that became extinct in 1928.

Norfolk Island Norfolk Island is a remote volcanic island about 1400 km east of Australia and midway between New Zealand and New Caledonia. The island was first settled by Polynesians but was long unpopulated when the British arrived in 1788. Since then it has been greatly affected by human pressures and a number of species and subspecies are now extinct. The Norfolk Island hawk-owl (Ninox novaeseelandiae undulata) had, by 1986, been reduced to a single female. In a programme to preserve at least some of her genes two male

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owls of the related New Zealand subspecies (N. n. novaeseelandiae) were released on to the island as potential mates for her. The attempt was successful and resulted in fledged chicks in 1989 and 1990. The original female disappeared in 1996, but by then there was a small hybrid population of around a dozen birds, which persists to this day. The Norfolk Island kaka (Nestor productus) was a type of parrot endemic to Norfolk and adjacent Philip Island. It went extinct in the mid- to late nineteenth century, with the last known living bird in captivity in London in 1851. The Norfolk Island parakeet (Cyanoramphus cookii) is confined to the region of Norfolk Island National Park. The Norfolk Island ground dove (Pampusana norfolkensis) is known only from a single painting. It is thought to have succumbed to introduced predators at the end of the eighteenth century. The Norfolk Island pigeon (Hemiphaga novaeseelandiae spadicea) was last seen in 1901. The Norfolk Island starling (Aplonis fusca) was last recorded in 1923, and was certainly extinct before 1968. The Norfolk Island thrush (Turdus poliocephalus poliocephalus) went extinct during the late 1970s, with the last confirmed record occurring in 1975. The Norfolk Island robin (Petroica multicolor) is confined to the vicinity of Norfolk Island National Park. The white-chested white-eye (Zosterops albogularis) is a type of passerine bird that is now possibly extinct, although a small population may still survive within Norfolk Island National Park. The Norfolk Island golden whistler (Pachycephala pectoralis xanthoprocta) is a type of passerine bird confined to the vicinity of Norfolk Island National Park.

Miscellaneous Islands The Percy Islands are a group of small islands located off north-eastern Queensland. The Percy Islands flying fox (Pteropus brunneus) is known only from a single specimen collected in 1874 from an unspecified island in the group, along with subsequent reports of a colony towards the end of the nineteenth century. Pearson Island is located off the coast of South Australia. It and nearby islands in the group have been protected since the 1960s. The Pearson black-flanked rock-wallaby (Petrogale lateralis pearsoni) is confined to Pearson Island. The Abrolhos Islands are a large group of islets and associated coral reefs located off the coast of southeastern Western Australia. The Abrolhos painted buttonquail (Turnix varius scintillans) is confined to a few islets within the Abrolhos Islands, where it is threatened by introduced rats and feral cats. The Recherche Archipelago is an island group located of the southern coast of Western Australia. Part of it is protected within the Recherche Archipelago Nature Reserve. The Recherche black-flanked rock-wallaby (Petrogale lateralis hacketti) is confined to the Recherche Archipelago.

Balance for the Australian Realm

Lancelin Island is a small, low-elevation island located off the south-western coast of Western Australia. The Lancelin skink (Ctenotus lancelini) is confined to the island, where it is rare and decreasing. Dirk Hartog Island is located off the central-western coast of Western Australia. The Dirk Hartog white-winged fairy wren (Malurus leucopterus leucopterus) is restricted to the island. Barrow Island is located off the Pilbara coast of Western Australia. The Barrow wallaroo (Macropus robustus isabellinus) and the Barrow white-winged fairy wren (Malurus leucopterus edouardi) are both restricted to Barrow Island.

Balance for the Australian Realm Compared with Africa and Asia prehistoric modern humans entered Australia relatively late, about 65,000–50,000 years ago, arriving by temporary land-bridges and short sea crossings from what is now South East Asia. The oldest human remains yet found there have been dated to around 41,000 years ago, and are believed to be the ancestors of modern indigenous Australians. These first inhabitants were the ancestors of modern Aborigines. The latter people developed a hunter-gatherer way of life and generally had little effect on the living nature around them apart from the grass fires that they periodically lit to catch lizards (a practice that still continues, and which can have devastating environmental effects). It was later immigrants – seafarers from New Guinea – about 5000 years ago who are thought to have introduced the dingo (Canis lupus dingo), the feral Australian dog. Until European settlers arrived with their domesticated dogs, all dingos were purebreds, living, breeding, and undergoing natural section in the wild. The animals had a devastating effect on many native species of fauna, in the long run much more serious than the grass fires mentioned above. Early European exploration of Australia was driven primarily by the Dutch. The western and northern coastlines had been known and charted by them since at least the beginning of the seventeenth century, and within a few decades the southern coastline too would be explored, culminating with the discovery of ‘Van Dieman’s Land’ (Tasmania) by Abel Tasman in 1642–43. Brief contact with the indigenous people had also been made, although no attempt at colonizing what they called ‘New Holland’ had been made. Nevertheless, a number of shipwrecks had left men either stranded or, as in the case of the Batavia in 1629, marooned for mutiny or murder, thus becoming the first Europeans to permanently inhabit the island continent. But it was not until later in the following century, when Captain James Cook’s Endeavour sailed along and mapped the eastern coast, that the European era in Australia truly began. The latter claimed this stretch of coast for Great Britain and christened it New South Wales. It was there, on 29 April 1770, that he would anchor within a wide bay about 16 km south of Sydney. Onboard were two naturalists, an Englishman named Joseph Banks and a Swede,

Daniel Solander, a disciple of Linnaeus. They made so many discoveries during the week they stayed there, and became so enthusiastic over all the new plants and trees they found, that the place was given the name Botany Bay. Not much of the wildlife, either plants or animals, that Banks and Solander found there two and a half centuries ago remains today. Cook and his men made several remarkable zoological discoveries during their visit, with Cook himself describing what was clearly a small kangaroo species. He is therefore often credited with ‘discovering’ the kangaroo (Dutch sailors had, however, found and described kangaroos in western Australia at least a century and a half earlier). During his visit Cook sailed along and mapped the east coast, which he named New South Wales and claimed for Great Britain. During the 1780s the British government sent a fleet of ships, the ‘First Fleet’, to establish a new penal colony at Botany Bay, near present-day Sydney. Another British settlement was established in Van Diemen’s Land, now known as Tasmania, in 1803, which became a separate colony in 1825. The United Kingdom formally claimed the western part of western Australia (the Swan River Colony) in 1828, and additional ‘free’ (i.e. non-penal) colonies were carved from parts of New South Wales: South Australia in 1836, Victoria in 1851, Queensland in 1859, and the Northern Territory in 1911. The last convict ship to New South Wales arrived in 1848. Most early convicts were transported for petty crimes, and assigned as labourers or servants upon arrival. While the majority settled into colonial society once emancipated, convict rebellions and uprisings were also staged, although invariably suppressed under martial law. Meanwhile, exploration continued. In 1798–99 Matthew Flinders circumnavigated Tasmania, proving its insularity, and in 1802–03 circumnavigated all of the continent. Much of the interior would remain unknown for considerably longer, and it wasn’t until 1860–61 that Robert O’Hara Burke and William Willis became the first to cross Australia from south to north, travelling from what is now Melbourne to the Flinders River. The arrival of Europeans with their sheep, cattle, rabbits, and many other foreign animals and plants in their wake upset an equilibrium that had lasted for tens of thousands of years. Previously the flora and fauna, including indigenous humans and in spite of the dingo, had fused into a more or less balanced ecosystem. It is true that some mammal species had declined considerably as a result of aboriginal hunting, as evidenced by the remains found in middens and caves. But it was only during the past 250 years that humans have had a truly devastating environmental impact. The aboriginal population itself, estimated to have been between 750,000 and 1,000,000 in 1788, would also undergo a serious decline over the next century and a half, falling to just 50,000 by 1930. This was primarily due to infectious disease (mainly smallpox), although thousands more would die as a result of frontier conflict with settlers. Moreover, a public policy of ‘assimilation’ beginning in the mid-nineteenth century may have also been a factor. Australia was the worst possible continent for

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the introduction of exotic plants and animals because of the vulnerability of its unique, highly specialized flora and fauna. No less than 17 types of wild mammal have been brought into Australia since 1788, the fatal year of the introduction of the rabbit. The destructive effect of these exotics combined with overgrazing by enormous numbers of livestock and predation by domestic cats and dogs has been monumental. Sheep, together with rabbits, certainly have done more to change Australian natural habitats and damage wildlife than any other introduced species. Deforestation has also affected the fauna, as has ruthless hunting for sport and for pelts with dogs and modern firearms. Today the main threats are feral cats and megafires.

In recent historical time (i.e. since A . D . 1500), the Australian Realm has lost at least 38 species/13 subspecies of vertebrates. Among the extinct forms 25 species/3 subspecies are mammals, 10 species/10 subspecies are birds and 3 species are amphibians. Another 2 species are possibly extinct. In addition, there are 225 species/28 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 70 species/9 subspecies are mammals, 37 species/16 subspecies are birds, 27 species/3 subspecies are reptiles, 56 species are amphibians and 35 species are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

25 species

1 species

~ species

70 species

3 subspecies

~ subspecies

~ subspecies

9 subspecies

28 taxa

1 taxon

~ taxa

79 taxa

10 species

1 species

~ species

37 species

10 subspecies

~ subspecies

~ subspecies

16 subspecies

20 taxa

1 taxon

~ taxa

53 taxa

~ species

~ species

~ species

27 species

~ subspecies

~ subspecies

~ subspecies

3 subspecies

~ taxa

~ taxa

~ taxa

30 taxa

3 species

~ species

~ species

56 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

3 taxa

~ taxa

~ taxa

56 taxa

~ species

~ species

~ species

35 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

~ taxa

~ taxa

35 taxa

38 species

2 species

~ species

225 species

13 subspecies

~ subspecies

~ subspecies

28 subspecies

51 taxa

2 taxa

~ taxa

253 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

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Chapter

8

The Polynesian Realm

The Polynesian Realm comprises all the islands within the vast expanse of the North and South Pacific east of the Australian, Papua-Melanesian and Indo-Malaysian realms and west of the Americas. Alternatively known as the Oceanian Realm, it is divided into three zoogeographic regions (Polynesian, Novozelandic, and Micronesian). The climatic range is considerable, ranging from tropical to subpolar. The Polynesian Realm is geologically quite young. While other terrestrial realms include ancient continental landmasses or fragments of continents, it is composed of mostly volcanic high islands and coral atolls that arose from the sea in relatively recent times, many of them during the Pleistocene. They were created either by volcanism or as island arcs pushed upward by the collision and subduction of tectonic plates. Thus, the islands are really tall pillars rising from the depths, constantly exposed to the destructive action of rain, sun, and sea. During countless ages, moreover, myriads of small corals have built up strong walls around almost every island. Sometimes the land has disappeared from inside an encircling atoll, leaving only a lagoon in the centre. Some low islands consist entirely of coral. Plants and animals have colonized and are still doing so both volcanic and coral islands, mainly from the west. The Polynesian sheath-tailed bat (Emballonura semicaudata) as a species ranges across the islands of the western and central Pacific, where it is threatened by loss of habitat and disturbance of its roosting caves. The nominate form (E. s. semicaudata) was historically found sporadically in the Samoan, Fiji, Tongan, and Caroline islands as well as Vanuatu. It has been extirpated from the Samoan Islands and was last collected from the Tonga Islands in 1989, and may now be largely or wholly confined to the Fiji Islands, particularly Rotuma. The black-footed albatross (Phoebastria nigripes) is a large seabird that ranges throughout the North Pacific Ocean (and occasionally south of the equator as well) but nests only on a dozen islands, virtually all of which are in the isolated northwestern Hawaiian Islands. Additional small colonies are found in a few islands south of Japan and (at least formerly) off the coast of Mexico. At one time heavily hunted, the species has recovered much of its former range and numbers but remains vulnerable to incidental capture from commercial fisheries. The Laysan albatross (Diomedea immutabilis) is a gulllike species that ranges across the North Pacific from Japan

to the Bering Sea and south to Mexico. It is currently confined to just 16 nesting sites, however, the vast majority of which are in the north-western Hawaiian Islands where during the early nineteenth century hundreds of thousands of birds were slaughtered by Japanese feather hunters. The important Wake Island and Johnston Atoll colonies were completely wiped out before protections were finally put in place. While once again common, the species has yet to fully reclaim its former range and numbers, and remains vulnerable to feral cat predation, longline fisheries, and plastic pollution. The Phoenix petrel (Pterodroma alba) is a type of seabird found throughout the central Pacific, where their breeding colonies are nevertheless highly threatened by habitat destruction, human exploitation, and predation by invasive species. The Polynesian storm petrel (Nesofregetta fuliginosa) breeds on a number of islands across the central Pacific but is highly vulnerable to introduced predators and other threats. Many of its former colonies have become extirpated. Bryan’s shearwater (Puffinus bryani) is a rare and littleknown type of seabird first described from a single individual collected in 1963 from Midway Island in the north-western Hawaiian Islands. It was not reported again until the early 1990s, when a few more were found in nesting burrows there, and since 2015 has also been found in the Bonin Islands of Japan. Its at-sea distribution remains unknown.

The Micronesian Region The Micronesian Region is comprised of thousands of small islands scattered across the western Pacific Ocean, north of the equator and west of the International Date Line. Major groups include the Mariana Islands in the north-west, the Caroline Islands in the centre, the Marshall Islands to the west, and the islands of Kiribati in the south-west. Geologically these islands are mainly coral atolls and the climate is entirely tropical with very little seasonal variation. Fauna generally consists of bats, birds, and a few reptiles. The Mariana flying fox (Pteropus mariannus) is confined to the Mariana Islands and to Ulithi, an atoll in the Caroline Islands, where it is threatened mainly by habitat destruction and hunting for food.

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The Polynesian Realm

Bannerman’s shearwater (Puffinus bannermani) is a small seabird that breeds only on Higashijima and perhaps Chichi Jima, as well as on Minami-Iwoto in the Volcano Islands. The Micronesian imperial pigeon (Ducula oceanica) is, as a species, found in the Caroline Islands, Marshall Islands, and Nauru. Its various subspecies will be discussed below. The Micronesian saw-tailed gecko (Perochirus ateles) was historically found throughout the Micronesian Region and remains fairly common despite have been extirpated from some islands, such as Guam. It is threatened mainly by competition from introduced Pacific house geckos (Hemidactylus frenatus) and by predation from feral cats. Boettger’s emo skink (Emoia boettgeri) is found primarily in the Caroline Islands, where it has been recorded from Pohnpei, Sapwuahfik Atoll, and the Mortlock Islands. The species also occurs marginally in the Marshall Islands, where it known from three small islets within Arno Atoll and historically from Majuro Atoll as well. It is everywhere experiencing declines due to forest loss and degradation as well as from predation by invasive mammalian predators.

The Bonin Islands Also known as the Ogasawara Islands (Ogasawara Gunto in Japanese), the Bonin Islands are a group of over 30 subtropical and tropical islands of volcanic origin about 1300 km southeast of Japan. Because they have never been connected to a continent, many of the animals and plants here have undergone unique evolutionary processes. This has led to the islands’ nickname of the ‘The Galápagos of the Orient’. The Bonin sambar deer (Rusa unicolor boninensis) is known only from the remains of two specimens collected in the early twentieth century. The Bonin flying fox (Pteropus pselaphon) is confined to a few islands in the archipelago, where the total population is thought to be less than 300.

Chichi Jima Chichi Jima is the largest island in the Bonin Islands group. At least four species have been driven to extinction on this island owing to enormous forest destruction and introduced pigs and rats. The Bonin wood pigeon (Columba versicolor) is known to have existed only on Chichi Jima and Nakondoshima. It was last recorded in 1889. The Bonin rufous night heron (Nycticorax caledonicus crassirostris) was last observed in 1889. The Bonin thrush (Zoothera terrestris) is known only from four specimens collected in 1828. It became extinct during the nineteenth century. The Bonin grosbeak (Carpodacus ferreorostris) is known only from two specimens collected in 1828. It became extinct during the nineteenth century.

Haha Jima Haha Jima is the second largest of the Bonin Islands.

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Figure 8.1 Illustration of a Bonin thrush from 1828. (Credit: F. H. von Kittlitz.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

Sturdee’s pipistrelle bat (Pipistrellus sturdeei) is a small species known from only a single specimen collected in 1915. It was said to have come from Haha Jima, but there is the possibility that it may have originated elsewhere.

The Volcano Islands The Volcano Islands (Kazan Retto in Japanese; also known as the Iwo Islands) are a group of three actively volcanic islands located south of the Bonin Islands. Matsudaira’s storm petrel (Hydrobates matsudairae) is fairly widespread throughout the eastern Pacific and central Indian oceans, but is only known to breed for certain in the Volcano Islands (Minami and perhaps formerly on Kita). In 2004 the total population was estimated at around 20,000.

Iwo Jima Iwo Jima is unusually flat and featureless for a volcanic island, apart from Mount Suribachi on its southern tip. The Iwo Jima white-browed crake (Amaurornis cinerea brevipes) was wiped out by introduced cats and rats. It was last collected in 1911 and last seen in 1924.

The Mariana Islands The Mariana Islands are a crescent-shaped arc of 10 uninhabited and mostly dormant volcanic peaks and 5 limestone and coral islands located about 2000 km east of the Philippines. Once covered by dense forests, all but one of the latter islands have been inhabited by Europeans for 350 years and are intensively cultivated. Cattle and pigs were introduced early. Probably several species of birds were extirpated before zoological exploration began in 1822, which may explain why there are far fewer species of bats, birds, and reptiles on the large Marianas than on the smaller islands of Caroline Islands group, many of which are highly threatened. The Mariana sheath-tailed bat (Emballonura semicaudata rotensis) was historically found on Guam, Rota, Tinian, Saipan, and Aguigan. It is now entirely confined to the latter island,

The Micronesian Region

where the total population of around 500 is known to roost in just three caves. The Micronesian scrubfowl (Megapodius laperouse) is a chicken-like bird divided into two subspecies. The Mariana scrubfowl (M. l. laperouse) has been exterminated on four of the Mariana Islands, including Guam, owing to habitat destruction and predation by rats, cats, and dogs, and barely survives on only five islands. Owston’s rail (Hypotaenidia owstoni) is a flightless species that was historically endemic to Guam, where it was extirpated by the late 1980s due to feral cat predation. Fortunately, a captive breeding programme was established, and populations have been introduced onto Rota and Cocos Island. The Mariana fruit dove (Ptilinopus roseicapilla) is still fairly common on four of the Northern Mariana Islands (Saipan, Tinian, Aguijan, and Rota). It has become extirpated from Guam owing to predation by the introduced brown tree snake (Boiga irregularis), although vagrants originating from Rota are reported on occasion. Threats include habitat destruction, hunting, and introduced predators. The Mariana swiftlet (Aerodramus bartschi) is a bat-like bird that was historically found on Guam, Saipan, Tinian, Rota, and Aguigan. Invasive species, pesticides and disturbance of its nesting caves all contributed to a significant decline during the mid-twentieth century, and the species has long been extirpated from Tinian and Rota. A small breeding colony was successfully introduced to the Hawaiian Island of Oahu in the 1960s, and a reintroduction is planned for Rota. The Mariana crow (Corvus kubaryi) is a very rare species confined to northern Guam and Rota in the Northern Mariana Islands, where it has declined steadily since the 1960s. It has nearly been extirpated on Guam, where it has been decimated by the introduction of the brown tree snake (Boiga irregularis). On Rota the primary threat is loss of habitat. A small number of birds from the latter island have been relocated to the Guam National Wildlife Refuge. The total world population is thought to be less than 250. The golden white-eye (Cleptornis marchei) is a type of passerine bird now confined to Saipan, Aguigan, and Sarigan (where it was successfully translocated in 2011–12), but which appears to have formerly occurred on Tinian and Rota as well. It is threatened by habitat destruction and by predation from introduced brown tree snakes (Boiga irregularis). The Northern Mariana white-eye (Zosterops saypani) is confined to Tinian, Saipan, and Aguijan, where it is fairly common. The Mariana emo skink (Emoia slevini) was historically found throughout the Mariana Islands, but has been extirpated from many of them due to introduced tree snakes. It apparently still survives on Guguan, Alamagan, Asuncion, Sarigan, and Maug.

Pagan Pagan is a double island consisting of two stratovolcanoes joined by a narrow strip of land, located about 320 km north

of Saipan. The inhabitants were all evacuated due to eruptions in 1981. The Pagan reed warbler (Acrocephalus yamashinae) likely became extinct in the early 1970s owing to human development and introduced feral ungulates. The volcanic eruption on the island in 1981 destroyed any lingering hope of its survival.

Saipan Saipan is the second largest of the Mariana Islands, after Guam. The western side is lined with sandy beaches and a large lagoon, while the eastern shore is composed primarily of rugged rocky cliffs. The highest point is a limestone mountain called Mount Tapochau. Remaining native forest occurs in small, isolated fragments on steep slopes. The Saipan reed warbler (Acrocephalus hiwae) is confined to Saipan and Alamagan, where it is seriously threatened on the former island by introduced brown tree snakes (Boiga irregularis).

Aguigan Aguigan (also known as Aguihan, Aguiguan, and Aguijan) is a small, uninhabited coralline island located 8 km north-west of Tinian. Feral goats have destroyed much of the native vegetation. The Aguigan reed warbler (Acrocephalus nijoi) has not been reported since 1995 despite extensive surveys, and is now believed to be extinct.

Tinian Tinian is located about 10 km south-west of Saipan. The Tinian monarch (Metabolus takatsukasae) is a type of passerine bird that had been reduced to about 40 individuals by the late 1960s, but has made a remarkable recovery since then and now numbers in the tens of thousands. Nevertheless, it remains vulnerable to typhoons and the potential introduction of the brown tree snake (Boiga irregularis) from nearby Guam. In 2015–16 it was successfully translocated to Guguan in the Northern Marianas.

Rota Rota is a small island located north of Guam. The Rota white-eye (Zosterops rotensis) is a type of passerine bird endemic to the island, where it was formerly common and widespread but now largely confined to the upper escarpments of a single plateau.

Guam The southernmost of the Mariana Islands, Guam is the largest island in Micronesia. From the seventeenth to the nineteenth centuries the Spanish introduced pigs, dogs, chickens, Philippine deer (Rusa marianna), and water buffalo (Bubalus bubalis). Later introductions included cane toads, the East African giant land snail, and various frog species. However, it was the accidental introduction of the brown tree snake (Boiga

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The Caroline ground dove (Pampusana kubaryi) is confined to the islands of Pohnpei, Chuuk, and Ant Atoll. The Caroline lorikeet (Trichoglossus rubiginosus) is nowadays confined to Pohnpei and nearby Ahnd Atoll, but is thought to have formerly had a wider distribution within the Caroline Islands. The Caroline goby (Sicyopterus eudentatus) is confined to a few streams on Pohnpei and Kosrae.

Pohnpei Figure 8.2 Guam flycatcher photographed in the 1940s. (Credit: Smithsonian Miscellaneous Collections.)

irregularis) near the end of World War II that has really devastated the endemic wildlife. During the twentieth century much of the island’s once-dense forests were replaced by thick tangan-tangan brush (Leucaena leucocephala). The Guam flying fox (Pteropus tokudae) is only known from three specimens, the last having been shot by hunters in 1968. There was a possible sighting in the late 1970s, but nothing since. The Guam kingfisher (Todiramphus cinnamominus) remains extinct in the wild, but is well-established in captivity. The Guam flycatcher (Myiagra freycineti) was considered common up until the 1970s, but quickly went extinct in 1983 along with most of the island’s endemic birds. The bridled white-eye (Zosterops conspicullatus) was a type of passerine bird endemic to Guam that is believed to have gone extinct in 1983 due to predation by introduced brown tree snakes (Boiga irregularis). The Guam reed warbler (Acrocephalus luscinius) was last seen in the Agana Swamp in 1969, despite having been reported as fairly common there just a year or two before.

Located in the eastern Caroline Islands, Pohnpei (formerly known as Ponape Island) is the largest and highest island in Micronesia and one of the few islands within the region to maintain its central volcano. The invasive introduction of the brown tree snake (Boiga irregularis) has had a devastating effect on avian populations in recent decades, as has habitat destruction. The Pohnpei flying fox (Pteropus molossinus) is known for certain only from Pohnpei as well as Ant and Pakin atolls. Reports from the Nomoi Islands appear to be erroneous. The Pohnpei imperial pigeon (Ducula oceanica townsendi) is confined to Pohnpei. The Pohnpei starling (Aplonis pelzelni) is (or was) endemic to Pohnpei. The species declined dramatically sometime after 1930, when about 60 specimens were collected in a 3-month period. It was not located during a survey conducted in 1983 and is considered extinct by some authorities. However, since the 1970s there been several possible sightings, including a specimen collected in 1995. The Pohnpei kingfisher (Todiramphus reichenbachii) is confined to the lowland forests of Pohnpei, where it remains relatively common but declining. The Pohnpei cicadabird (Edolisoma insperatum) is confined to Pohnpei. The Pohnpei emo skink (Emoia ponapea) is confined to the forests of Pohnpei.

The Caroline Islands

Kosrae

The Caroline Islands number about 500 tiny, widely scattered islands extending some 2600 km across the Pacific Ocean to the north of New Guinea. Most of them are low, flat coral atolls, but a few are mountainous and of volcanic origin. Several have become infested by rats brought long ago by whalers. The pelagic flying fox (Pteropus pelagicus), as a species, is found on the Chuuk and Nomoi islands, where it is seriously threatened by rising sea levels due to climate change. The Chuuk flying fox (P. p. insularis) was historically known from the Chuuk Islands and Namonuito Atoll, although the latter population may now be extirpated. The Caroline Islands sheath-tailed bat (Emballonura semicaudata sulcata) is confined to the islands of Chuuk and Pohnpei. The Palau imperial pigeon (Ducula oceanica monacha) is confined to Yap and the Palau Islands.

Kosrae (formerly known as Kusaie) is the easternmost of the Caroline Islands. It is mountainous and still largely unspoiled. The Kosrae flying fox (Pteropus ualanus) declined rapidly in 1927 owing to a disease epidemic. A few colonies of 100–400 bats were reported in 1983, after which significant numbers were harvested for export to Guam as food (it is not locally hunted). The species was reported to be uncommon in 1989 but relatively stable. The Kosrae crake (Zapornia monasa) was a flightless species known only from two specimens collected from coastal swamps in 1827–28. It is believed to have gone extinct within half a century due to introduced rats. The Kosrae imperial pigeon (Ducula oceanica oceanica) is confined to Kosrae. The Kosrae starling (Aplonis corvina) is known only from two specimens collected from mountain forests in 1828. It was extinct by 1880 due to introduced rats.

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The Palau sheath-tailed bat (Emballonura semicaudata palauensis) is found on the islands of Koror, Peleliu, Babelthuap, and Anguar, where it is reasonably common. The Palau scrubfowl (Megapodius laperouse senex) has been exterminated on two islands and barely survives on four others. The Palau ground dove (Pampusana canifrons) occurs throughout the Palau Archipelago but is rare on most islands. Surveys in the late 1970s estimated a total population of around 500. However, there have been very few records in recent years, in part because of the inaccessibility of the bird’s habitat along with their secretive nature. Cats and rats are known to be present on at least some of the islands. The Rock Islands The Rock Islands are a collection of limestone and coral uprises located in Palau’s Southern Lagoon. They were declared a world heritage site in 2012. The Seventy Islands scaly-toed gecko (Lepidodactylus paurolepis) is known only from five specimens collected from three small islands in the Ngerukeuid Islands Wildlife Preserve.

Figure 8.3 Illustration of a Kosrae starling from 1831. (Credit: F. H. von Kittlitz.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

Anguar Anguar is the southernmost of the Palau Islands. The Palau forest dragon (Hypsilurus godeffroyi) is known only from two specimens of uncertain provenance collected in 1867. Subfossil remains of an agamid lizard here considered to be conspecific have been discovered on Anguar.

Yap Island

The Chuuk Islands

Yap Island is actually made up of four contiguous (although separated by water) islands surrounded by a common coral reef. The Yap flying fox (Pteropus yapensis) is confined to the four adjacent main islands of Yap, where it is seriously threatened by hunting. The Yap reed warbler (Acrocephalus astrolabii) is known only from specimens collected in 1838 or 1839, the precise origins of which remain unclear. The Yap cicadabird (Edolisoma nesiotis) is fairly widespread on Yap, but in very low numbers.

The Chuuk Islands (formerly known as the Truk Islands, and also known as Chuuk Lagoon) are a group of 11 major islands and dozens of smaller ones centered around a central lagoon with a common fringing coral reef. The islands have been heavily deforested. The Chuuk imperial pigeon (Ducula oceanica teraokai) is confined to Chuuk. The Chuuk monarch (Metabolus rugensis) is a type of passerine bird widely but sparsely distributed on all, or nearly all, of the high lagoon islands as well as some of the outer reef islets.

The Palau Islands

The Faichuk Islands The Faichuk Islands are located in western Chuuk Lagoon. The teardrop white-eye (Rukia ruki) is a type of passerine bird confined to four tiny islands in the Faichuk group. In 1984 the total population was estimated at 530.

Located about 900 km east of Mindanao, the Palau Islands are an archipelago of about 340 coral-reef islands forming the western chain of the Caroline Islands. The large Palau flying fox (Pteropus pilosus) is known only from two specimens collected prior to 1874, and extensive surveys over the years since have failed to locate it. The precise causes of its presumed extinction are unknown, although hunting for food by local people along with forest degradation seems most likely. The small Palau flying fox (P. pelewensis) is found on the main islands of Palau as well as on a few more distant islets, where it is seriously threatened by hunting.

Namoluk Atoll Namoluk is a small atoll located about 185 km south-east of Chuuk. The Toimon scaly-toed gecko (Lepidodactylus oligoporus) is known only from a few specimens collected on Toimon Island, part of Namoluk Atoll.

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The Polynesian Realm Figure 8.4 Wake Island rail photographed in 1936. (Credit: Unknown.)

The Nomoi Islands The Nomoi Islands (also known as the Mortlock Islands) are a group of three large atolls located about 250 km south-east of the Chuuk Islands. The Nomoi flying fox (Pteropus pelagicus pelagicus) is confined to the Nomoi Islands, where it has been extirpated from at least two islands.

The Marshall Islands The Marshall Islands, located near the equator and slightly west of the International Date Line, are spread out over 29 coral atolls and comprise 1156 individual islands and islets. During the mid-twentieth century they were the setting for much nuclear testing on the part of the Americans. The latter began in 1946 on Bikini Atoll after residents were evacuated. Over the years, 67 weapon tests were conducted, including the 15megaton Castle Bravo hydrogen bomb test, which produced significant fallout in the region. The testing concluded in 1958. Over the years, just one of over 60 islands were cleaned by the U.S. government, and many of the islanders and their descendants still live in exile as their islands remain contaminated with high levels of radiation. Many other low-lying islands are additionally threatened now by rising sea levels due to global warming, which threaten to submerge them entirely. The Marshall Islands imperial pigeon (Ducula oceanica ratakensis) is confined to the Marshall Islands.

Wake Island Wake Island is an isolated low atoll in the central Pacific between the Marianas and Hawaii. With limited freshwater resources, no harbour, and no plans for development, it remained uninhabited during the early twentieth century. It

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did, however, boast a large seabird population, which unfortunately attracted Japanese feather poachers. The Wake Island rail (Hypotaenidia wakensis) was a nearly flightless species that was the only native land bird on the island. It was exterminated during its occupation by Japanese troops in World War II, probably in 1945.

Nauru Nauru is a tiny, isolated atoll located in the south-western Pacific. Once heavily vegetated, its environment was largely destroyed by strip-mining for its phosphorous deposits and introduced species. The Nauru reed warbler (Acrocephalus rehsei) is confined to the island, where it remains relatively common.

The Gilbert Islands The Gilbert Islands (formerly known as the Kingsmill Islands) are a chain of 16 coral atolls located about 1500 km north of Fiji. The islands have no known endemic species, although it is possible that a now-extinct population of Micronesian imperial pigeon (Ducula oceanica) represented a distinct subspecies. In addition, three species of threatened petrel (Pterodroma) as well as the bristle-thighed curlew (Numenius tahitiensis) are found on the islands.

Balance for the Micronesian Region Modern human settlement of the Micronesian Region began several thousand years ago, although there are competing theories about where these people came from and when they first arrived. The Marianas were the first islands in all of Oceania to be colonized by Austronesian peoples, having

The Novozelandic Region

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

2 species

~ species

~ species

8 species

~ subspecies

~ subspecies

~ subspecies

6 subspecies

2 taxa

~ taxa

~ taxa

14 taxa

12 species

1 species

1 species

23 species

2 subspecies

~ subspecies

~ subspecies

5 subspecies

14 taxa

1 taxon

1 taxon

28 taxa

1 species

~ species

~ species

5 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

1 taxon

~ taxa

~ taxa

5 taxa

~ species

~ species

~ species

~ species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

~ taxa

~ taxa

~ taxa

~ species

~ species

~ species

1 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

~ taxa

~ taxa

1 taxon

15 species

1 species

1 species

37 species

2 subspecies

~ subspecies

~ subspecies

11 subspecies

17 taxa

1 taxon

1 taxon

48 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

been settled by voyagers from the Philippines in approximately 1500 B . C . These populations gradually moved southwards until they reached the Bismarck Archipelago, the Solomons and coastal New Guinea by 1300 B . C . By 1200 B . C . they were once more crossing open seas beyond interisland visibility, reaching Vanuatu, Fiji, and New Caledonia before continuing eastward to become the ancestors of the Polynesian people. Further migrations by other Austronesians also followed, most likely from Sulawesi, settling Palau and Yap by around 1000 B . C . The earliest known contact between these Micronesian indigenous peoples and Europeans was in 1521, when a Spanish expedition under Ferdinand Magellan reached the Mariana Islands. Further contact was made during the sixteenth century, although initial encounters between Europeans and natives were often very brief. In the early seventeenth century Spain colonized Guam, parts of the Marianas, and the Caroline Islands, creating the Spanish East Indies. During the early nineteenth century American missionaries began to have an influence, and thereafter the colonies would be shared among American, German, and British interests until most achieved independence during the twentieth century. Today, most of Micronesia consists of independent states with the exception of the Northern Mariana Islands, Guam, and Wake islands, all of which continue to be U.S. territories.

In recent historical time (i.e. since A . D . 1500), the Micronesian Region has lost at least 15 species/2 subspecies of vertebrates. Among the extinct forms 2 species are mammals, 12 species/2 subspecies are birds, and 1 species is a reptile. One other species is possibly extinct, and 1 species is currently extinct in the wild. In addition, there are 37 species/11 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 8 species/6 subspecies are mammals, 23 species/5 subspecies are birds, 5 species are reptiles, and 1 species is a freshwater fish.

The Novozelandic Region The Novozelandic Region consists of the main North, South, and Stewart islands of New Zealand in the subtropical and temperate zones, along with a number of smaller sub-Antarctic island groups. Geologically it consists of fragments of the ancient supercontinent Gondwana, but has been separated from other landmasses for tens of millions of years. Together they comprise the most isolated large landmass in the world, there being no evidence that they were ever connected with either Australia or Antarctica. As a result, many of New

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The Polynesian Realm

Zealand’s plants and animals followed lines of evolution that have made them unique. Indeed, in its virgin state, New Zealand was a sort of vast living museum. Zoologically speaking, New Zealand is strikingly different from all other terrestrial realms and regions, because it includes no native mammals except for bats and sea lions. There are a few reptiles (most notably the tuataras, lone survivors of an ancient order that disappeared elsewhere a hundred million years ago), along with some primitive amphibians and fishes. But it is the birds that dominate, having replaced terrestrial mammals and filled a wide range of habitats in the process. The most spectacular of these were six genera and nine species of giant flightless moas, the two largest of which reached a height of about 3.6 m with neck outstretched and weighed about 230 kg, thus putting them among the largest birds ever known. They were grazing birds utilizing the grasslands as kangaroos do in Australia and antelopes in Africa. The first humans to reach New Zealand, the Polynesians, arrived sometime between A . D . 1250 and 1300, bringing with them dogs and rats which would ultimately wipe out a great many species. Sometime prior to the arrival of Europeans a subsequent wave of immigration had already established a distinct Maori culture there. Although the islands still consisted mainly of pristine forests and savanna by this point, the indigenous people had already largely wiped out the large, flightless moas discussed above, having hunted them for food as is shown clearly by many finds made on both the North and South Islands. Historically these birds would have had few or no enemies, a fact which no doubt made them unwary and easy to catch. Their huge eggs must also have been coveted as food and for domestic utensils. Many other species, mostly birds, were also exterminated by the Maori. The New Zealand sea lion (Phocarctos hookeri) has a highly restricted distribution for a marine mammal, being nowadays confined to coastal areas of the southernmost islands of New Zealand and their surrounding waters. Its principal breeding colonies are in the Auckland Islands, with most of the remaining ones on Campbell Island. Small numbers occur on the southern coast of South Island and on Stewart Island. The species was at one time found throughout New Zealand but is now perhaps the world’s rarest sea lion, with a total population of around 10,000. The northern royal albatross (Diomedea sanfordi) ranges widely across the southern oceans as far north as Peru, but breeds only in the Chatham and Auckland islands and at Taiaroa Head on the Otago Peninsula, South Island (the latter being the only albatross colony on a human-inhabited mainland in the Southern Hemisphere). Hutton’s shearwater (Puffinus huttoni) is a medium-sized seabird that ranges throughout both New Zealand and Australian waters but currently breeds only in two natural colonies within the Seaward Kaikoura Range in north-eastern South Island, with a third (protected and artificial) having been established near the town of Kaikoura itself. Six other natural colonies, in both the Seaward and Inland Kaikoura ranges,

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have been wiped out in recent decades by introduced pigs and stoats. The yellow-eyed penguin (Megadyptes antipodes) breeds on the eastern and south-eastern coasts of South Island, Stewart Island and outliers, the Auckland Islands and the Campbell Island group. It is threatened by disease, invasive predators, and fisheries by-catch. The New Zealand crested penguin (Eudyptes pachyrhynchus) now breeds only along the south-western coast of South Island and on Stewart Island and its outliers. At other times the species disperses widely through the sub-Antarctic islands and occasionally as far as Tasmania and south-eastern coastal Australia. It is threatened mainly by introduced predators such as dogs, cats, rats, and stoats, as well as by human disturbance of its nesting sites. The New Zealand swan (Cygnus sumnerensis) is an extinct species known only from fossil remains discovered during the nineteenth century on South Island and the Chatham Islands. Two subspecies will be discussed below. The New Zealand brown teal (Anas chlorotis) was formerly widespread throughout the mainland and offshore islands, but has been severely affected by the loss of swamps, ponds, and forests. It occurred in the Chatham Islands until about 1925, and on Stewart Island until the early 1970s. It has since disappeared from South Island as well, and is now largely restricted to northern North Island. In recent years populations have been reintroduced to Kapiti and Mana Islands and to the Zealandia Sanctuary near Wellington. In 2011 the total population was estimated at 1500–2500. The Auckland merganser (Mergus australis) at one time occurred on South and Stewart islands, but was already extinct there when Europeans arrived. It disappeared from the Auckland Islands after settlers and their domestic animals invaded the group. The last sighting was in 1902. The nominate form of the white-necked petrel (Pterodroma cervicalis cervicalis) occurs throughout a large part of the Pacific, but for breeding purposes is almost entirely confined to Macauley Island in the Kermadec Islands. It formerly bred on Raoul Island as well, but has been extirpated from there. A second small breeding colony has been established on Philip Island, near Norfolk Island. The New Zealand shore plover (Thinornis novaeseelandiae) was extirpated from mainland New Zealand during the nineteenth century, and today breeds only in South East Island in the Chatham Islands (with vagrants to Pitt Island). The wrybill (Anarhynchus frontalis) is a type of plover with a curious, sideways-bent bill that breeds only along the rivers of central South Island, before returning to coastal areas throughout New Zealand. It is threatened by habitat destruction. The weka (Gallirallus australis), also known as the Maori woodhen, is a flightless rail endemic to New Zealand. The buff weka (G. a. hectori) historically occurred in the eastern districts of South Island where the Maoris used to kill large numbers for food, sometimes bagging as many as 2000 in

The Novozelandic Region

a single hunt. It became extinct there about 1925. The subspecies also occurred on Chatham and Pitt islands, but was extirpated there before 1868. Fortunately, in 1905 specimens were reintroduced from South Island, where they increased rapidly and ultimately became fairly abundant. Attempts to translocate a population back to the Canterbury region of South Island in 1961 failed, but another introduction to Mao Waho Island (in Lake Wanaka), and from there to Pigeon and Pig Island (in Lake Wakatipu) have been much more successful. Fishes of the genus Galaxias are scaleless and tubular in body form. Many are amphidromous, with the fry going to sea after hatching and returning as juveniles to freshwater where they grow to adulthood. The giant galaxias (G. argenteus) occurs in coastal streams, wetland lakes, and lagoons across North Island, South Island, Stewart Island, and the Chatham Islands. The species has undergone a significant decline due to loss of habitat and has been extirpated from many parts of its former range.

The Kermadec Islands Located about halfway between New Zealand and Tonga, the subtropical, volcanic Kermadec Islands (Rangitahua in Maori) include four main islands and some isolated rocks. There are no native land mammals, but the arc is an important breeding site for several species of seabirds. The Kermadec red-fronted parakeet (Cyanoramphus novaezelandiae cyanurus) was historically found throughout the islands, but was eradicated from the principal island of Raoul in the early nineteenth century as a result of introduced cats, goats, and rats. It thereafter survived only on the nearby Herald Islets and on Macauley Island. In 2008, following lengthy island restoration efforts that included the removal of invasive predators, the subspecies began to recolonize Raoul on its own from the Herald Islets.

New Zealand Main Islands The main islands of New Zealand consist of North Island and South Island, separated from one another by the narrow Cook Strait, and Stewart Island. The New Zealand greater short-tailed bat (Mystacina robusta) is known from subfossil remains from both South Island and North Island. By the time of European settlement it was probably already restricted to a few small islands off the coast of Stewart Island, where it was last seen on Big South Cape Island in 1967. There have been unconfirmed reports in recent years, but the species is certainly very rare if not already extinct. The New Zealand lesser short-tailed bat (M. tuberculata), one of the most terrestrial of all bats, is divided into three subspecies historically found throughout North Island, South Island, and their satellites. The nominate form (M. t. tuberculata) is confined to a small area of southern North Island, western and southern South Island, and Codfish Island. It is threatened by forest clearance and introduced predators.

The New Zealand long-tailed bat (Chalinolobus tuberculatus) is still found patchily on North Island, South Island, and Stewart Island, but is now rare or absent from many areas where it was formerly common. It is threatened by habitat destruction and introduced species. The New Zealand king shag (Leucocarbo carunculatus) is a large, black and white cormorant confined to a few small islets and rock stacks lying across the outer reaches of Marlborough Sound, north-western South Island. Shooting by fisherman historically destroyed many of their former breeding colonies, but the birds have long been fully protected. The population (about 1000) has been stable for over half a century, with the only major threat being fishing nets. The bronze shag (L. chalconotus) breeds in a small and decreasing number of colonies along south-eastern South Island and on Stewart Island. Cook’s petrel (Pterodroma cookii) breeds only on three small islands (Little Barrier Island, Great Barrier Island, and Codfish Island), at other times dispersing across the Pacific Ocean as far as the western coast of the Americas. It is threatened by introduced pigs, dogs, rats, and cats, which attack their nests and burrows. Pycroft’s petrel (P. pycrofti) breeds under forest cover on a dozen small offshore islands along the coast of New Zealand, with subfossil evidence indicating that it historically did so on Norfolk and Lord Howe islands as well). It too disperses widely across the Pacific at other times. Translocations have been undertaken with both species. The black petrel (Procellaria parkinsoni) ranges widely across the Pacific Ocean from Australia to Central America and the north-western coast of South America, but currently breeds only on Great Barrier Island and Little Barrier Island. Colonies were formerly found in the mountains of North Island and South Island as well, but were extirpated during the mid-twentieth century by introduced pigs and feral cats. The New Zealand storm petrel (Fregetta maoriana) was long known only from putative fossil material and three specimens collected during the nineteenth century, two from the eastern coast of North Island and another of unknown provenance but thought to be from South Island. Thought to be extinct, it was rediscovered in 2003 off north-western North Island and a small breeding colony was subsequently discovered on Little Barrier Island by researchers. The black-fronted tern (Chlidonias albostriatus) breeds only along riverbanks on South Island, from where it disperses at other times to coastal areas from Stewart Island to southern North Island. It is threatened mainly by introduced species. The New Zealand fairy tern (Sternula nereis davisae) breeds only in coastal areas of northern New Zealand, where it is threatened by introduced predators. The black-billed gull (Larus bulleri) is found widely but patchily along the rivers and coasts of North Island and South Island. It is threatened by introduced predators as well as human disturbance of its nesting colonies.

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The southern red-breasted plover (Charadrius obscurus) is a type of shorebird historically found on Stewart Island and the southern tip of South Island, but is now restricted, when breeding, to the former. By the early 1990s the species had been reduced to just 62 birds, but thanks to a feral cat eradication programme it recovered rapidly. Unfortunately, despite management the population has since declined once more for reasons that are unclear, and currently numbers around 30–40 breeding pairs. The New Zealand bittern (Ixobrychus novaezelandiae) was a type of heron only known for certain from South Island, although subfossil material has been discovered on North Island as well (old reports of living birds there were likely based on misidentifications). On South Island only a few specimens were ever recorded, the last in the 1890s. The reason for its extinction is unknown. Hodgen’s waterhen (Tribonyx hodgenorum) was a type of flightless rail that, to judge from subfossil remains, was historically widespread on both North Island and South Island. It was driven extinct during the seventeenth century as a result of human hunting and rat predation. The black stilt (Himantopus novaezelandiae) is a seriously threatened wading bird that formerly bred and wintered across both North Island and South Island. Following a long-term decline due to habitat destruction and predation from introduced species it is now confined, during the nesting season, to the upper Waitaki Valley in south-central South Island, where in 2020 the total world population was estimated at less than 200. There is a small but successful captive-breeding programme. Finsch’s duck (Chenonetta finschi) was a large, flightless, terrestrial species that was historically widespread and common on both North Island and South Island. It is thought to have been driven extinct by a combination of human hunting and predation from introduced species, particularly rats, sometime between the fifteenth and seventeenth centuries, although an account of a ‘goose’ killed in 1870 suggests that it may have survived far longer. The blue duck (Hymenolaimus malacorhynchos) was historically widespread across North Island and South Island, but since the arrival of Europeans its range has become highly fragmented. It is now largely confined to wetland areas within the forested mountain ranges of central North Island and western South Island, where in 2010 the total population was estimated at 2500–3000. The laughing owl (Ninox albifacies) is a now-extinct species divided into two subspecies. The South Island laughing owl (N. a. albifacies) was native to South and Stewart Islands, with bones known from the Chatham Islands as well. It was plentiful in many areas during the early nineteenth century but started to disappear thereafter. The last specimens were collected in 1914, with unconfirmed reports until the 1960s. The cause of the decline was habitat changes and predators; for the latter this ground-nesting owl was an easy prey. The kakapo or owl parrot (Strigops habroptila) is a large, heavy species that has long been one of New Zealand’s most

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Figure 8.5 A live South Island laughing owl photographed c.1889–1910. (Credit: H. C. C. Wright.)

seriously threatened birds. Flightless and nocturnal, it lives on the ground and climbs trees from which it is able to glide. It was historically widely distributed over both the North and South islands as well as on Stewart Island, but suffered considerably from the destruction of forests and introduced animals. It seems to have been exterminated on North Island by the 1920s and occurred only in a few isolated areas on South Island. One of its last refuges was in Fiordland, where by the 1940s it was becoming scarce. In the 1950s the New Zealand Wildlife Service was established and began to make regular expeditions to search for kakapos, both in the south-west and north-west of South Island, but only a few recent signs were found. Finally, in 1958, a single individual was caught and released in the Milford Sound catchment area in Fiordland. Six more were captured in 1961, one of which was released and the others transferred to aviaries of the Mount Bruce Bird Reserve near Masterton, on North Island. Within a few months four of these birds died, and the fifth about four years later. Over the next dozen years regular expeditions again found few signs of kakapos. Only one was captured, in 1967, which died the following year. By the early 1970s it was uncertain whether the species still survived. But 14 birds were discovered between 1974 and 1976, all of which, unfortunately, were males. In 1977 the species was reported from Stewart Island, where soon after a large number were located. These were being decimated by feral cats and, after efforts to control the latter, it was ultimately decided to translocate all surviving

The Novozelandic Region Figure 8.6 An illustration of a kakapo from 1873. (Credit: John G. Keulemans.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

birds to predator-free islands (Maud, Codfish, Mana, and Little Barrier) where they could be closely monitored and protected. Sixty-five kakapo (43 males and 22 females) were successfully transferred onto the four islands. Little Barrier Island was eventually deemed unsuitable due to its rugged landscape and continued presence of rats, and its birds (along with those on Mana) were evacuated in 1998. The entire population on Codfish Island was temporarily relocated in 1999 to Pearl Island while rats were being eliminated, and in 2005 all the kakapo on Pearl and Chalky islands were moved to Anchor Island. In 2012 seven birds were transferred back to Little Barrier Island. As of 2018 the total known adult population was around 150. The New Zealand kaka (Nestor meridionalis) is a type of large parrot divided into two subspecies. The South Island kaka (N. m. meridionalis) is mostly found west of the Southern Alps on South Island as well as on Stewart Island and several offshore islands. It was much reduced in the past by habitat destruction and hunting, and continues to be rare and declining due to introduced predators. Malherbe’s parakeet (Cyanoramphus malherbi) once occurred throughout New Zealand and its offshore islands, but was eradicated everywhere due to introduced rats. It is now confined to a small area of forest in South Island’s Southern Alps. Iredale’s snipe (Coenocorypha iredalei) was historically found on South Island and Stewart Island along with a few satellite islands. Following the introduction of rats by the Polynesians it was extirpated on South Island and Stewart Island, and thereafter survived on approximately nine small

islands off Stewart Island, from where it was progressively wiped out during the nineteenth and twentieth centuries due to the arrival of rats and other introduced predators. By the early 1960s it was entirely confined to Big South Cape Island, then still predator-free, but in 1964 rats accidentally came ashore by way of a visitor’s boat. In August, 1964, two of the birds were moved to a nearby islet free of rats, but both died. In December of the same year searchers failed to find any snipe left on Big South Cape Island, and none were ever reported again. The little spotted kiwi (Apteryx owenii) is a small ratite bird that was historically found in forested areas on both North Island and South Island, but was wiped out from there by introduced predators. It survived only on Kapiti, an island located about 5 km off the western coast of lower North Island. In recent years some have been translocated from there to other small offshore islands and mainland reserves protected by pest-exclusion fences. The New Zealand quail (Coturnix novaezelandiae) was reported to be very common on the North Island in the 1770s. On the South Island it was abundant until 1865, when it declined rapidly; by 1880 the species was gone throughout New Zealand. Reports from the period 1865–80 agree that extinction resulted from fires that destroyed food, cover, and the birds themselves, combined with predation by dogs, cats, and rats. Later analyses suggest that diseases imported with pheasants and other game birds may also help to explain the extraordinarily rapid disappearance of this species. The South Island kokako (Callaeas cinereus) is (or was) a crow-like bird that was abundant on South Island as well as on

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The Polynesian Realm

Stewart Island as late as 1888, but disappeared from the latter during the 1940s. It seems to have been confined to the Fiordland area thereafter, with the last confirmed sightings taking place in 2007, and before that in 1967. It is almost certainly extinct. The South Island saddleback (Philesturnus carunculatus) is a type of wattlebird that once ranged throughout South and Stewart islands as well as the South Cape Islands (Big South Cape, Solomon, and Pukeweka islands). Predation by introduced mammals (mainly ship rats) led to its extirpation on South and Stewart islands by 1900. When rats invaded the South Cape Islands as well in the early 1960s the New Zealand Wildlife Service successfully translocated 36 individuals from Big South Cape Island to nearby Big and Kaimohu islands in 1964, thereby averting the extinction of the species. They have since been translocated to around 20 offshore islets and one sanctuary on the mainland. By 1969 the birds had become extirpated on Solomon, Pukeweka, and Big South Cape islands, but rats have now been eliminated from the latter and the species has been reintroduced there. The yellowhead (Mohoua ochrocephala) is a rare type of passerine bird that was once widespread on South Island and Stewart Island. It suffered significant declines and extirpations due to introduced predators, leaving only a few surviving populations in scattered lowland forest fragments on South Island. Fortunately, the species has now been successfully translocated to a few safe offshore islands, and the majority of its mainland range is now subject to predator control measures. Lyall’s rockwren (Traversia lyalli) historically occurred on both North Island and South Island, but was extirpated there centuries ago by rats brought by Polynesians. A flightless species, it was particularly vulnerable to introduced predators and was ultimately wiped out on Stephens Island, its last refuge, by feral cats. The well-known story that it was systematically exterminated by a single domestic cat belonging to the local lighthouse keeper is at best an exaggeration. While this particular cat did kill one of the last birds seen, a few more were obtained in the following years before the species finally became extinct in 1895. Pelzeln’s bushwren (Xenicus gilviventris) was historically found on North Island prior to European settlement but is now confined to a few scattered areas on South Island, where it is declining due to nest predation. The tuatara (Sphenodon punctatus) is an ancient reptile endemic to New Zealand. Superficially resembling lizards, they in fact form a distinct lineage of their own and are thus of enormous interest in the study of evolution. The single surviving species once occurred from the northern tip of North Island to near the southernmost part of South Island. They apparently disappeared from the two main islands before the arrival of the Europeans. It is possible that the Maoris contributed to the disappearance, because subfossil remains from their kitchen middens show that they ate it. Maori dogs may also have been involved. But it was primarily

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introduced rats that have been implicated in the decline. Tuatara eggs take more than 15 months to hatch, making them extremely vulnerable, and adults themselves are also easy prey. Fortunately, a few populations managed to survive on a number of small coastal islands to the north and south of North Island. The northern tuatara (S. p. punctatus) occurs mainly on islands off the northern coast of North Island as well as, marginally, on islands in the Cook Strait. They have long enjoyed official protections, and have been successfully translocated to new islands. In 2005 a population was introduced into a heavily fenced sanctuary on mainland North Island near Wellington, where the animals are now permanently established. The Cook Strait gecko (Hoplodactylus stephensi) is confined to Stephens and Maud islands as well as to disjunct coastal forest remnants in the northern Coromandel Peninsula of North Island. The Marlborough green gecko (Naultinus manukanus) is confined to shrubby coastal areas in northern South Island as well as on some of the Cook Strait and Marlborough Sound islands. Hamilton’s frog (Leiopelma hamiltoni) was historically found throughout much of North Island and South Island, but has long been confined to a single rock tumble on Stephens Island, where the population is vulnerable to predation by tuataras and black rats. Translocation to an adjoining area made into a man-made refugium was attempted in 1992, but with limited success as many of the frogs homed back to the original site. A more successful translocation to a nearby island, Nukuwaiata, occurred in 2004 and 2006. The New Zealand grayling (Prototroctes oxyrhynchus) was a type of amphidromous (saltwater and freshwater) smelt that was historically widespread and common in the lowland rivers and streams of both North Island and South Island, where it would go to spawn. A combination of habitat destruction and introduced fish species resulted in a noticeable decline by the late 1870s, and the last known specimens were caught sometime during the late 1920s to early 1930s. It was finally given full legal protection in 1951. The bluegill bully (Gobiomorphus hubbsi) is a type of small goby found patchily in coastal rivers across North Island, South Island, and Great Barrier Island. It is threatened by habitat destruction and degradation. The torrentfish (Cheimarrichthys fosteri) is a migratory species with a complex life cycle: the fry go to sea after hatching and return as juveniles to shallow, fast-flowing rivers across North Island and South Island. It is threatened by habitat destruction and degradation. The Gollum galaxias (Galaxias gollumoides) is a nonmigratory species confined to the southern part of South Island and Stewart Island. The dwarf New Zealand galaxias (G. divergens) is found on North Island and South Island where it is fairly widely dispersed in higher-elevation rivers and springs. The shortjawed galaxias (G. postvectis) is found patchily on North Island, South Island, and a number of

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offshore islands. All are threatened by habitat destruction and degradation and introduced salmonid species. The brown mudfish (Neochanna apoda) is found patchily in shallow swamp-forest wetland fragments in southern North Island and north-western coastal South Island.

North Island North Island (Te Ika-a-Maui in Maori) is the most populous of New Zealand’s landmasses, once covered by a mantle of semitropical and temperate forests rich in interesting species, many of which are not gone. The Kauri lesser short-tailed bat (Mystacina tuberculata auporica) is confined to north-western North Island and Little Barrier Island. The Volcanic Plateau lesser short-tailed bat (M. t. rhyacobia) is found patchily throughout central North Island. The North Island laughing owl (Ninox albifacies rufifacies) was probably exterminated by rats and cats. The last specimens were collected in 1889, with unconfirmed reports until the 1930s. The North Island kaka (Nestor meridionalis septentrionalis) is a type of large parrot found patchily on North Island and on the offshore islands of Kapiti and Little Barrier Island. Two species of large, flightless rails known as takahes (Porphyrio) historically inhabited New Zealand, where they appear to have been widespread. The decline of both is thought to have been due to climate change combined with hunting by early Polynesians. Introduced browsing species such as red deer and wapiti also contributed to it by destroying vegetation. As the reproductive rate of these birds is very low, they were unable to easily recover their losses. The North Island takahe (P. mantelli) was an inhabitant of high-altitude alpine grasslands. Known only from subfossil material and a single possible historic record from 1894, it was exterminated by humaninduced habitat changes and hunting. The North Island snipe (Coenocorypha barrierensis), to judge by subfossil evidence, was historically found throughout North Island but was extirpated there after the arrival of the Polynesians and the associated introduction of Pacific rats (Rattus exulans). It is known to have survived on Little Barrier Island until 1870, when two specimens were found and one captured (later to die in captivity). The birds apparently disappeared soon after, when cats were introduced to the island. The North Island weka (Gallirallus australis greyi) is a type of rail confined to two widely separated population pockets on North Island (the Northland region and Poverty Bay on the eastern coast). The northern brown kiwi (Apteryx mantelli) is a chickensized species that remains relatively common within isolated and fragmented areas of North Island and some adjacent offshore islands. It is threatened mainly by introduced predators such as feral dogs and ferrets. The huia (Heteralocha acutirostris) was a type of wattlebird notable for the fact that the bills of the male and female looked

Figure 8.7 A pair of huia (male in front of female) by John Gerrard Keulemans, 1888. (Credit: J. G. Keulemans.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

as although they belonged to two, totally different species. The male had a short, arched, thick starling-like bill, while the female’s bill was slender, long, and curved like that of a nectar-feeding bird. These differences suggest different feeding techniques: Both sexes fed on insects, but apparently the male chopped rotten bark from trees while the female probed crevices. Endemic to southern North Island, the last confirmed sighting was in 1907, although there were credible reports as late as the early 1960s. It appears to have required large tracts of primary, undisturbed forest. The North Island kokako (Callaeas wilsoni) is a type of wattlebird that still survives in patches of relict forests in the northern part of the North Island. Formerly it occurred throughout the island, but the clearing of forests and predation by opossums, ermines, and rats have led to its decline. The North Island piopio (Turnagra tanagra) was a thrushlike bird that was reportedly common in southern North Island during the 1870s. By the following decade it had become increasingly rare due to habitat destruction and predation by introduced species. The last confirmed record dates from 1902, although occasional purported sightings occurred up until 1970.

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The stitchbird (Notiomystis cincta) is a type of small passerine bird that was relatively common throughout North Island when Europeans arrived, but by 1890 had been extirpated there. Long confined to Little Barrier Island, where it is threatened by introduced cats, it has since been introduced to three other island sanctuaries and to two sites on the North Island mainland. The North Island saddleback (Philesturnus rufusater) is a type of passerine bird that was abundant and widespread on North Island at the time of European contact, and also occurred on a number of coastal islands. It declined rapidly to extirpation on the North Island mainland, however, following the introduction of predatory mammals, especially ship rats and stoats. By the early 1900s the species was confined to a single population on Hen Island (Taranga), off the northeastern coast. In the 1960s the New Zealand Wildlife Service began a series of translocations, and there are now about 15 populations on various coastal islands along with 5 at predator-fenced mainland sites. The long-footed bushwren (Xenicus longipes) was a small, almost flightless passerine bird that occurred in three nowextinct subspecies on each of the three larger islands of New Zealand. In each case introduced predators were the reason for the decline. The North Island long-footed bushwren (X. l. stokesi) was last reliably reported from the southern Rimutaka Range in 1918 and the Ureweras up to 1955, with additional probable sightings in 1911 on Kapiti Island, 1949 near Lake Waikareiti, and several times in the first half of the twentieth century within the Huiarau Range. Apparently, the last population lived in the area where Te Urewera National Park was established, just around the time of its extinction. Delcourt’s giant gecko (Hoplodactylus delcourti), the largest of all geckos, was known only from the eastern part of North Island. It is thought to have gone extinct around the mid-nineteenth century. Several skinks of the genus Oligosoma endemic to North Island have been decimated by loss of habitat and predation from introduced species. The robust skink (O. alani) is a large, rare species that was once widespread on North Island, but is now confined to six small islands off the north-eastern coast. Whitaker’s skink (O. whitakeri) is found on two small, predator-free islands off northern North Island (Middle Island and Castle Island). There is an additional mainland population at Pukerua Bay, near Wellington, and there are plans to translocate some of these to nearby Mana Island. Macgregor’s skink (O. macgregori) is confined to a small area of coastal north-western North Island and in the Cavalli Islands, and to Mana Island off the south-western coast. The striped skink (O. striatum) is found in a few scattered areas of western and north-western North Island and on Great Barrier and Little Barrier islands. The small-scaled skink (O. microlepis) occurs in small, isolated populations throughout central North Island, including one on Motutaiko Island in Lake Taupo.

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Archey’s frog (Leiopelma archeyi) is confined to a few areas of North Island, with the largest population occurring in the Coromandel Range. A smaller population is found in the Whareorino Forest of western North Island, from which a few individuals were translocated to Pureora Forest Park in central North Island in 2006. The Inanga galaxias (Galaxias gracilis) is a land-locked species confined to less than a dozen lakes on the northwestern coast of North Island. It was introduced into Lake Ototoa in the 1980s. It is threatened mainly by introduced fish species. The Northland mudfish (Neochanna heleios) and the black mudfish (N. diversus) are both confined to swampy areas and wetlands in far north-western North Island, where they are seriously threatened by habitat destruction and degradation as well as by introduced fish species. The Barrier Islands The Barrier Islands are located north-west of North Island. They include Great Barrier Island, Little Barrier Island, and a number of smaller islets. The chevron skink (Oligosoma homalonotum) is confined to Great and Little Barrier Islands, where it is threatened by introduced rats. Three Kings Islands The Three Kings Islands are a group of 13 uninhabited islands located about 55 km north-west of North Island. Falla’s skink (Oligosoma fallai) is confined to the Three Kings Islands. The Poor Knights Islands The Poor Knights Islands are located off the north-eastern coast of North Island. Buller’s shearwater (Ardenna bulleri) is a type of large seabird that breeds only on the Poor Knights Islands, specifically the two main islands and five small islets, although at other times it disperses widely throughout the Pacific Ocean. It is threatened mainly by fisheries by-catch, and potentially vulnerable to the accidental introduction of nest predators in its breeding colonies and to stochastic events.

South Island South Island (Te Waipounamu in Maori) is the larger of the two main islands of New Zealand. More mountainous and alpine than North Island, with a great chain of mountains more than 3000 m high (known as the Southern Alps) dominating the western and central areas, its wide range of habitats and smaller population has provided a haven of sorts for many species of birds exterminated from the latter. Prior to the arrival of Europeans it was still largely covered by temperate forests, of which only remnants now exist, along with areas of rainforest. The Westland petrel (Procellaria westlandica) is a large seabird that breeds only in the densely forested coastal foothills

The Novozelandic Region

near Punakaiki, South Island, where it is threatened by habitat degradation due to erosion and landslips. At other times it migrates as far as the eastern coast of Australia and the western coast of South America. The South Island swan (Cygnus sumnerensis sumnerensis) is thought to have been wiped out by the first Polynesian setters around 1450. The kea (Nestor notabilis) is a large, cold-adapted parrot from the forested and alpine regions of South Island. A known carrion-eater, it was once killed for bounty due to unfounded concerns by sheep-farmers that it attacked livestock. It has been fully protected since 1986, and today numbers between 3000 and 7000. The South Island takahe (Porphyrio hochstetteri), the world’s largest living rail, was thought to have been extinct since 1898 after the last four specimens were collected. In 1948, however, a small colony of 250–300 was unexpectedly discovered near Lake Te Anau in the Murchison Mountains of south-western South Island. The government immediately closed off the area covering the remaining habitat, which fortunately already lay within Fiordland National Park (New Zealand’s largest). Unfortunately, this protection has not been enough to ensure a stable recovery. By 1970 the population was estimated at around 400, but within a decade had been reduced to a fluctuating number of around 100–160 owing to competition with feral deer. The latter are now controlled, with varying degrees of success, by means of hunting from helicopters. The colony was devastated once again in 2007–08 by disease brought on by introduced stoats (Mustela erminea), reaching a low of just 80 individuals by 2014. Fortunately, a captive-rearing (later, captive breeding) programme had been initiated at a special facility near Lake Te Anau, which provided a means to replenish the wild stock. In addition, beginning in the mid-1980s populations had begun to be translocated to eight other, predator-free mainland sites and islands (Mana Island, Tiritiri Matangi, Cape Sanctuary, Maungatautari, Motutapu Island, Tawharanui, Rotoroa Island, and one undisclosed site). In 2016 the total population consisted of some 280 mature individuals, with approximately 87 breeding pairs. The western weka (Gallirallus australis australis) is a type of rail found mainly in the northern and western regions of South Island. Several kiwis (Apteryx) endemic to South Island have lost much of their former habitat and continue to be threatened by predation from introduced predators. The great spotted kiwi (A. haastii) is found patchily in western and north-western South Island, and is the only known kiwi species without any secure island populations. The southern brown kiwi (A. australis) is divided into two subspecies. The South Island brown kiwi (A. a. australis) is found patchily in south-western South Island. The Okarito brown kiwi (A. rowi) is confined to the Okarito Forest on the central-western coast of South Island, with further translocated populations on Mana, Motuara, and Blumine islands in the Cook Strait region.

Figure 8.8 Credit: South Island Takahe (Porphyrio hochstetteri) in the Karori Wildlife Sanctuary, © Pseudopanax.

The South Island piopio (Turnagra capensis) is a nowextinct type of passerine bird divided into two subspecies. The nominate form (T. c. capensis) was reported to be fairly common up until the 1860s, but declined very rapidly in the 1880s. The last definite record was from 1905. There have been unconfirmed reports since, although none later than 1963. The South Island long-footed bushwren (Xenicus longipes longipes) was last authentically recorded from Arthur’s Pass in 1966 and Nelson Lakes National Park in 1968. There have been a few unsubstantiated reports since then from Fiordland and Nelson Lakes, but the subspecies is nowadays generally considered extinct. The black-eyed gecko (Mokopirirakau kahutarae) is an alpine species first discovered in 1970, and now known from a few disjunct mountain peaks in northern South Island. Three skinks of the genus Oligosoma are threatened by introduced predators and habitat degradation. The scree skink (O. waimatense) occurs in a few rocky areas in central and northern South Island. The Otago skink (O. otagense) and the grand skink (O. grande) are each confined to a few scattered localities in south-central and south-eastern South Island. The Canterbury mudfish (Neochanna burrowsius) is confined to wetland fragments in central-eastern South Island, where it is considered to be one of New Zealand’s most threatened freshwater fishes. Several species of galaxias (Galaxias) are endemic to South Island, where they are threatened by loss of habitat and predation by introduced salmonids. The bignose galaxias (G. macronasus) is confined to a few localities within the Waitaki River catchment of south-central South Island. Eldon’s

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The Maud Island frog (Leiopelma pakeka) was historically confined to Maud Island, where at its lowest point it was confined to a single small area of remnant forest. In recent years populations have been successfully translocated to other parts of Maud Island as well as to nearby islands and to a protected area near Wellington. The Brothers The Brothers are a group of small islands in the Cook Strait comprised of two main islands and a number of islets. The Brothers tuatara (Sphenodon punctatus guntheri) occurs naturally only on North Brother Island, where the total population numbers around 400. Since the 1990s they have been successfully translocated to three other islands (Titi Island, Somes Island, and Long Island).

Stewart Island

Figure 8.9 A living South Island long-footed bushwren (Xenicus longipes longipes) taken in 1911. (Credit: Herbert Guthrie-Smith.)

galaxias (G. eldoni), one of the most range-restricted of all non-migratory galaxids, is confined to a few tributaries within the Taieri and Tokomairiro catchments in south-eastern coastal South Island. The dusky galaxias (G. pullus) and the central Otago roundhead galaxias (G. anomalus) are both confined to the Taieri and Clutha catchments in south-eastern South Island. The lowland longjaw galaxias (G. cobitinis) is confined to the Kauru River, a tributary of the Kakanui River, and to parts of the upper Waitaki catchment in south-central South Island. The flathead galaxias (G. depressiceps) occurs within the Taieri, Waikouaiti, Shag, Akatore, and Tokomairiro river catchments of south-eastern South Island. The upland longjaw galaxias (G. prognathus) is found patchily in mid- to high-altitude rivers and streams on the eastern slopes of the Southern Alps. Stephens Island Stephens Island (Takapourewa in Maori) lies at the northernmost tip of South Island and is home to several threatened species, including tuataras. The Stephens Island piopio (Turnagra capensis minor) was driven extinct by 1897 due to feral cat predation. Maud Island Maud Island (Te Hoiere in Maori) is located in the Marlborough Sounds at the northern tip of South Island. During the twentieth century much of the forest was cleared for farming, although today it serves as an important, predator-free nature reserve.

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Stewart Island (Rakiura in Maori) is located 30 km south of South Island. The third largest island of New Zealand, it is surrounded by three larger as well as numerous smaller islands. The Stewart Island weka (Gallirallus australis scotti) is confined to the Stewart Island archipelago, with an additional introduced population on Kapiti Island. The Stewart Island brown kiwi (Apteryx australis lawryi) is confined to Stewart Island. The Stewart Island long-footed bushwren (Xenicus longipes variabilis) was very likely the last surviving of the three subspecies of long-footed bushwren. Historically found on Stewart Island and a few nearby islets, it is known to have survived on the former until 1951 but was exterminated soon after by feral cats. It survived on Solomon Island until the early 1960s, and on Big South Cape Island until the rat invasion of 1964. The New Zealand Wildlife Service attempted to save it by relocating all the birds that they could capture (six in total) to rat-free Kaimohu Island, but the translocation unfortunately failed. The last one died in 1972. The southern skink (Oligosoma notosaurus) is confined to Stewart and Codfish islands, with an additional population on Betsey Island. Codfish Island Codfish Island (Whenua Hou in Maori) is located to the west of Stewart Island. Following the eradication of brushtail possums (Trichosurus vulpecula) and rats in 1998, it has served mainly as a predator-free bird sanctuary and, in particular, the focus of kakapo (Strigops habroptila) recovery efforts. The Codfish Island fernbird (Poodytes punctatus wilsoni) is confined to this one small island, where it is sensitive to disturbances.

The Chatham Islands The Chatham Islands, about 680 km south-east of New Zealand, are the largest of the island groups belonging to the

The Novozelandic Region

continental shelf of New Zealand. They consist of about 10 islands within a 40-km radius, the largest of which are Chatham Island and Pitt Island. The latter are the only inhabited islands, the remaining smaller islands being conservation reserves with restricted or prohibited access. Several birds have already vanished from these islands and quite an assemblage of species is obviously going the same way. All the factors that have contributed to the extermination of birds on New Zealand have also operated on the Chatham Islands; Europeans introduced rabbits, goats, and cats, and destroyed the native vegetation. In addition, commercial collectors of rare birds have been at work on the islands, driving a number of species extinct. The Chatham Island swan (Cygnus sumnerensis chathamensis) is known only from fossils, and is believed to have been driven extinct by Polynesian settlers around 1650. The Chatham penguin is known only from subfossil bones, but may have become extinct as recently as the late nineteenth century, as a bird kept captive at sometime between 1867 and 1872 might refer to this taxon. It appears to have been a distinct species (‘Eudyptes chathamensis’), with a thin, slim, and low bill, but at time of writing has yet to be formally described. The Chatham albatross (Thalassarche eremita) breeds only on The Pyramid, a 1.7-ha stack in the Chatham Islands. Two cormorants of the genus Phalacrocorax are endemic to the Chatham Islands. Onslow’s shag (P. onslowi) is confined to Chatham, Star Keys, Rabbit, and Pitt islands. Featherston’s shag (P. featherstoni) is confined to Chatham, Pitt, Mangere, Little Mangere, South East, Star Keys, the Pyramid, Big and Middle Sister, Murumurus, the Castle, and Rabbit islands. The Magenta petrel (Pterodroma magentae) is one of the world’s rarest birds. First collected at sea in 1867, midway between New Zealand and South America by the Italian ship Magenta, it was not recorded again for another 111 years and was feared extinct. However, in 1978 it was rediscovered on Chatham Island, where it took another 10 years to locate one of its breeding burrows within a forested valley. In 2012 the total population was thought to be around 150–200. The Chatham petrel (P. axillaris) breeds only on South East Island and two predator-protected sites on Chatham and Pitt islands. Hawkins’ rail (Diaphorapteryx hawkinsi) was known from Chatham and Pitt islands. It was exterminated by hunting sometime after 1895. Dieffenbach’s banded rail (Hypotaenidia dieffenbachii) occurred on Chatham, Mangere, and Pitt islands. Already rare when the type specimen was collected in 1840, it was extinct by 1872. The Chatham rail (Cabalus modestus) occurred on Chatham and Mangare islands. It went extinct between 1893 and 1895. The Chatham snipe (Coenocorypha pusilla) is confined to South East, Star Keys, Mangere, and Little Mangere.

The Chatham oystercatcher (Haematopus chathamensis) is confined to South East, Pitt, Mangere, and a few smaller islands and stacks. The Chatham parakeet (Cyanoramphus forbesi) is confined to Mangere and Little Mangere islands. The Chatham pigeon (Hemiphaga chathamensis) was formerly found on Mangere and Pitt islands as well as on Chatham Island, but is now confined to the latter. The Chatham black robin (Petroica traversi) had, by 1980, the smallest population of any bird for which precise figures are known (two males and three females), and seemed doomed to extinction. A last-ditch conservation effort succeeded in saving the species, although it remains rare and restricted to South East and Mangere islands. All surviving individuals are the descendants of a single breeding pair. The Chatham fernbird (Poodytes rufescens) occurred on Pitt and Mangare islands. It is thought to have gone extinct around 1892 when the last specimen was collected. The Chatham bellbird (Anthornis melanocephala) occurred on Chatham, Mangere, and Little Mangere islands. It was last seen on Little Mangere Island in 1906, and a search for it in 1936 was unsuccessful. The Chatham tui (Prosthemadera novaeseelandiae chathamensis) is a type of passerine bird confined to the Chatham Islands.

Novozelandic Sub-Antarctic Islands The sub-Antarctic islands of the South Pacific are chiefly comprised of six groups south of New Zealand that have been collectively designated a UNESCO World Heritage Site. Most lie near the south-eastern edge of the largely submerged continent centered on New Zealand known as Zealandia, which was riven from Antarctica between 130 million and 85 million years ago, and from Australia 85–60 million years ago. Until 1995 scientific research staff were stationed permanently at a meteorological station on Campbell Island. Since then, all of the islands have been uninhabited, although they continue to be periodically visited by researchers and tourists. The southern royal albatross (Diomedea epomophora), with its more than 3-m wingspan, is one of the largest of all albatrosses. The vast majority of the world’s population nest on rat-free Campbell Island (around 8200–8600 pairs), although there are a few minor colonies within the Auckland Islands. During the non-breeding season the birds disperse throughout the southern oceans, where they are vulnerable to fisheries by-catch. The Antipodean albatross (D. antipodensis) is divided into two subspecies. The nominate form (D. a. antipodensis) breeds on the Antipodes Islands, Campbell Island, and marginally on Pitt Island in the Chatham Islands. Salvin’s albatross (Thalassarche salvini) breeds colonially within the Crozet Islands of the southern Indian Ocean and in a few islands and a number of islets in sub-Antarctic New Zealand (the Bounty Islands, the Chatham Islands, and The Snares). At other times the birds disperse as far as South Africa

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and the western coast of South America, where they are threatened mainly by fisheries by-catch. The erect-crested penguin (Eudyptes sclateri) now breeds only on the Antipodes and Bounty islands. During the midtwentieth century the species was recorded also breeding on the New Zealand mainland, although these were individual pairs that were not part of a larger population. Outside the breeding period the birds have been reported throughout the coastal areas of New Zealand, southern Australia, and even as far as the Kerguelen Islands and Falkland Islands. They are mainly threatened by fisheries by-catch.

The Snares The Snares is a small island group located about 200 km south of Stewart Island. It consists of the main North East Island and the smaller Broughton Island, as well as the Western Chain Islands some 5 km away. All are biologically important and have been minimally impacted by humans, with no introduced predators. The Snares crested penguin (Eudyptes robustus) breeds only on The Snares, but ranges at other times as far away as southern Australia and Argentina. The Snares snipe (Coenocorypha huegeli) was historically confined to The Snares. In recent years translocated colonies have also been established on Putauhinu Island on the coast of Stewart Island and on Codfish Island. The Snares fernbird (Poodytes caudatus) is confined to the forests of North East Island. The Snares tomtit (Petroica dannefaerdi) is confined to The Snares in areas of tussock grassland.

thick forests of southern rata (Metrosideros umbellata). Despite the strong, almost constant west winds, domestic animals were introduced. In 1807 pigs were released, in 1850 cattle and goats, and in 1900 sheep. Rabbits were also introduced. The goats were particularly destructive to the vegetation, and the pigs, cats, and dogs have had a disastrous effect on groundnesting birds, which were generally trusting and unafraid, as they had never before met mammalian predators. The Auckland albatross (Diomedea antipodensis gibsoni) breeds only on the Auckland Islands (Adams, Disappointment, and Auckland). The Auckland shag (Leucocarbo colensoi) is confined to the Auckland Islands and adjacent waters, where colonies are present on Auckland, Enderby, Rose, Ewing, and Adams Islands. The Auckland rail (Lewinia muelleri) was thought to have been exterminated in 1865 by introduced domestic animals. However, in 1966 it was rediscovered on Adams Island, and in 1993 on Disappointment Island. The total population is around 2000. The Auckland teal (Anas aucklandica) was exterminated on Auckland Island but still survives on Ewing, Enderby, Rose, Ocean, Adams, Disappointment, and Dundas islands.

The Campbell Island Group

Located in sub-Antarctic waters about 860 km south-east of Stewart Island, the volcanic Antipodes Islands consist of a main island surrounded by a series of offshore islets and stacks. The Antipodes parakeet (Cyanoramphus unicolor) is common on the main island and Bollons Island, and occurs in small numbers on Leeward, Inner Leeward, and Archway islets. The population has long held steady at between 2000 and 3000 birds, but the species remains vulnerable should rats come ashore. Hochstetter’s parakeet (C. hochstetteri) is similarly confined to the Antipodes Islands.

About 235 km south-east of the Auckland Islands lies mountainous, uninhabited Campbell Island, together with a number of smaller satellite islets. The main island’s rugged cliffs and inhospitable and harsh climate did not prevent man from introducing sheep and cattle there, which greatly modified the vegetation. Fortunately, the island was declared a nature reserve in 1954. The domesticated animals have long since been removed, and rats eliminated, all of which has led to a recovery of native species. The Campbell albatross (Thalassarche impavida) breeds only on the northern and western coasts of Campbell Island and on the nearby offshore islet of Jeanette Marie. Numbers appear to be increasing, but the species is still highly vulnerable. The Campbell shag (Leucocarbo campbelli) is confined to Campbell Island and adjacent offshore islands and stacks. The Campbell flightless teal (Anas nesiotis) is one of the world’s rarest ducks. Between 1866 and 1944 it was collected only three times and was long feared extinct until its breeding grounds on Dent Island were discovered in 1975. Since then, a successful captive-breeding programme has allowed for the translocation of individuals first to Codfish Island and later to Campbell Island itself. The total population is now in excess of 200. The Campbell snipe (Coenocorypha aucklandica perseverance) was only just discovered by chance in 1997 on Jacquemart Island. It has since recolonized Campbell Island following the eradication of rats, where it is thriving.

The Auckland Islands

Macquarie Island

The Auckland Islands lie 360 km south of Stewart Island. Tussock grasses dominate the vegetation, but there are also

Macquarie Island, almost halfway between New Zealand and Antarctica (although belonging to Australia), is treeless, with

The Bounty Islands The Bounty Islands are a small group of 13 uninhabited granite islets and numerous rocks. They lie about 670 km eastsouth-east of the South Island of New Zealand, and 530 km south-west of the Chatham Islands. The Bounty shag (Leucocarbo ranfurlyi) is a type of cormorant confined to narrow cliffside ledges. The total population is believed to be less than 1000.

The Antipodes Islands

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tussock grasses and mosses forming vegetation. It nevertheless serves as a vitally important breeding area for seabirds and marine mammals. The ecology of the island was affected soon after the beginning of European visits in 1810. The fur seals, elephant seals, and penguins were killed for their fur and blubber, and rats and mice were inadvertently introduced from the ships. Later, cats were deliberately introduced as well in an effort to control the rodents. Then, in about 1870, rabbits were left on the island by sealers to breed for food, which caused tremendous damage. Beginning in the late twentieth century efforts were made to eliminate the feral cats, which had had a devastating impact on the native seabirds in particular, although rodents and rabbits remained a problem until April 2014, when the island was officially declared pest-free. The royal penguin (Eudyptes schlegeli) is only known for certain to breed on Macquarie Island and nearby Bishop and Clerk islets. However, small numbers of similar-looking birds occasionally appear on other sub-Antarctic islands (such as South Georgia and Kerguelen Island), perhaps indicating that they may breed elsewhere. The species was historically hunted for its oil, with an average of 150,000 birds (both royal and king penguins) being taken each year. At the peak of the industry in 1905 the plant established on Macquarie Island was processing 2000 at a time. When the hunt was finally ended the species gradually recovered its numbers. While not currently directly threatened it remains vulnerable to stochastic events. The Macquarie shag (Leucocarbo purpurascens) is a type of marine cormorant that breeds only on Macquarie Island and nearby Bishop and Clerk islets. The Macquarie buff-banded rail (Gallirallus philippensis macquariensis) was said to be common until 1880, but was gone by 1894. The Macquarie red-fronted parakeet (Cyanoramphus erythrotis) was exterminated by 1913.

Balance for the Novozelandic Region New Zealand was one of the last major landmasses to be settled by humans. Radiocarbon dating, evidence of deforestation, and mtDNA variability within Maori populations suggest that Polynesians first arrived on these islands between A . D . 1250 and A . D . 1300. This represented the culmination of a long series of voyages through the Pacific. At some point during the following centuries a group of these settlers migrated to the Chatham Islands. Europeans had known of New Zealand since the mid-seventeenth century, when the Dutch explorer Abel Tasman first ‘discovered’ the main islands. They would remain little visited over the next two centuries. During 1769–70 the English explorer James Cook circumnavigated North Island and South Island, mapping almost the entire coastline and proving they were not part of a larger ‘Terra Australis Incognita’. Following Cook, New Zealand was visited by numerous European and North American whaling, sealing, and trading ships, greatly transforming Maori culture in the process. In 1791–95 George

Vancouver and his expedition discovered the Chatham Islands and The Snares. In 1806 the British mariner Abraham Bristow reached the Auckland islands, and in 1810 the Australian sealer Frederick Hasselborough visited the Campbell and Macquarie islands. From the early nineteenth century on Christian missionaries began to settle in New Zealand, which had by then become a British colony. They would eventually convert most of the native population that, by then, had begun to decline dramatically due to introduced diseases and armed conflict. As discussed previously, prior to the beginning of European colonization a number of species had already been driven to extinction by early Polynesians. Yet New Zealand’s forests, at least, remained mostly intact. It was after the arrival of the Europeans, however, that an arguably even greater destruction of New Zealand’s flora and fauna than anything done previously occurred. Indeed, the negative impact of the European settlers on the region in just two centuries is comparable to what was inflicted upon Australia, North America, and the Caribbean over the same period. Not only were pigs, cattle, goats, and other domestic animals brought in to provide meat but also wild mammals, such as deer and rabbits, for hunting and sport. One mistake led to another. Several species soon began destroying grain, forest, and native animals. Not less than 53 exotic mammals have been introduced, of which 34 are still there. When the rabbits, introduced before 1838, became too abundant, predators such as ermines, polecats, and weasels were brought in. These quickly established themselves, and like rats and dogs, turned on the native birds, reptiles, and amphibians. Another serious mistake was the deliberate introduction of the common brushtail possum (Trichosurus vulpecula) in 1858 as a valuable fur-bearing animal. In New Zealand unlike Australia, this creature, which is herbivorous, caused tremendous destruction in forests and orchards. It increased rapidly and soon the New Zealanders declared a war of extermination against it, killing over 900,000 in 1945 alone. Above all, it is the tens of millions of sheep that have changed the islands beyond recognition, causing serious problems with erosion in the process. Added to this the aforementioned destruction of the forests, the burning of scrub, and the drainage of swamps has led to numerous species and subspecies becoming extinct during the last two centuries alone. Some may yet be wiped out in the near future. Today only a very small amount of mostly planted forest remains in New Zealand, with only remnants of the originals here and there, mainly in the mountains. Elsewhere there are only pastures and agricultural lands. It is no exaggeration to say that the consequences for native plants and animals have been a catastrophe. Moreover, these surviving scraps of wilderness have been greatly harmed by the numerous exotic as well as domestic species introduced over the past thousand years. Thus, the original nature of New Zealand has been destroyed forever, and has been replaced by a composite nature from all continents.

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Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

~ species

1 species

~ species

3 species

~ subspecies

~ subspecies

~ subspecies

2 subspecies

~ taxa

1 taxon

~ taxa

5 taxa

23 species

~ species

~ species

65 species

6 subspecies

~ subspecies

~ subspecies

1 subspecies

29 taxa

~ taxa

~ taxa

76 taxa

1 species

~ species

~ species

14 species

~ subspecies

~ subspecies

~ subspecies

1 subspecies

1 taxon

~ taxa

~ taxa

15 taxa

~ species

~ species

~ species

3 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

~ taxa

~ taxa

3 taxa

1 species

~ species

~ species

17 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

1 taxon

~ taxa

~ taxa

17 taxa

25 species

1 species

~ species

102 species

6 subspecies

~ subspecies

~ subspecies

14 subspecies

31 taxa

1 taxon

~ taxa

116 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

In recent historical time (i.e. since A . D . 1500), the Novozelandic Region has lost at least 25 species/6 subspecies of vertebrates. Among the extinct forms 23 species/6 subspecies are birds, 1 species is a reptile, and 1 species is a freshwater fish. One other species is possibly extinct. In addition, there are 102 species/14 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 3 species/2 subspecies are mammals, 65 species/11 subspecies are birds, 14 species/1 subspecies are reptiles, 3 species are amphibians, and 17 species are freshwater fishes.

The Polynesian Region The Polynesian Region comprises the largest section of the Polynesian Realm. It extends from the Hawaiian Islands in the north to the Tonga and Cook islands in the south-west and Pitcairn and Easter islands to the south-east, and encompassing all of the islands of the central Pacific. The oceanic parrot (Eclectus infectus) is known from bones found on three islands in the Tonga Archipelago, and which presumably relate to a drawing of a parrot made by an expedition in 1793. It may have occurred on Vanuatu and Fiji as well.

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The blue lorikeet (Vini peruviana) was historically endemic to some two dozen islands around Tahiti in the Society Islands but is now confined to about eight (Motu, Manuae, Tikehau, Rangiroa, Aratua, Kaukura, Apataki, and possibly Maupihaa, Harvey Island, and Manihi). The species is also present on the northern atolls of the Tuamotu Archipelago, and has been introduced to the island of Aitutaki in the Cook Islands. It is threatened mainly by introduced rats and feral cats. The Polynesian imperial pigeon (Ducula aurorae) was historically found on Tahiti in the Society Islands and on Makatea in the Tuamotu Archipelago. It is thought to have been extirpated from the former island, where it was long confined to two valleys where there have been no reports for many years. On Makatea the population was estimated at around 1200 in 2009. The Polynesian ground dove (Pampusana erythropterus) is divided into two subspecies. The nominate form (P. e. erythropterus) was historically common throughout the Society Islands, eastern Tuamotu and, to judge by fossil material, the Cook Islands as well. It is now extirpated from the Society Islands and most of the Tuamotus, and likely survives only in the remote Acteon Islands group, where the total population is thought to be around 200. Stair’s ground dove (P. stairi) is found discontinuously in the Samoan, Fiji, Tonga,

The Polynesian Region

and Wallis and Futuna islands. All are seriously threatened by loss of habitat and introduced rats and feral cats. The barred emo skink (Emoia trossula) is known from around 20 small and medium-sized islands in the Fiji, Cook, and Tonga islands. The olive small-scaled emo skink (E. lawesi) occurs in the Samoan and Tonga islands as well as Niue. Steindachner’s emo skink (E. adspersa) is found sporadically in the Samoan, Tonga, Cook, and Wallis and Futuna Islands as well as from Tuvalu, where it may perhaps be extirpated. All are threatened by habitat destruction and predation from introduced species. The Samoan stiphodon (Stiphodon hydroreibatus) is a type of freshwater goby known only from a small number of specimens collected from the Samoan Islands and Futuna Islands.

The Hawaiian Islands The Hawaiian Islands (Mokupuni o Hawai’i in Hawaiian) are an archipelago of eight major islands, several atolls, and numerous smaller islets in the northern Pacific Ocean, extending some 2400 km from the island of Hawaii in the south to northernmost Kure Atoll. The islands are actually the tops of what is perhaps the world’s mightiest mountain chain. The depths around them are very impressive, the average being just under 5500 m. If measured from its base at the bottom of the sea Mauna Kea, a volcano on Hawaii, is the highest mountain in the world. The islands are geologically rather young and there have been many volcanic eruptions that, several times, probably exterminated all the plants and animals that had managed to reach them. Gradually, beginning in the northwest, volcanic activity died down. Thus Oahu, on which Honolulu is located, has probably not had an eruption since the Polynesian first discovered the islands around A . D . 500, but Hawaii, the youngest island in the group, has volcanoes such as Mauna Loa, the largest active volcano in the world. The islands are interesting as well for their unique biodiversity. Long isolation has created species found nowhere else, and as an illustration of the process of evolution these islands are as important as the Galápagos. It is therefore tragic that so many endemic species have been exterminated, and that so many others remain threatened. What remains of the natural biodiversity is among the most magnificent on Earth. The great volcanoes on Maui and Hawaii rise from a humid, tropical area into a cold, dry alpine zone. It is a mystery how the alpine plants reached the islands. Moreover, none of the alpine plants on the volcanoes of Hawaii exist elsewhere and their relations to other alpine species are so distant that their origin cannot be traced with certainty. About 90 per cent of all the native plants on the Hawaiian Islands are endemic. Many birds in these islands are also not found anywhere else. One of the most beautiful examples of the evolution of species and the colonization of hitherto unoccupied environments is provided by the Hawaiian honeycreepers (Mohoidae). All are probably descended from one species, perhaps even only one pair, that once upon a time reached Hawaii from North America. Before the arrival of humans the only vertebrates that had reached

Hawaii were birds and a bat. Nearly all these immigrants came from North America, except three, two of which can be traced to Australia and Polynesia, while the origin of the third, a rail, is unknown. Unfortunately, this unique fauna has suffered shockingly during the past two centuries. Cultivation, the destruction of forests, disease, and the introduction of a large number of exotic plants and animals was a catastrophe for the native fauna. Domestic mammals such as goats, sheep, cattle, pigs, cats, and dogs as well as rats have seriously damaged vegetation and preyed directly on native birds. In addition, Europeans brought the small Indian mongoose (Herpestes auropunctatus) to Hawaii in 1883, making the same terrible mistake as they had made elsewhere in the world. Rabbits, axis deer (Axis axis), and even mouflons (Ovis gmelini) were also introduced, the latter as late as 1954. The mongoose occurs on all major islands except Kauai, Lanai, and Niihau, the axis deer on Molokai, Oahu, and Lanai, and the mouflon on Lanai alone. However, feral goats are abundant on most islands and destroy much vegetation. Likewise, feral pigs constitute a threat to groundnesting birds. The number of Hawaiian birds that have become extinct or are threatened is thus tragically high, particularly because almost all of them represented various stages of evolution and scientifically made the Hawaiian Islands a living museum. Thirty species/3 subspecies have been exterminated and 31 species/5 subspecies are threatened. Indeed, no other comparable area of the world shows such a negative faunal balance as Hawaii. Most of the vanished species are land birds, which could not survive on volcanic islands once the forest cover was destroyed. Sea birds generally survived on the coral limestone atolls. The Hawaiian monk seal (Neomonachus schauinslandi) was ruthlessly slaughtered by sealers, whalers, plumage hunters, and guano diggers during the latter half of the nineteenth century, and was nearly driven extinct. Complete protection saved them, and from their few remaining retreats on the more remote islands and atolls the species is now once again found throughout the Hawaiian Islands, with occasional sightings outside of the main range (Johnston Atoll, Palmyra Atoll, and Wake Island). Nevertheless, the total population (less than 1000) remains small. The Hawaiian hoary bat (Lasiurus cinereus semotus) was formerly found on the islands of Hawaii, Molokai, Maui, Oahu, and Kauai, but appears to be confined now to Hawaii and Kauai. The Hawaiian dark-rumped petrel (Pterodroma sandwichensis) ranges across the central Pacific but breeds only in the Hawaiian Islands. Long harvested by early Polynesians, it may have already been restricted to its current breeding range (mainly Haleakala Crater on Maui, with smaller colonies on Mauna Loa, Hawaii, Waimea Canyon, Kauai, and on Lanai and possibly Molokai) when Europeans arrived. The total population possibly exceeds 10,000, but is threatened by loss of habitat and introduced predators. Newell’s shearwater (Puffinus newelli) is a type of seabird that nests principally on the mountains of Kauai, with smaller

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additional colonies on Molokai, Hawaii, and possibly Maui, Oahu, and Lanai. The Hawaiian black-necked stilt (Himantopus mexicanus knudseni) is a long-legged, slender shorebird that still occurs locally on all the main islands, but is threatened by the ongoing building boom in the lowlands. The total population is thought to be around 2000. The Hawaiian hawk (Buteo solitarius) breeds only on Hawaii, although vagrants are occasionally seen on Molokai, Oahu, and Kauai. By the mid-twentieth century it had been reduced to fewer than 200 birds as a result of indiscriminate shooting. Legal protection helped to increase its number to over 1000, where it has remained stable. The Hawaiian short-eared owl or pueo (Asio flammeus sandwichensis) is found throughout the Hawaiian Islands but is everywhere decreasing, particularly on the island of ,Oahu where it was formerly common. The birds are strongly affected by light pollution and are often killed in vehicular accidents in which they dive toward the headlights of cars, possibly in an attempt to hunt. The Hawaiian goose or nene (Branta sandvicensis) historically occurred on all the main Hawaiian Islands. From a population of about 25,000 in the late nineteenth century it was reduced almost to extinction by hunting, introduced predators, and habitat destruction. By the mid-twentieth century, when it became the focus on an international conservation effort, perhaps as few as 30 birds remained. At the forefront of this campaign was Sir Peter Scott’s successful captive breeding programme at the Wildfowl Trust in Slimbridge, England. Since 1960 over 2400 geese, raised in England and at other facilities in the United States, have been reintroduced into protected areas on the islands of Hawaii, Maui, Molokai, and Kauai. In 2011 the total world population was in excess of 2500. Unfortunately, the mortality rate

remains high (particularly in times of drought), and the majority of birds outside of Kauai do not breed in the wild state. The Hawaiian duck or koloa (Anas wyvilliana) was formerly found on all of the main Hawaiian Islands except for Lanai and Kahoolawe. A significant population decline in the early twentieth century brought on by introduced predators, habitat destruction, and hunting reduced it to Kauai and Niihau. It has since been reintroduced to wetland areas on Oahu, Hawaii, and Maui, but continues to be seriously threatened by hybridization with feral mallard ducks (A. platyrhynchos). The total population in 2007 was about 2200. The Laysan duck (A. laysanensis) was also formerly widespread within the Hawaiian Islands, but eventually became confined to Laysan. The species was near extinction at the beginning of the twentieth century, with just 20 birds left in 1923 and about 33 in 1950. After rabbits were eliminated from the island, however, the population increased to around 500 by 1987. With the population approaching maximum carrying capacity, 42 were translocated to two islands of Midway Atoll National Wildlife Refuge in 2004 and 2005. A tsunami in March 2011 hit both the Laysan and Midway populations hard, but both have since rebounded. As an additional safeguard, 28 individuals were translocated from Midway to Kure Atoll, a predator-free island about 90 km to the north-west, in 2014. The Laysan rail (Zapornia palmeri) was confined to the north-western Hawaiian Islands, where it inhabited grass tussocks and thickets. Birds were introduced to Pearl and Hermes Reef and to Eastern Island in the Midway Atoll in 1891 and 1913 and from there to Sand Island in 1910, with failed reintroductions reportedly also attempted at Lisianski Island, Laysan, and the main Hawaiian group. However, it became extinct on Laysan between 1923 and 1936, on Pearl and Hermes Reef in 1930, and on Sand Island in 1943. The last population, on Eastern Island, was exterminated by introduced rats in 1944. Figure 8.10 Laysan rails by John Gerrard Keulemans. (Credit: John Gerrard Keulemans.)

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The Hawaiian gallinule (Gallinula galeata sandvicensis) is a chicken-sized wading bird originally found on most of the main Hawaiian Islands. Population numbers and range declined during the nineteenth century and first half of the twentieth century, mainly due to hunting, introduced species, and the draining of the lowland wetlands it favours. It was last reported on Hawaii in 1887 (subsequent attempts to reintroduce the birds in the late 1920s and again in the 1950s failed), and disappeared from Molokai sometime after the 1940s (a 1983 reintroduction was also unsuccessful, with five of the six birds being shot for food). At the species’ lowest point in the 1950s only 57 remained on Kauai and Oahu. With better protection the numbers have steadily increased since and, while still small, appear to be stable. The Hawaiian coot (Fulica alai) had declined dramatically during the twentieth century but is still to be found on all the main islands, where the total population fluctuates between 2000 and 4000. Drainage of wetlands for cultivation and developments remain a threat. The olomao (Myadestes lanaiensis) is a type of thrush divided into two subspecies. The Molokai olomao (M. l. rutha) was historically found on both Maui and Molokai. It was last recorded on the former island in 1933 and thought to be extinct on Molokai as well. In 1968, however, it was found to still exist in a single unaltered forest, where it was very rare. Nevertheless, the last well-documented record was in 1994. The Nihoa finch (Telespiza ultima) once occurred on Molokai and perhaps other islands, but has long been confined to Nihoa. The population is estimated at around 4500. The ou (Psittirostra psittacea) was a chunky, finch-like honeycreeper historically found throughout all the larger Hawaiian Islands except Oahu until the 1890s, although it declined precipitously thereafter. The last recorded sighting was in 1989 on Kauai, where it is now almost certainly extinct, although there are still occasional unconfirmed reports from the Kilauea Volcano area of Hawaii island. Bishop’s oo (Moho bishopi) was extinct on Molokai by 1915 as a result of commercial exploitation of the bird’s beautiful plumage, decimation by rats, and destruction of forests. The species was believed to live on Maui as well, where a single bird was observed in 1981 on the north-east slope of Haleakala. The ula-ai-hawane (Ciridops anna) was a small honeycreeper known only from five specimens and last definitely recorded in 1892, although with a possible sighting as late as 1937. Fossils indicate that it (or related species) also occurred at one time on Kaui, Molokai, and Oahu. The Maui Nui akialoa (Akialoa lanaiensis) is known only from three specimens collected on Lanai in 1892. Fossil material perhaps relatable to this species has been found on Maui and Molokai. The crested honeycreeper or akohekohe (Palmeria dolei) was extirpated on Molokai after 1907, and is now confined to the north-eastern slopes of Haleakala on Maui, where it is extremely rare, although stable.

The black mamo (Drepanis funerea) is a type of honeycreeper that was only ever observed on Molokai, although fossils are known from adjacent Maui. Last collected in 1907, intensive searches in the subsequent decades found no sign of it. The iiwi (D. coccinea) once occurred on all the main islands, but is now extinct on Lanai. Only relict populations remain on Oahu and Molokai. The Hawaiian spinecheek sleeper goby (Eleotris sandwicensis) is confined to lowland streams of the larger Hawaiian Islands. The Hawaiian freshwater goby (Lentipes concolor) is a little-known species from the mountain streams of Hawaii, Molokai, and Maui. It is important to native people as a food fish.

Hawaii Known as ‘the Big Island’, Hawaii (Hawai’i in Hawaiian) is the largest and south-easternmost of the chain. It is also the youngest, having been built from five separate shield volcanoes. The Hawaii rail (Zapornia sandwichensis) was apparently confined to upland forest clearings on the eastern side of Hawaii Island, although it may formerly have occurred on Molokai as well. Last seen in 1884 and possibly in 1893, it was exterminated by introduced cats. The Hawaii crow (Corvus hawaiiensis) is confined to the mountain slopes of Mauna Loa and Hualalai on Hawaii Island, where it was still reported to be common in 1891. Highly vulnerable to introduced rats, mongooses, and feral cats, by the 1970s the population had been reduced to less than two dozen birds living in protected areas. The last two known wild individuals disappeared from Mauna Loa in 2002, rendering the species extinct in the wild. Over 100 remain in captive breeding facilities, however, and a reintroduction plan is being developed. The Hawaii thrush or omao (Myadestes obscurus) is a robin-like bird formerly found throughout the island, but now restricted to rainforests on the southern and eastern slopes. The Hawaii akialoa (Akialoa obscura) was a type of finch that was not uncommon until 1895, but which declined rapidly thereafter. Last reported in 1940, it appears to have been driven to extinction by habitat destruction. The Hawaii elepaio (Chasiempis sandwichensis) is a type of small flycatcher confined to montane areas of Hawaii. Three subspecies are recognized. The Kona elepaio (C. s. sandwichensis) inhabits mesic forest, where its population appears to be stable at around 60,000–65,000. The volcano elepaio (C. s. ridgewayi) occurs in montane rainforest. The Mauna Kea elepaio (C. s. bryani) is confined to dry forests on the leeward slopes of Mauna Kea. The total population is between 2000 and 2500. The Hawaii creeper or alawi (Manucerthia mana) is confined to three montane dry forest areas of Hawaii.

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The Hawaii akepa (Loxops coccineus) is a small passerine bird confined to three montane forest areas on Hawaii, specifically in Hakalau Forest National Wildlife Reserve on Mauna Kea, the upper forest areas of Kau in the southern part of the island, and on the northern slope of Hualalai. The total population is thought to be around 14,000. The koa finches (Rhodacanthis) were large, seed-eating honeycreepers, all of which are now extinct. The lesser koa finch (R. flaviceps) is known only from a handful of skins, and was last recorded in 1891. The greater koa finch (R. palmeri) was still common during the late nineteenth century, but was last collected in 1896. The Hawaii mamo (Drepanis pacifica) was a type of honeycreeper last recorded in 1898 above Hilo and in 1899 nea Kaunana. The greater amakihi (Viridonia sagittirostris) was a type of honeycreeper found only along the Wailuku River above Hilo, where it became extinct in 1901 after most of its habitat was converted into agriculture.

Maui

Figure 8.11 Hawaii oo by John Gerrard Keulemans. (Credit: John Gerrard Keulemans.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The Hawaii oo (Moho nobilis) was a type of honeycreeper that was frequently shot for its beautiful feathers, which were used by native people in making robes. As late as 1898 hunters were still able to slaughter over 1000, but after that the population declined rapidly. The last known sighting was in 1934 on the slopes of Mauna Loa. The kioea (Chaetoptila angustipluma) was a type of large honeycreeper historically confined to Hawaii Island, although fossil remains are also known from Oahu and Maui. Only four specimens were ever collected. Last seen in 1859, it was presumably exterminated by logging of its montane plateau forest habitat. The palila (Loxioides bailleui) is a finch-billed type of honeycreeper formerly found throughout the Hawaiian Islands, but now confined to the forests of the upper slopes of Mauna Kea. In 2016 the total population was estimated at 1900. The akiapolaau (Hemignathus wilsoni) is a type of honeycreeper endemic to Hawaii Island, where it was formerly widespread in old growth and wet forests. Surveys in the late 1970s and early 1980s estimated around 1500 birds, although many populations have since been extirpated, mainly due to habitat destruction and introduced predators.

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The second largest of the Hawaiian Islands, Maui’s diverse landscapes are the result of a unique combination of geology, topography, and climate. It is home to a large, still relatively pristine rainforest on the north-eastern flanks of Haleakala Volcano. However, the vibrant dry forest on the island’s leeward side has been destroyed by human activities, while agricultural and industrial land use has had an adverse effect on most of the coastal regions. The Maui akepa (Loxops ochraceus) was last observed in 1988. It is likely extinct, although possible audio evidence gives hope that the species may still exist on Haleakala. The Maui nukupuu (Hemignathus affinis) is (or was) a type of honeycreeper confined to the eastern and north-eastern slopes of Haleakala. Considered extinct since 1896, it was rediscovered with three or four individuals in 1967. The last confirmed sighting was in 1989, although there have been other possible records since then. The black-faced honeycreeper or po’o-uli (Melamprosops phaeosoma) was first discovered in 1973 in the Ko’olau Forest Reserve on the north-eastern flanks of Haleakala. At the time it was estimated to number fewer than 200 birds. By 1995 only 5–7 individuals were left and by 1997 only 3 could be found (2 male and 1 possible female). Two of these were not seen again after 2003, and the following year the remaining individual was captured but died soon after. The Maui parrotbill (Pseudonestor xanthophrys) is a type of honeycreeper confined to a small forested area on the northeastern slopes of Haleakala. The population of around 500 birds is fully protected and has remained more or less stable since the 1970s. The Maui alauahio (Paroreomyza montana) is divided into two subspecies. The nominate form (P. m. newtoni) is found only in two areas of eastern Maui, where it is vulnerable to habitat degradation by feral goats and fires.

The Polynesian Region

Oahu Oahu (O’ahu in Hawaiian) is the third largest of the Hawaiian Islands. It is comprised of two separate shield volcanoes, with a broad valley or ‘saddle’ between them. The Oahu thrush or amaui (Myadestes woahensis) was a type of thrush known only from a single specimen collected in 1825, and is believed to have gone extinct around 1850. The Oahu oo (Moho apicalis) was last seen in 1837. The Oahu akepa (Loxops wolstenholmei) was last seen in 1930. The Oahu akialoa (Akialoa ellisiana) is known only from two specimens collected in the mountains of Oahu in 1837. There were undocumented reports in 1937 and 1940, but the species is now considered extinct. The Oahu nukupuu (Hemignathus lucidus) was considered common in 1860, but became extinct by the end of the nineteenth century. The Oahu alauahio (Paroreomyza maculata) was a type of honeycreeper last definitely reported in 1985. Subsequent searches have failed to find it, and the species is most likely extinct. The Oahu amakihi (Chlorodrepanis flava) is restricted to two small mountainous areas. The Oahu elepaio (Chasiempis ibidis) is a small flycatcher confined to a few isolated mountain areas, where the total remaining population is about 1250.

Kauai Known as the ‘Garden Isle’, Kauai (Kaua’i in Hawaiian) lies north-west of Oahu. Two small thrushes (Myadestes) were historically endemic to Kauai. The kamao (M. myadestinus) was once the most common forest bird on the island, but by 1928 was confined to the Alaka’i Wilderness Preserve. The last definite record was in 1985. The puaiohi (M. palmeri) is confined to a single upland area, where it is somewhat protected within the Alaka’i Wilderness Preserve. In 2007 the total population was estimated at 200–500. A captive-breeding programme has been successful. The Kauai akialoa (Akialoa stejnegeri) was widespread on the island at the beginning of the twentieth century. By the 1960s it was very rare and confined to the Alaka’i Wilderness Preserve, where it was last recorded in 1969. The Kauai oo (Moho braccatus) was historically common, but by the 1970s was confined to the Alaka’i Wilderness Preserve. In 1981 a single pair remained, the female ultimately disappearing after a hurricane the following year and the male last seen in 1985. The last report, of vocalizations only, was in 1987. The Kauai nukupuu (Hemignathus hanapepe) was last definitely reported in 1899, although occasional, unconfirmed sightings have continued up to the present century. The Kauai elepaio (Chasiempis sclateri) is a small flycatcher that lost roughly half its population owing to a hurricane in 1992, but has since largely recovered.

The akekee (Loxops caeruleirostris) has declined rapidly over the past few decades. The total population is about 1000. The Kauai creeper or akikiki (Oreomystis bairdi) is confined to a single upland area, where the total population is about 500. The Kauai amakihi (Chlorodrepanis stejnegeri) is confined to a few mountainous areas, although still relatively common. The anianiau (Magumma parva) is a type of honeycreeper confined to montane forests on Kauai.

Lanai Also known as ‘Pineapple Island’ owing to its past as an islandwide pineapple plantation, today Lanai (Lana’i in Hawaiian) is mostly privately owned. The Lanai olomao (Myadestes lanaiensis lanaiensis) was last seen in 1933. The Kona grosbeak (Chloridops kona) was a type of honeycreeper already rare at the time of its discovery, being restricted to a small area of forest on lava flows. It was last collected in 1894. The Lanai hookbill (Dysmorodrepanis munroi) is known only from a single specimen collected in 1913, along with single sightings in 1916 and 1918. The Lanai alauahio (Paroreomyza montana montana) was extinct around 1937.

Molokai Located east of Oahu and north of Lanai, Molokai (Moloka’i in Hawaiian) developed from two separate shield volcanoes. The Molokai alauahio (Paroreomyza flammea) was last recorded in the early 1960s. A survey in 1979 failed to find it.

Niihau The westernmost of the inhabited Hawaiian Islands, Niihau (Ni’ihau in Hawaiian) lies about 30 km west of Kauai. Little more than an extinct volcano rising from the sea floor and quite arid, its intermittent playa lakes, however, provide important wetland habitats for threatened waterfowl such as Hawaiian coot (Fulica alai), Hawaiian black-necked stilt (Himantopus mexicanus knudseni), and Hawaiian duck (Anas wyvilliana). Barren of trees for centuries, in recent years the owners have begun an ambitious reforestation campaign.

Nihoa Nihoa is a small, steep, rocky island. It is the tallest of 10 islands in the uninhabited north-western Hawaiian Islands. The millerbird (Acrocephalus familiaris) is a type of warbler divided into two subspecies. The Nihoa millerbird (A. f. kingi) is historically endemic to Nihoa, where numbers have fluctuated from less than 50 to more than 800. In recent years individuals have been translocated to Laysan, where the population stands at around 150.

Laysan Located 1500 km north-west of Honolulu, Laysan (Kauo in Hawaiian) is an atoll of sorts, although the land completely

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The Line Islands The Line Islands, named from their location on the equator, are a chain of atolls about 1500 km south of Hawaii. Polynesians landed on these islands but did not settle there, so it is entirely white whalers and traders that have changed the face of the islands by digging for guano, drying copra, and merciless hunting. The Line Islands reed warbler (Acrocephalus aequinoctialis) is confined to Kiritimati (Christmas Island) and Teraina (Washington Island). It formerly occurred on Tabueran (Fanning Island) as well, but was extirpated there in 1972.

Teraina Teraina (Washington Island) is a small coral atoll notable for the presence of a freshwater lake. The Teraina gadwall (Mareca strepera couesi) is a now-extinct type of dwarf duck formerly confined to Washington Lake. It was of considerable zoological interest, in that a species so widely distributed throughout the northern hemisphere should have adapted to a tiny mid-ocean lake and evolved a subspecies there. Humans exterminated it in or shortly after 1874.

Kiritimati

Figure 8.12 Laysan millerbird by John Gerrard Keulemans. (Credit: John Gerrard Keulemans.)

Kiritimati (Christmas Island) is a raised coral island located in the northern Line Islands. It is notable for having the greatest land area of any coral atoll in the world. The Kiritimati sandpiper (Prosobonia cancellata) is known only from its type specimen, and appears to have been exterminated by introduced cats and rats by around 1850.

The Marquesas Islands surrounds a shallow central lake some 2.4 m above sea level. The island’s native vegetation was almost completely destroyed due to the introduction of rabbits in the 1890s. The latter were finally exterminated after 1923, but by then three endemic bird taxa were either already extinct or soon would be. The Laysan millerbird (Acrocephalus familiaris familiaris) was estimated to number about 1500 birds in 1915, but was extinct before 1923 due to habitat destruction caused by rabbits. A population of the surviving subspecies (A. f. kingi) from Nihoa has since been successfully introduced to the island. The Laysan apapane (Himatione fraithii) was a type of honeycreeper that became extinct in 1923 due to loss of habitat, with the final blow being a violent storm in which what are thought to have been the final three individuals were killed. A short, silent film clip of one of the latter is still in existence. The Laysan finch (Telespiza cantans) is today largely confined to Laysan, with additional small populations on two very tiny islands in the Pearl and Hermes Atoll. In 1903 the introduction of rabbits to Laysan led to a serious decline, but the species recovered rapidly after the former were exterminated in 1923. An introduced population on Midway Island succumbed to rats during the 1940s.

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The Marquesas Islands (Îles Marquises in French) are a remote volcanic island group located about 1400 km north-east of Tahiti. They fall into two main divisions: a northern one centered around the large island of Nuku Hiva, and a southern one clustered around Hiva ’Oa. In contrast to the lush vegetation of other Polynesian islands they are remarkably dry and, until relatively recently, little affected by European contact. The Marquesan swamphen (Porphyrio paepae) is known from Hiva Oa and Tahuata, where it is thought to have been exterminated by hunting and introduced predators. In Paul Gauguin’s 1902 painting Le Sorcier d’Hiva Oa ou le Marquisien à la cape rouge there is a bird that resembles native descriptions of the Marquesas swamphen being killed by a dog. The explorer Thor Heyerdahl may have seen one as well in 1937, but the species must have gone extinct soon after. The Marquesan imperial pigeon (Ducula galeata) has been heavily depleted by introduced species and hunting, and is today largely confined to isolated valleys in the west and north of Nuku Hiva, with a small translocated population on Ua Huka. The Marquesan ground dove (Pampusana rubescens) is currently confined to two uninhabited, cat-free islets (Hatuta’a and Fatu Huku). Subfossil specimens indicated that

The Polynesian Region

it formerly occurred on Nuku Hiva and at least three other islets as well, suggesting that the species formerly ranged throughout the Marquesas. The Marquesan kingfisher (Todiramphus godeffroyi) historically occurred on Hiva Oa and Tahuata, but was extirpated from the former by the late 1990s. In 2003 the total population was estimated at between 400 and 500. The Marquesan monarch (Pomarea mendozae) is a type of flycatcher divided into two subspecies. The Hiva Oa monarch (P. m. mendozae) was historically found on Hiva Oa and Tahuata. It is now extinct, having last been reported in 1975.

Nuku Hiva Nuku Hiva is the largest of the Marquesas. Once almost entirely forested, today the central part consists of a high grassy plateau dotted with pine forest plantations. The red-moustached fruit dove (Ptilinopus mercierii) is a now-extinct species divided into two subspecies. The Nuku Hiva red-moustached fruit dove (P. m. mercierii) is known only from a single specimen collected from Nuku Hiva between 1836 and 1839. The Nuku Hiva monarch (Pomarea nukuhivae) was considered uncommon as early as 1922 and rare by the 1930s, when the last confirmed sightings took place. It was not seen in 1972 or 1975 despite several weeks of intensive searches (although there was an unconfirmed report in the latter year made by hunters of a bird matching its description). Further unsuccessful searches in the late 1990s seem to confirm its extinction. The northern Marquesan reed warbler (Acrocephalus percernis) is not considered to be threatened as a species, although four of its subspecies are. The Nuku Hiva reed warbler (A. p. percernis) is confined to Fatu Hiva. The Nuku Hiva coastal stream goby (Stenogobius caudimaculosus) is confined to Nuku Hiva.

Ua Huka Ua Huka is located in the northern Marquesas Islands, approximately 40 km east of Nuku Hiva. While small, it still has around 30 per cent of its original forest cover and remains uninfested with black rats, although feral goats are a problem. The ultramarine lorikeet (Vini ultramarina) was formerly common throughout the Marquesas but is now found only on Ua Huka. In 2012 the total population was estimated at less than 2500. The Ua Huka monarch (Pomarea iphis) is confined to the dry forests of Ua Huka. The Ua Huka reed warbler (Acrocephalus percernis idae) is confined to Ua Huka.

Hiva Oa Hiva Oa is the second largest of the Marquesas Islands and the largest of the southern Marquesas group. The Hiva Oa red-moustached fruit dove (Ptilinopus mercierii tristrami) is known from around a dozen specimens, the last of which was collected in 1922. A report from 1980 appears

to be an error, but in any event the form is now certainly extinct, a victim of introduced predators. The Hiva Oa coastal stream goby (Stenogobius marqueti) and the Atuana goby (Sicyopterus marquesensis) are confined to Hiva Oa.

Fatu Hiva Fatu Hiva is the southernmost of the Marquesas Islands. The Fatu Hiva monarch (Pomarea whitneyi) was long considered secure and relatively common despite being confined to one tiny island. However, since the arrival of black rats in 2000 it has declined rapidly and is presumably facing extinction. In 2009 the total population was estimated at just 67 birds. The southern Marquesan reed warbler (Acrocephalus mendanae) is divided into three subspecies. The Fatu Hiva reed warbler (A. m. fatuhivae) is confined to Fatu Hiva.

Eiao Eiao is a small, uninhabited island located in the northern Marquesas. It and the surrounding rocks were proclaimed a nature reserve in 1992, but have since suffered extensive habitat destruction by feral goats, sheep, and pigs. The Eiao monarch (Pomarea fluxa) was last observed in 1977, and is now considered extinct. It is thought to have fallen victim to rats, feral cats, and perhaps disease brought by the introduced chestnut-breasted manakin (Lonchura castaneothorax). The Eiao reed warbler (Acrocephalus percernis aquilonis) is confined to dry upland forest on Eiao, where the total population in 1987 was estimated at between 100 and 200. It is now possibly extinct.

Ua Pou Ua Pou is located about 50 km south of Nuku Hiva in the northern Marquesas. The Ua Pou monarch (Pomarea mira) was last observed in 1985 and thought to be extinct. However, an unconfirmed sighting of a single specimen in 2010 has raised hopes that the species may still survive in very small numbers. The Ua Pou reed warbler (Acrocephalus percernis dido) is confined to Ua Pou. The Ua Pou coastal stream goby (Stenogobius squamosus) is confined to Ua Pou.

Mohotani Mohotani is an uninhabited island located south-east of Hiva Oa and east of Tahuata, in the southern Marquesas. The Mohotani monarch (Pomarea mendozae motanensis) is confined to Mohotani, where the population is small but apparently stable. The Mohotani reed warbler (Acrocephalus mendanae consobrina) is confined to Mohotani.

Hatutu Hatutu is a small island located approximately three kilometres north-east of Eiao, in the northern Marquesas Islands.

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The Hatutu reed warbler (Acrocephalus percernis postremus) is confined to Hatutu.

The Savai’i white-eye (Zosterops samoensis) is a type of passerine bird confined to Savai’i, where it is not uncommon.

Wallis and Futuna Islands

The Society Islands

Located in the South Pacific about two-thirds of the way between Hawaii and New Zealand, the Wallis and Futuna Islands (Îles Wallis-et-Futuna in French) consist of three main volcanic tropical islands along with a number of tiny islets, divided into two separate archipelagos lying about 260 km apart. Four species of freshwater goby are endemic to small estuarine streams on Futuna Island, where they are highly threatened by habitat alteration due to taro farming. Futuna’s emperor (Akihito futuna) and the Wallis and Futuna stiphodon (Stiphodon rubromaculatus) are both confined to a single short stream, where they are very rare. Sasal’s sicyopus goby (Smilosicyopus sasali) and Keletaona’s coastal stream goby (Stenogobius keletaona) are both known from two streams but, like the previous two species, are believed to have been formerly more widespread.

The Society Islands (Îles de la Société in French) are located in the south-central Pacific. Consisting of an eastern (Windward) and western (Leeward) group, these islands have suffered the extermination of more vertebrate species than any other island group of the South Seas except for the Hawaiian Islands. Paradoxically, this sad record seems to be partly explained by the unusual friendliness towards Europeans of the Polynesians on the islands. As a result, European explorers, whalers, and traders frequented Tahiti much more than other islands. At each visit, rats went ashore and eventually exterminated a number of birds. The Tahiti red-billed rail (Hypotaenidia pacifica) was extirpated from Tahiti by 1844, and last recorded on Mehetia in the 1930s.

The Samoan Islands

Tahiti is the highest and largest island in French Polynesia and is divided into two parts: the bigger, north-western Tahiti Nui and the smaller, south-eastern Tahiti Iti. Formed from volcanic activity and surrounded by coral reefs, it was originally settled by Polynesians between A . D . 300 and A . D . 800. The Tahiti crake (Zapornia nigra) was a type of rail known from two illustrations, one by Forster from Tahiti during Cook’s second voyage (1772–75), and the other by Miller in 1784. It presumably became extinct soon after.

The Samoan Islands lie in the central South Pacific about 800 north-east of Fiji. The larger islands are volcanic in origin, mountainous, and covered in tropical moist forests. Some of the smaller islands are coral atolls with black sand beaches. The insular mouse-eared bat (Myotis insularum) is known only from a perhaps incorrectly labelled type specimen, said to have originated from Samoa. The Samoan tooth-billed pigeon (Didunculus strigirostris) was a superficially parrot-like species that was very common when first discovered during the mid-nineteenth century, but near extinction 50 years later due to introduced predators. Today it is found only in forested areas of Savai’i, Upolu, and Nu’ulua. The mao (Gymnomyza samoensis) is a large honeyeater that still occurs in small numbers on Savai’i and Upolu. It may have formerly been found on Tutuila as well. The Samoan emo skink (Emoia samoensis) is confined to the Samoan Islands. The Salele flagtail (Kuhlia salelea) is a type of freshwater fish confined to a few Samoan rivers.

Tahiti

Savai’i Savai’i is comprised of a shield volcano and is the largest and highest of the Samoan Islands. It is home to the Central Savai’i rainforest, the largest continuous area of moist forest in Polynesia. The Samoan moorhen (Pareudiastes pacificus) is (or was) an almost flightless species that was long thought to have been exterminated by introduced cats, compounded by hunting, not having been observed since 1873. In 1984, however, there were two possible sightings in upland forest west of Mount Elietoga, and in 2003 a possible sighting of two individuals on Mount Sili. Nevertheless, the species is now most likely extinct.

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Figure 8.13 Early 1770s illustration of a Tahiti red-billed rail by Johann Georg Adam Forster. (Credit: Johann Georg Adam Forster.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The Polynesian Region

Figure 8.15 Illustration of a Raiatea starling by Johann Georg Adam Forster, 1774. (Credit: Johann Georg Adam Forster.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

Figure 8.14 Painting of a Tahiti crake by Johann Georg Adam Forster. (Credit: Johann Georg Adam Forster.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

The Tahiti sandpiper (Prosobonia leucoptera) was originally collected by Johann Reinhold Forster and painted by his son in 1773. Another specimen, now lost, was taken in 1777. Introduced rats are believed to have caused the extinction. The Tahiti black-fronted parakeet (Cyanoramphus zealandicus) was endemic to Tahiti, where it is known from three specimens collected on Cook’s voyage in 1773, a fourth in 1842, and a fifth in 1844. It was extinct soon after. The so-called Liverpool pigeon (Caloenas maculata) is a poorly known species known from two museum specimens of unknown providence, the first collected between 1783 and 1823 and now lost, and another still in a museum in Liverpool. The latter is believed to have originated in Tahiti, but the species has not been reported there since 1928, when the only possible sightings were made. It is presumably extinct, a likely victim of hunting. The Tahiti monarch (Pomarea nigra) was extremely rare throughout the twentieth century and has long been reduced to just four valleys, where the total population in 2015 was thought to be less than 50. The Tahiti reed warbler (Acrocephalus caffer) is confined to Tahiti, where it is absent from the eastern peninsula and localized to between 6 and 12 valleys. In 1993 the total population was estimated at around a few hundred individuals.

Raiatea Raiatea is the second largest of the Society Islands. The Raiatea parakeet (Cyanoramphus ulietanus) is known only from two specimens collected in 1773. The Raiatea fruit dove (Ptilinopus chrysogaster) breeds on Raiatea, Tahaa, Bora Bora, and Maupiti. The total population is thought to be less than 2500. The Raiatea starling (Aplonis ulietensis) is known only from a 1774 painting of the type specimen, now lost, as well as from contemporary descriptions and a few field notes. An expedition to the island in 1850 failed to find the species, which likely became extinct following the arrival of rats.

Figure 8.16 Moorea sandpiper by William Ellis, painted during Captain James Cook’s Second Voyage (1772–75). (Credit: William Ellis.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

Two subspecies of Society Islands reed warbler (Acrocephalus musae) formerly occurred in the Society Islands. The Raiatea reed warbler (A. m. musae) was confined to Raiatea, where it went extinct sometime during the late nineteenth or early twentieth centuries.

Moorea Moorea is a high island located 17 km north-west of Tahiti. The Moorea sandpiper (Prosobonia ellisi) was a type of shorebird that has not been recorded since two original specimens were collected in 1777. Both are now lost, but paintings based on them survive. The Moorea kingfisher (Todiramphus youngi) is confined to Moorea where it is generally uncommon, although abundant at one locality. The Moorea reed warbler (Acrocephalus longirostris) may perhaps be extinct, although there have been two unconfirmed records this century which suggest it could still exist in very small numbers.

Maupiti Maupiti is a small coral atoll with a volcanic island in its midst.

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The Polynesian Realm

The Maupiti monarch (Pomarea pomarea) is known only from the type specimen collected in 1823. It presumably became extinct as a result of habitat destruction and invasive species.

Huahine Huahine is located in the Leeward Islands group. The Huahine reed warbler (Acrocephalus musae garretti) was confined to Huahine, where it went extinct sometime during the late nineteenth or early twentieth centuries.

The Tuamotu Archipelago The Tuamotus (Îles Tuamotu in French) consist of 76 small islands and atolls. Together these form the largest chain of atolls in the world, stretching as they do over an area of ocean roughly the size of Western Europe. All are low coral islands, being little more than high sand bars built upon coral reefs. The Tuamotu sandpiper (Prosobonia parvirostris) is a small wading bird historically widespread in the Tuamotu Archipelago but long restricted to a few predator-free, usually uninhabited islands. Currently only five islands are known to support populations (Tenararo, Morane, Reitoru, Tahanea, and Raraka), where the total number in 2003 was thought to be around 1300. The Tuamotu ground dove (Pampusana erythropterus albicolis) was formerly found throughout the Tuamotu Archipelago but is now confined to Hao and probably Tahanea atolls.

Niau Niau is a small atoll notable for its remnant tropical forests. The Tuamotu kingfisher (Todiramphus gambieri) is divided into two subspecies. The Niau kingfisher (T. g. niauensis) is confined to Niau.

Makatea Makatea is a raised coral atoll located in the northwestern Tuamotus. The Makatea fruit dove (Ptilinopus chalcurus) is confined to Makatea, where it remains relatively common.

The Gambier Islands Although geographically part of the Tuamotus, the Gambier Islands (Îles Gambier in French), at the south-eastern extreme of the archipelago, are geologically distinct. They consist of a small group of volcanic islands, remnants of a caldera, along with islets on the surrounding fringing reef. The Mangareva kingfisher (Todiramphus gambieri gambieri) became extinct prior to 1922.

The Tonga Islands The Tonga Islands are an archipelago of about 170 islands, of which 36 are inhabited. They are located directly south of Samoa and about two-thirds of the way from Hawaii to New Zealand. The Tongan scrubfowl (Megapodius pritchardii) was once widespread in the Tonga Islands but is now confined to

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Niuafo’ou and Fonualei, the latter population having been introduced. The Tongan ground skink (Tachygyia microlepis) has not been recorded since the early nineteenth century, and is presumably extinct.

Eua Eua is a hilly and still heavily forested island, much of which is protected as Eua National Park. The Eua forest gecko (Lepidodactylus euaensis) is confined to the island, where it remains common.

The Cook Islands Located north-east of New Zealand between French Polynesia and American Samoa, the Cook Islands comprise 15 major atolls (sunken volcanoes topped by coral growth) and two submerged reefs, divided into a northern and southern growth. There is a large introduced population of ship rats that have drastically reduced bird populations. The Cook Islands reed warbler (Acrocephalus kerearako) remains fairly common on Mangaia and Miti’aro.

Rarotonga The volcanic island of Rarotonga stands over 4500 m above the ocean floor. The interior is dominated by eroded volcanic peaks cloaked in dense vegetation. The Cook Islands fruit dove (Ptilinopus rarotongensis) is divided into two subspecies. The Rarotonga fruit dove (P. r. raotongensis) is confined to Rarotonga, where it is moderately common. The Rarotonga monarch (Pomarea dimidiata) was thought to be extinct in the early 1900s and was long considered one of the world’s rarest birds. A survey in 1983 located an estimated 35–50 individuals in a few isolated valleys, although since that time the species has made a remarkable recovery. The Rarotonga starling (Aplonis cinerascens) occurs in the rugged interior of the island. In 1973 it was still not uncommon, although by 1984 the population had been considerably reduced. In 2011 it was estimated at 2350. The Rarotonga emo skink (Emoia tuitarere) is confined to a single locality, where it is still fairly common.

Mauke Mauke is characterized by a central volcanic plateau, deep underground caves and lakes, and jagged fossilized coral extending far inland. The Mauke starling (Aplonis mavornata) is known only from the type specimen collected in 1825 from Mauke. The species was not found on the next ornithological visit to Mauke in 1975, and the species is believed to be extinct, a victim of introduced brown rats.

Atiu Atiu is located 190 km north-east of Rarotonga. The north of the island is home to the largest wetlands area in the Cook Islands.

The Polynesian Region

The Atiu fruit dove (Ptilinopus rarotongensis goodwini) is confined to Atiu, where it is moderately common. The Atiu swiftlet (Aerodramus sawtelli) is confined, when breeding, to just two caves on Atiu, and is therefore vulnerable to stochastic events and human activities.

Mangaia Mangaia is the most southerly of the Cook Islands and the second largest, after Rarotonga. Introduced species include the aggressive common myna (Acridotheres tristis), cats, and rats. The Mangaia kingfisher (Todiramphus ruficollaris) is confined to the island, although its population is considered to be stable.

The Austral Islands The Austral Islands (Îles Australes in French) are located 640 km south of Tahiti, in south-western French Polynesia. They are comprised of two separate archipelagos, the Tubuai Islands (consisting of Tubuai, Raivavae, Rimatara, Rurutu, and the uninhabited Íles Maria) and the Bass Islands (consisting of Rapa Iti and Marotiri).

Tubuai Tubuai (also known as Tupua’i) is the main island of the Tubaui Island group. It features two volcanic domes and is surrounded by a barrier reef and numerous islets. Randall’s coastal stream goby (Stenogobius randalli) is confined to Tubuai.

Rimatara Rimatara is characterized by its fern-covered central hills, forests, and numerous swamps. Kuhl’s lorikeet (Vini kuhlii) was originally found only on Rimatara, but was introduced to Teraina (Washington Island), Tabuaeran (Fanning Island), and Kiritimati (Christmas Island) by Polynesians, and more recently to Atiu in the Cook Islands. It appears to have been historically present on at least five of the southern Cook Islands as well, but was exterminated there due to excessive exploitation for its beautiful red feathers. The total population in all areas is perhaps 2000. The Rimatara reed warbler (Acrocephalus rimitarae) is confined to the island, where it remains relatively common.

The Rapa Iti shearwater (Puffinus myrtae) is a type of seabird that historically bred on Rapa Iti and surrounding islets, but was extirpated from the main island by introduced rats, goats, and feral cats. The Rapa Iti fruit dove (Ptilinopus huttoni) is confined to undisturbed forest fragments on the island, where the total population was estimated at around 275 individuals in 1990. Two freshwater fish species, the Rapa Iti goby (Sicyopterus rapa) and Julien’s stiphodon (Stiphodon julieni), are confined to three small rivers all less than 3 km in length.

The Pitcairn Islands The Pitcairn Islands are a group of four volcanic islands spread out over a wide area of ocean in easternmost Polynesia. The Henderson petrel (Pterodroma atrata) is now known to breed only on Henderson Island, but may also have bred on Pitcairn Island in the recent past. The species was wiped out on Dulcie Island by invasive rats in 1922. While its non-breeding range is not well known, it has been sighted on Easter Island.

Pitcairn Island Pitcairn is the only inhabited island in the Pitcairn Islands group. The Pitcairn reed warbler (Acrocephalus vaughani) is endemic to Pitcairn, where it is threatened by introduced cats and rats.

Henderson Island Henderson Island is a small, uninhabited, raised-reef island whose remoteness and unsuitability for human habitation have long made it the subject of scientific study. It features a remarkable number of endemic species, including at least three species of pigeon driven to extinction by early Polynesians around A . D . 1000. All are vulnerable to habitat destruction and the accidental introduction of invasive species. The Henderson crake (Zapornia atra) is a type of flightless rail confined to Henderson Island. Stephen’s lorikeet (Vini stepheni) is confined to Henderson Island. The Henderson fruit dove (Ptilinopus insularis) is confined to Henderson Island. The Henderson reed warbler (Acrocephalus taiti) is confined to Henderson Island.

Rurutu

Easter Island

The northernmost of the Austral Islands, Rurutu is notable for its many caves. The Rurutu stiphodon (Stiphodon discotorquatus) is a type of freshwater goby known only from a single short river on Rurutu. It was last recorded in 1985, and may be extinct.

Easter Island (Rapa Nui in Polynesian/Isla de Pascua in Spanish) is located in the south-eastern Pacific Ocean. Famous for its monumental statues, Polynesians lived there for more than 1000 years and built a thriving civilization. Scientists have found evidence of extensive deforestation and soil erosion, however, indicating that a massive ecological disaster took place here due most likely to overpopulation. By the time of European arrival in 1722 the island’s population had dropped to just two or three thousand, and introduced disease and Peruvian slave-trading reduced that further, to a

Rapa Iti Rapi Iti is the largest and only inhabited island in the Bass Islands. It is essentially the peak of a sinking volcano, the caldera forming a protected central bay.

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low of only 111 inhabitants in 1877. Today, Rapa Nui is one of the most remote inhabited places on Earth. There are no longer any native terrestrial vertebrate species on Easter Island, although at least five undescribed forms (two rails, a heron, and two parrots) are known to have gone extinct there at some point in the past. The complete destruction of the island’s unique subtropical broadleaf forests is the likely cause.

Balance for the Polynesian Region The first people to arrive in this region are believed to be related to the sea-migrating Austronesian people who most likely originated in Taiwan several thousand years ago. Remarkable navigators, they had migrated slowly south and east from Asia before ultimately sweeping out over most of the Pacific islands by means of outrigger canoe, finally reaching the western Polynesian islands by 900 B . C ., and the eastern ones a thousand years later. Everywhere that they went they brought with them domestic animals such as chickens, dogs, and pigs, which would certainly play a role in changing the environment. They also carried with them the Polynesian rat (Rattus exulans), which quickly had fatal consequences for several ground-nesting bird species. Yet on the whole the incursions were not yet too destructive. These people employed a garden type of agriculture, with groves of breadfruit, coconuts, and bananas being easily produced and usually without any effect on the soil. The interior of the islands were often left untouched by the settlers, who were also dependent upon the sea for most of their food. This form of agriculture combined with fishing was the basis for an idyllic way of human life, but did not allow a strong population growth all over the Pacific islands. Tribal wars, cannibalism, and voluntary limitation of excess children kept the human population in balance with the carrying capacity of the soil and sea.

When Europeans first began to colonize these islands about 250 years ago, however, all sense of biological harmony was destroyed. They latter brought with them exotic plants and insects in large numbers and introduced sheep, goats, and pigs almost everywhere. Foreign diseases spread among the native population. New methods of farming, unsuitable to the islands, caused increasing erosion. Most disastrous of all, their ships carried even more disastrous invasive rodents than those previously brought by the Polynesians, namely brown and black rats (Rattus norvegicus and R. rattus), as well as feral cats, all destroyers of defenceless endemic birds. The destruction of natural vegetation and animal life upset the ecological equilibrium in many places, making it more and more difficult for human beings to survive. The native fauna began to be wiped out. More recently the Pacific islands have undergone a massive population explosion and global climate change threatens to return the more low-lying islands to the sea once more, making the very future of humans on these islands uncertain. In recent historical time (i.e. since A . D . 1500), the Polynesian Region has lost at least 46 species/8 subspecies of vertebrates. Among the extinct forms 1 species is a mammal, 44 species/8 subspecies are birds, and 1 species is a reptile. Another 8 species are possibly extinct. In addition, there are 86 species/17 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 1 species/1 subspecies are mammals, 64 species/16 subspecies are birds, 6 species are reptiles, and 15 species are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

1 species

~ species

~ species

1 species

~ subspecies

~ subspecies

~ subspecies

1 subspecies

1 taxon

~ taxa

~ taxa

2 taxa

44 species

7 species

1 species

64 species

8 subspecies

~ subspecies

~ subspecies

16 subspecies

52 taxa

7 taxa

1 taxon

80 taxa

1 species

~ species

~ species

6 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

1 taxon

~ taxa

~ taxa

6 taxa

~ species

~ species

~ species

~ species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

~ taxa

~ taxa

~ taxa

~ species

1 species

~ species

15 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

1 taxon

~ taxa

15 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

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Balance for the Polynesian Realm

(cont.)

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Total vertebrates

46 species

8 species

1 species

86 species

8 subspecies

~ subspecies

~ subspecies

17 subspecies

54 taxa

8 taxa

1 taxon

103 taxa

Note: ~, not applicable.

Balance for the Polynesian Realm As we have seen, during the two and a half centuries that Europeans have been exploring, trading, whaling, and settling in the island world of the South Seas they have managed to wreak considerable havoc. Considering the small land area these islands represent, the figures are frightening. How much remains now that Western civilization with its diseases, introduced species and commercialization has swept over the region? Because there are thousands of islands involved, it is difficult to give a simple answer. Almost every one has its own special character, or at least certain features, even when only a narrow sound separates two neighbouring islands. On isolated islands or in the remote mountain regions of even the larger ones remnants may still be found of the original flora and fauna. Unfortunately, the number of human inhabitants continues to grow at an alarming rate.

In recent historical time (i.e. since A . D . 1500), the Polynesian Realm as a whole has lost at least 86 species/16 subspecies of vertebrates. Among the extinct forms 3 species are mammals, 79 species/16 subspecies are birds, 3 species are reptiles, and 1 species is a freshwater fish. Another 10 species are possibly extinct, and 2 species are currently extinct in the wild. In addition, there are 231 species/42 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 13 species/9 subspecies are mammals, 157 species/32 subspecies are birds, 25 species/1 subspecies are reptiles, 3 species are amphibians, and 33 species are freshwater fishes.

Vertebrate Class

Extinct

Possibly Extinct

Extinct in the Wild

Threatened

Mammals

3 species

1 species

~ species

13 species

~ subspecies

~ subspecies

~ subspecies

9 subspecies

3 taxa

1 taxon

~ taxa

22 taxa

79 species

8 species

2 species

157 species

16 subspecies

~ subspecies

~ subspecies

32 subspecies

95 taxa

8 taxa

2 taxa

189 taxa

3 species

~ species

~ species

25 species

~ subspecies

~ subspecies

~ subspecies

1 subspecies

3 taxa

~ taxa

~ taxa

26 taxa

~ species

~ species

~ species

3 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

~ taxa

~ taxa

~ taxa

3 taxa

1 species

1 species

~ species

33 species

~ subspecies

~ subspecies

~ subspecies

~ subspecies

1 taxon

1 taxon

~ taxa

33 taxa

86 species

10 species

2 species

231 species

16 subspecies

~ subspecies

~ subspecies

42 subspecies

102 taxa

10 taxa

2 taxa

273 taxa

Birds

Reptiles

Amphibians

Freshwater fishes

Total vertebrates

Note: ~, not applicable.

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Chapter

9

The Nearctic Realm

The Nearctic Realm, as here defined, consists of most of the continental regions of the United States and Canada, along with parts of northern Mexico and certain satellite islands such as Bermuda. It is bordered by the Arctic Realm to the north and by the Caribbean and Neotropical realms to the south. The physical features and the major biogeographic regions of North America are almost a mirror of those of Palearctic Eurasia. In the northern part, below the polar areas, are vast coniferous forests succeeded by deciduous woodlands, grass prairies, deserts, chaparral, and subtropics. Cutting through these from north to south in western North America is an enormous mountain chain that divides midway into two separate ranges. Today, the climate ranges from subarctic in the north, through temperate to subtropical in Florida and Mexico. When the ancient supercontinent of Pangaea split into two about 180 million years ago, North America remained joined to Eurasia as part of the supercontinent of Laurasia, while South America was part of the supercontinent of Gondwana. North America later split from Eurasia. The latter has, at various times, been joined by land bridges to Asia, Central and South America since then, which allowed for an exchange of plant and animal species between them (known to science as the Great American Interchange). As a consequence, the Nearctic Realm has relatively few endemics but a rich biodiversity. As in Eurasia, vast Pleistocene ice sheets advanced southward from the polar regions, reaching beyond the Great Lakes and thus covering at their maximum more than half of the continent (curiously, parts of Alaska escaped glaciation). Many large animals, including horses, camels, tapirs, mammoths, mastodonts, ground sloths, sabre-tooth cats (Smilodon), the giant short-faced bear (Arctodus simus), and at least two species of American cheetah (Miracinonyx), became extinct in North America at the end of the ice ages. At the same time, evidence of early modern humans appears and marks the beginning of the ongoing Holocene extinction event. Previously, megafaunal extinctions were believed to have been caused by the changing climate, but many scientists now believe that, while climate change contributed to these extinctions, the primary cause was hunting by these newly arrived humans or, in the case of some large predators, indirectly as a result of their prey becoming scarce. The American bison (Bison bison), brown or grizzly bear (Ursus arctos), moose

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(Alces alces), and elk or wapiti (Cervus canadensis) all entered North America around the same time as the first humans and explanded rapidly, filling ecological niches left empty by the newly extinct North American megafauna. The ‘grizzly bear’, mentioned above, is the name used for various populations of North American brown bear (Ursus arctos), which first arrived on the continent some 50,000 years ago after crossing the Bering Stait land bridge from eastern Asia. Historically found continuously, and in a very wide ecological range, from the arctic coast of Alaska in the north to Mexico in the south and from the eastern edge of the Great Plains westward to the Pacific, it was in particular an inhabitant of the entire extent of the Rocky Mountains. The almost continual human warfare that has been waged against it, however, along with massive habitat destruction, has resulted in the grizzly having long since vanished from most of these areas. Today it survives only in the large wilderness regions of Alaska and north-western Canada, as well as in some national parks and reserves in the northern continental United States, where it lives mostly in the mountains above the timber line. While the precise taxonomy remains confused and disputed, many of the more isolated populations are known to be under serious threat and a few have already become extinct. Two examples of this include the California grizzly bear (U. a. californicus), which lived in the Sierra Nevada and some coastal areas. The last known individual was shot in 1922, although another individual may have been spotted in Sequoia National Park in 1924. The so-called Mexican grizzly bear, which is now generally thought to have been a population of mainland grizzly (U. a. horribilis), historically occurred in Arizona, New Mexico, Texas, and northern Mexico. Owing to human persecution, by the 1930s this range had decreased to just three isolated mountains in central Chihuahua, Mexico where, by 1960, only 30–40 were left. Hunting, trapping, and poisoning continued unabated, unfortunately, and by the beginning of the 1980s none remained. Whether or not it was actually a distinct subspecies, the loss of these magnificent creatures from so large an area is no less tragic. The grey or timber wolf (Canis lupus) has undergone an enormous reduction in range over the past 150 years due to human persecution and habitat destruction, but still occurs in reasonable numbers in remoter areas. Of the four generally

The Nearctic Realm Figure 9.1 ‘Mexican grizzlies’ in the Field Museum, Chicago, in the early twentieth century. (Credit: Field Museum.)

recognized extant subspecies in North America, only one, the Mexican wolf (C. l. baileyi), is currently considered to be threatened. Two others have gone extinct and are dealt with separately in this text. The cougar, puma, or mountain lion (Puma concolor) remains one of the most widespread mammals in the Western Hemisphere, being found in a variety of habits across the length of the Americas. In the past a great many subspecies were described, although today only six or seven are generally recognized. The North American cougar (P. c. couguar) includes those populations found in the United States, Canada, Mexico, and most of Central America (possibly as far as South America north-west of the Andes). These large cats were intensively hunted following the European colonization. By the end of the nineteenth century they had been largely extirpated in the eastern part of their range, an area extending from the Maritime provinces of Canada to the Great Plains and southward to Georgia and Alabama. Only remnants survived in remote, mainly northern areas as well as in south Florida, where the ‘Florida panther’ (sometimes considered a distinct subspecies) is critically endangered. It is dealt with separately in this book. The American bison or buffalo (Bison bison) is divided into two subspecies. The tragic yet ultimately hopeful story of the plains bison (B. b. bison) has been told so many times that it is almost unnecessary to give a detailed account of it here. When Europeans first penetrated into the interior of North

America this subspecies occurred south of the range of the wood bison (B. b. athabascae) and east of the Rockies to the north-eastern Mexican states of Durango and Nuevo León, east to the Atlantic seaboard from New York to Georgia (and perhaps as far as Florida). Its centre of distribution was the Great Plains, both in lowlands and highlands. Hence, it had an immense range and constituted a tremendous natural resource, converting prairie grass into tons of meat and hides. The enormous herds utilized the prairies without destroying them and made seasonal migrations on a scale rarely seen in terrestrial mammals. Indeed, at the time of the European arrival in North America the bison were the largest aggregation of land animals on that continent, and perhaps in the world. Only the herds of ungulates that once roamed the savannas of Africa could compare with them in numbers. It is generally believed that the native people had hunted bison for centuries before the European settlers streamed westward. In reality it was at a rather late stage that they began to base their economy on these animals. Hunting bison on foot was certainly not productive. Most cultures of the few tribes inhabiting the prairies were based on the cultivation of maize, beans, and squash. However, when horses became available to native people of the prairies during the seventeenth and eighteenth centuries, quite a new culture developed. Tribes from the mountains converged on the prairies, making use of horses, and quickly changed their economy so that it became based on the bison. Because of the horse the Indians could pursue the

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bison effectively, but their hunting was still without any negative effect on the overall population. At the beginning of the nineteenth century, when Europeans started to push westwards over the plains, the slaughter of bison commenced, but it was not until the construction of the Union Pacific Railroad in the 1860s that the herds began to be annihilated. Professional hunters moved in and millions of animals were killed. Many deplored this wanton destruction, but any measure to stop or regulate the carnage was opposed by politicians, who saw in the destruction a way to get rid of the ‘Indians’. In the 1870s, Idaho attempted to protect the bison as well as other hoofed animals. Another law giving protection to the bison was passed in Congress, but President Ulysses S Grant never signed it and the massacres continued. Despite this, a herd numbering several million was reported as late as 1871. However, from then on the number dwindled rapidly. Between 1870 and 1875 at least two and a half million bison were killed every year. In 1883 the last important herd, about 10,000 animals, was destroyed. Only small, stray herds remained in remote areas. One group in Colorado was destroyed by taxidermists in 1897. In 1899 a census put the number of plains bison at just 541, and in Canada there were only a few. Most of these were collected onto various private ranches, with the last-known wild population, consisting of less than 30 animals, living in the area that later became Yellowstone National Park. Although it was the official policy of the United States government to minimize or exterminate the species, and most farmers considered it to be a pest or a nuisance, some people were concerned about the demise of this American icon and took steps to protect it. Some did so with the express purpose of ranching or hunting the animals, but others, such as the American Bison Society, endeavoured to save the species and to reintroduce it into a least part of its former range. In 1908 the National Bison Range in Montana was established, and later other reserves were set aside in the United States and Canada. With growing numbers bison were reintroduced to a number of other protected areas, thus saving it from extinction at the very last moment. Since then it has continued to increase steadily to its current total of around 20,000, although it remains entirely conservation-dependant and largely restricted to national parks and reserves. Merriam’s elk (Cervus canadensis merriami) was a large deer that inhabited the mountains of California, New Mexico, and Arizona. Competition with cattle and reduction by hunting led to its extermination in about 1906. The Roosevelt elk (C. c. roosevelti) is the largest surviving subspecies of elk in North America, historically found over much of the Pacific Northwest, extending to parts of northern California, where at one time it was heavily hunted. The desire to save it from possible extinction was one of the primary reasons behind the establishment of the Mount Olympus National Monument in 1909 by President Theodore Roosevelt (now Olympic National Park). It was introduced to parts of Alaska in 1928 and reintroduced to coastal British Colombia during the 1980s. The Tule elk (C. c. nannodes) is a notably small subspecies endemic to northern

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and central California, where it historically ranged from the grasslands and marshlands of the Central Valley to the grassy hills of the coast. When the Europeans first arrived an estimated 500,000 roamed these regions, but by 1870 they were thought to have been wiped out by market hunters during the gold rush. However, in 1874–75, a single breeding pair was discovered in the tule marshes of Buena Vista Lake in the southern San Joaquin Valley. These were protected by a local cattle baron named Henry Miller. A sizeable number were built up on Miller’s ranch, but on his death the latter was subdivided and hunting resumed. By 1895 the population had been reduced to 28. In 1933, rancher Walter Dow took a small group of penned elk to his own ranch in the Owens Valley, east of the Sierra Nevada. Although not native habitat for the animals, they nevertheless thrived there. Today there are over 4000 elk scattered among some two dozen reintroduced herds across the state. The Columbia white-tailed deer (Odocoileus virginianus leucurus) was once distributed in the north-west from the Columbia River in the north to southern Oregon. Hunting and habitat destruction had reduced it to just a few hundred individuals by the mid-twentieth century, but it has since recovered and is no longer considered to be threatened. The bighorn sheep (Ovis canadensis) is, as a species, widely distributed throughout the mountains of western North America from southern Canada to Mexico. The desert bighorn sheep (O. c. nelsoni) originally occurred throughout the southwestern United States and north-western Mexico. Heavily hunted, it is now restricted to just three states (Sonora, Baja California, and Baja California Sur) and to Tiburon Island in the Gulf of California. It is unclear whether the desert and peninsular forms represent distinct subspecies, but they are at the very least distinct populations of concern. The pronghorn (Antilocapra americana) is the only survivor of the Antilocapridae, a family of North American antelope whose closest living relatives are the giraffe and okapi. It is also the fastest hoofed animal in the world, able to achieve speeds of up to 96 km per hour. The species formerly occurred in large herds on the American plains, deserts, and tablelands from Alberta in the north to the Pacific slopes in the west and the Mexican plateaus in the south, where they were said to be as numerous as the bison. During the nineteenth century pronghorns were hunted senselessly, and by 1910 only small, scattered groups remained. Like the bison, those of the prairies escaped extinction at the last minute thanks to legal protection. Three subspecies inhabiting desert areas of the south-western United States and Mexico are still considered to be threatened, however. The Baja California pronghorn (A. a. peninsularis) is confined to a small area of the central Baja California Peninsula, Mexico, where the total population is around 200. The Sonoran pronghorn (A. a. sonoriensis) occurs in southwestern Arizona and north-western Mexico (Sonora). The Mexican pronghorn (A. a. mexicana) is found patchily in north-central Mexico (Chihuahua, Coahuila, Durango, Zacatecas, and San Luis Potosí).

The Nearctic Realm

The eastern spotted skunk (Spilogale putorius) was historically common over a wide area of North America, where at one time it was heavily trapped for its fur. Three subspecies are recognized. The Appalachian eastern spotted skunk (S. p. putorius) is now confined to higher-elevation habitats within the Appalachians, although formerly it ranged down into the piedmont and coastal plain of Virginia and the Carolinas. The south-eastern spotted skunk (S. putorius ambarvalis) is found in Florida, Georgia, Alabama, and perhaps elsewhere, while the midwestern spotted skunk (S. p. interrupta) ranges from south-central Canada through the central United States and into north-eastern Mexico. The New England cottontail rabbit (Sylvilagus transitionalis) was formerly widespread in the north-eastern United States and southern Canada, but has undergone a massive decline due to hunting and habitat destruction. It is now confined to a few scattered areas. The endemic ground squirrel (Urocitellus endemicus) and the brown ground squirrel (U. brunneus) are both confined to small areas of Idaho, where they are threatened by loss of habitat. Nelson’s antelope squirrel (Ammospermophilus nelsoni) is confined to the central and western San Joaquin Valley of California as well as to adjacent areas of the inner Coast Ranges. It is threatened by loss of habitat. The California condor (Gymnogyps californianus), a type of vulture, is the largest land and soaring bird in North America. In prehistoric times it was found across the United States as far as Florida, but with the extinction of the Pleistocene megafauna at the end of the last glacial period its range was significantly reduced. Five hundred years ago it still roamed across the south-west and West Coast from the Columbia River to Mexico and Texas. Human persecution and habitat destruction brought a further tremendous decrease in range in numbers during the twentieth century. By the midtwentieth century it had been confined to the coastal mountains of Santa Barbara and Ventura counties in southern California, where it had only limited protection. When it was clear that these populations too were becoming untenable an ambitious conservation plan was put in place by the United States government. As part of this, all remaining wild condors – just 27 birds – were captured by 1987 and for a time the species became extinct in the wild. The survivors were bred at the San Diego Wild Animal Park and the Los Angeles Zoo. Numbers rose through captive breeding and, beginning in 1991, the species was reintroduced back to the wild at three release sites: northern Arizona and southern Utah (including the Grand Canyon area and Zion National Park), the coastal mountains of central and southern California, and northern Baja California. These populations have continued to grow, but the species remains one of the world’s rarest birds. As of 2017 the total population was 463. The bald eagle (Haliaeetus leucocephalus) is a type of sea eagle that, after suffering a precipitous decline during the midtwentieth century, is once again found in wetland habitats

across most of North America. Overuse of the pesticide DDT, which destroyed its eggs, was the primary cause, although direct human persecution in the form of hunting was also a factor. At its lowest point in the 1950s the species was largely restricted to Alaska, the Aleutian Islands, northern and eastern Canada, and Florida. The North American osprey (Pandion haliaetus carolinensis) is a large fish-eating hawk that, although notably widespread, was at one time seriously threatened by overcollection of its eggs and hunting, and later by the spraying of pesticides. It has since made a remarkable recovery and is no longer considered threatened. The spotted owl (Strix occidentalis) is divided into a number of subspecies found patchily from south-western Canada and the western and south-western United States to central Mexico. The California spotted owl (S. o. occidentalis) was historically found in central and southern California and on the Baja California Peninsula, but has been extirpated from the latter. The northern spotted owl (S. o. caurina) is found from south-western British Columbia to northern California. The south-western spotted owl (S. o. huachucae) occurs from Utah and Colorado to Arizona, New Mexico, and extreme western Texas. All are threatened by logging and, to a lesser extent, competition with the barred owl (S. varia), human disturbance, and disease. The plight of the whooping crane (Grus americana) has long been well-known in North America, resulting in one of the most ambitious conservation recovery efforts in history. This magnificent bird formerly bred in the isolated marshes and bogs of north-western Canada south to Alberta, Manitoba, North Dakota, and Iowa, with an additional breeding population on the coast of Louisiana. Its winter quarters were located along the coast of the Gulf of Mexico from Florida to Mexico. It is thought that there were more than 20,000 birds prior to European settlement of these areas, but relentless shooting and disturbances in both the breeding and wintering grounds reduced this total to around 1300–1400 by 1870, and to just a handful by the 1930s. The latter were confined to the southern Mackenzie River region, fortunately within the confines of Wood Buffalo National Park on the border between Alberta and the Northwest Territories. From there they would migrate southward each year to Texas and Louisiana. After 1938 the entire population wintered in the Aransas National Wildlife Refuge in Texas, which had been established especially for them. This provided an opportunity to take a precise census of the birds each wintering season. When the record-keeping began there were just 14. After years of fluctuating, the population reached a high of 38 in 1961. The following year, however, disaster struck – six birds were lost and no young were born. But in the autumn of 1964 the cranes brought 10 young back to Texas, making a total of 42. In 1970 the figure was 57. By this point a small captive-breeding programme was also underway and numbered around two dozen. Since then the efforts have continued unabated. As of 2017 there were four wild populations totalling some 483

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birds. This includes the original natural population along with three reintroduced ones in the eastern United States that are not as yet self-sustaining. The latter includes a large flock that migrates between Wisconsin and Louisiana and two smaller, non-migratory flocks in Florida and Louisiana, respectively. The captive population, meanwhile, totals around 150. All told it has been a remarkable recovery from near and almost certain extinction, but is still a precariously small number to ensure the survival of a long-distance migrant. The red-crowned Amazon (Amazona viridigenalis) is a type of parrot naturally confined to a small area of northeastern Mexico (Tamaulipas, San Luis Potosí and Querétaro), with an additional, possibly feral population in the Rio Grande Valley of southern Texas, where it is threatened by loss of habitat and illegal collection for the international pet trade. The species has been introduced to parts of Florida, California, Puerto Rico, and Oahu in the Hawaiian Islands. The story of the passenger pigeon (Ectopistes migratorius) is one of the most tragic in the annals of conservation history. Prior to the permanent settlement by humans in North America, the breeding area of this species covered a vast region from the Great Plains in the west to the Atlantic coast, and from Manitoba and Quebec in the north to the Appalachians and northern Mississippi in the south. Much has been written about it. There is no doubt that the seemingly fantastic accounts of flocks darkening the sky or of their weight breaking great branches from trees where they perched are reliable. The well-known ornithologist Alexander Wilson estimated 2,230,272,000 birds in a single flock that he saw in 1832. Audubon witnessed another immense flock passing over him for hours on end, in such a concentration that the sunlight was almost blotted out and the sky in all directions, as far as the eye could see, was filled with flying pigeons. He calculated that there were well over one billion. A. W. Schorger found as many as 136 million pigeons in a concentrated nesting area in Wisconsin as late as 1871. When the European settlers pushed westwards they hunted passenger pigeons mercilessly, but it was not until professional hunters began to earn their livelihood by killing the birds that the species started to decrease. The annual slaughter of tens of millions in the 1860s and 1870s was more than it could endure. By the 1880s it had become evident that the species was doomed if hunting remained unregulated. Nobody seems to have heeded the warning signs, however, and in the 1890s the species faded out in the wild. The precise cause of the extinction is difficult to determine, but widespread clearance of forests, combined with the expansion of the railway and telegraph networks that enabled the efficient location and harvesting of nomadic nesting colonies, were clearly important factors. Others include disease and, in the final years, breakdown of social facilitation. The last fully authenticated wild specimen was shot near Oakfield, Illinois, on 12 March 1901, although it is possible that another was killed near Laurel, Indiana on 3 April 1902. Various sight records were made up to 1907. The very last individual of the species, ‘Martha’, died in the Cincinnati

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Figure 9.2 ‘Martha’, the last passenger pigeon, taken in life in 1912. (Credit: Enno Meyer.)

Zoo in 1914. Thus, in less than 50 years, humans had succeeded in wiping out one of the most abundant birds on Earth. The tricoloured blackbird (Agelaius tricolor) is relatively widespread along the Pacific coast of North America from northern California to the upper Baja California Peninsula in Mexico. It also remains extremely common and indeed forms the largest breeding colonies of any Nearctic land bird, with one such colony in the 1930s estimated to have contained around 300,000 individuals. Nevertheless, the species has undergone massive declines in recent years due to loss of habitat and persistent insecticide use. The rusty blackbird (Euphagus carolinus) remains widespread, breeding across the boreal zone of North America from New England to Alaska and wintering across the southwestern United States. Nevertheless, the species has been undergoing massive declines since the mid-twentieth century, the reasons for which are poorly understood. Bendire’s thrasher (Toxostoma bendirei) is a type of passerine bird found over a relatively wide area of the southwestern United States and north-western Mexico. It is declining everywhere due to loss of habitat and, perhaps, drought.

Mountains and Highlands

Belding’s yellowthroat (Geothlypis beldingi) is a type of warbler divided into two subspecies. The nominate form (G. b. beldingi) has been drastically affected by loss of habitat and is now confined to a few small marshes on the southernmost part of the Baja California Peninsula. The northern Belding’s yellowthroat (G. b. goldmaii) is found patchily in the wetlands of the central Baja California Peninsula. The black-capped vireo (Vireo atricapilla) is a small songbird found over a relatively wide area of the southern United States and Mexico. It has undergone rapid population declines in the past due to habitat destruction and degradation. The North American box turtle (Terrapene carolina) is divided into a number of subspecies found collectively throughout North America from south of the Great Lakes and east of the Rocky Mountains to southern Mexico. The eastern box turtle (T. c. carolina) is found in southern Canada and in northern and eastern United States. The three-toed box turtle (T. c. triunguis) occurs from eastern Texas to southeastern Kansas, southern Missouri and south-central Alabama. Baur’s box turtle (T. c. bauri) is found in southeastern Georgia and peninsular Florida, along with the Keys and barrier islands of the Gulf of Mexico. The Gulf Coast box turtle (T. c. major) is found in southern Mississippi, southern Alabama, and western Florida. All are threatened by habitat destruction, roadkill, and capture for the pet trade. The blunt-nosed leopard lizard (Gambelia sila) was historically found in the San Joaquin Valley and adjacent foothills of southern California, where it has been seriously threatened by habitat destruction. It is currently confined to a few scattered parcels of undeveloped land. Slowinski’s corn snake (Pantherophis slowinskii) is a littleknown species confined to eastern Texas, western Louisiana, and possibly southern Arkansas. The southern hog-nosed snake (Heterodon simus) is a harmless species that inhabits the coastal plain of the southeastern United States from North Carolina south to Lake Okeechobee in Florida and west to Mississippi. The species has disappeared from many areas due to a multitude of factors including loss of habitat, predation of eggs and hatchlings by fire ants, pesticides, road mortality, and general human persecution. It is now very rare or possibly extirpated in the western part of its range in Mississippi and Alabama. The Houston toad (Anaxyrus houstonensis) historically occurred in a variety of habitat types across the central coastal region of Texas, but is now confined to a few isolated populations. It disappeared from the Houston area itself during the 1960s, following an extended period of drought and rapid urban expansion. The arroyo toad (A. californicus) is confined to the south-western United States (southern California) and north-western Mexico (Baja California). It is threatened by habitat destruction and predation by introduced fishes and bullfrogs. The Shasta salamander (Hydromantes shastae) is known only from a few localities within a small area of northern

California, typically (although not always) near limestone outcrops. It is threatened by habitat destruction and human disturbance. The limestone salamander (H. brunus) is confined to a small area of the western foothills of the Sierra Nevada in central California, where part of its habitat is protected by a special reserve. The clouded salamander (Aneides ferreus) is found patchily on Vancouver Island (British Columbia), south-western Oregon, and north-western California. It has suffered significant declines due to intensive, short-rotation logging practices, which result in increasing scarcity of coarse woody debris on the forest floor. The Del Norte salamander (Plethodon elongatus) is confined to rocky areas within redwood or Douglas fir forests in south-western Oregon and north-western California.

Mountains and Highlands Like a gigantic barrier, the North American Cordillera runs along the western part of the Nearctic Realm from Alaska into Mexico, its highest peak (Denali, formerly known as Mount McKinley) exceeding 6000 m in height. It consists of three main belts: the Pacific Coast Ranges in the west, the Nevadan belt in the middle (including the Sierra Nevada), and the Laramide belt (including the Rocky Mountains) in the east. Apart from this extensive system of mountains and plateaus there is the much more ancient Appalachian Mountains of the eastern United States and Canada, the peaks and valleys of the Great Basin, and various smaller, isolated massifs located mainly in the southeastern United States. The Utah prairie marmot (Cynomys parvidens) is confined to a small, high-elevation area of south-central Utah. Historically much more widespread, pest control measures in the early twentieth century devastated its range and numbers. By the 1960s it had been reduced to just nine ‘prairie dog towns’ containing less than 3000 animals. The species has recovered somewhat with the banning of poison, but remains threatened. Townsend’s ground squirrel (Urocitellus townsendii) and Nancy’s ground squirrel (U. nancyae) are both confined to small areas of high-desert shrubland in Washington state, where they are threatened by loss of habitat. The robust cottontail rabbit (Sylvilagus robustus) is found patchily in the mountains of south-eastern New Mexico, western Texas, and northern Mexico (Coahuila). The New Mexico shrew (Sorex neomexicanus) appears to be confined to Capitan, Manzano, and Sandia mountains in south-central New Mexico. The Gunnison grouse (Centrocercus minimus) is a chicken-like bird historically found in the highland areas of the west-central United States. Owing to intensive hunting and habitat destruction it is now confined to a few fragmented populations in Colorado and Utah. The pinyon jay (Gymnorhinus cyanocephalus) is a permanent resident of the foothills and lower mountains slopes of the

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western and south-western United States and north-western Mexico (Baja California). It is threatened by the destruction of its preferred habitat (pinyon-juniper woodland). Bicknell’s thrush (Catharus bicknelli) is one of North America’s rarest and most secretive birds, with a breeding range entirely restricted to the north-eastern part of the continent. A habitat specialist, it favours coniferous mountaintops in south-eastern Quebec and the Maritime provinces of Canada, along with parts of New England, from where it migrates south to the Caribbean. The Wyoming toad (Anaxyrus baxteri) was common in and around the lakes of the high-elevation Laramie Plains in the 1950s, but underwent a major decline in the following decades. It is currently extinct in the wild. There is a nonreproducing population in Mortenson Lake National Wildlife Refuge, which is maintained with captive-reared individuals.

The Sierra Nevada Mountains The Sierra Nevada Mountains are located in the western United States between the Central Valley of California and the Great Basin. The vast majority lies in central and eastern California, although the Carson Range spur lies primarily in Nevada. The range is home to three major national parks (Yosemite, Sequoia, and Kings Canyon) and a number of smaller protected areas and national monuments. The Sierra Nevada bighorn sheep (Ovis canadensis sierrae) is now confined to these mountains. While not threatened, it appears to have historically ranged as far west as the California Coastal Ranges. An account of ‘wild sheep’ in the vicinity of the Mission San Antonio near Jolon, California, and the mountains around San Francisco Bay dates to around 1769. The Yosemite toad (Anaxyrus canorus) is confined to the wet mountain meadows of the Sierra Nevada Mountains in California, where it has undergone a serious decline in recent decades. The Sierra Nevada yellow-legged frog (Rana sierrae) was historically found throughout the Sierra Nevada Mountains but has undergone severe declines due to introduced predators, disease, and pollution. The species is now extirpated from Nevada, and only a small number of fragmented populations remain in California. The diabolical slender salamander (Batrachoseps diabolicus) is confined to a small area of north-central Califorina. The Kings River slender salamander (B. regius) is known only from two localities within Kings Canyon National Park in central California. The Kern Canyon slender salamander (B. simatus) is confined to a small area of central California largely within Sequoia National Forest. The Sequoia slender salamander (B. kawia), Kern Plateau slender salamander (B. robustus), and the relictual slender salamander (B. relictus) are each confined to a small area of south-central California. All are threatened by habitat disturbance. The Piute cutthroat trout (Oncorhynchus clarkii seleniris) is confined to two tiny streams in the eastern Sierra Nevada.

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The Pacific Coast Ranges The Pacific Coast Ranges are a series of mountain ranges stretching along the western coast of North America from Alaska south to northern and central Mexico. The Pacific giant salamander (Dicamptodon ensatus), along with its close relatives in the Pacific Northwest, are the largest of all salamanders that have a terrestrial stage. This species is found discontinuously in the damp coastal forests of northern California, where it is threatened in some areas by habitat destruction and degradation. The California tiger salamander (Ambystoma californiense) is found discontinuously in the coast ranges and surrounding foothills of west-central California, where it is rapidly declining.

The Olympic Mountains The Olympic Mountains are located on the Olympic Peninsula of western Washington state. Its western slopes are among the wettest places in North America. Most of the mountains are protected within the bounds of Olympic National Park and adjoining segments of the Olympic National Forest. The Olympic torrent salamander (Rhyacotriton olympicus) is confined to cold, high-elevation streams within the Olympic Mountains.

The Cascade Range The Cascade Range extends from southern British Columbia through Washington and Oregon to northern California. It includes both volcanic and non-volcanic mountains. The Cascade Mountains wolf (Canis lupus fuscus) was a cinnamon-coloured subspecies confined to the Cascade Range, where it became extinct in 1940. It has recently been replaced by a different subspecies of grey wolf. The Oregon slender salamander (Batrachoseps wrighti) is confined to the Cascade Mountains in north-central Oregon. The Cascade torrent salamander (Rhyacotriton cascadae) occurs in cold mountains streams and spring seepages on the western slopes of the Cascades in Washington and Oregon, where it remains fairly common.

The Klamath Mountains The Klamath Mountains are located in north-western California and south-western Oregon. The Siskiyou Mountains are the northermost and largest subrange of the Klamath Mountains. Storm’s salamander (Plethodon stormi) and the Scott Bar salamander (P. asupak) are each confined to a small area of the Siskiyou Mountains.

The California Coast Ranges The California Coast Ranges run parallel to the Pacific coast in north-central California, and consist of separate Northern Coast and Southern Coast ranges. Both have a predominantly Mediterranean climate.

Mountains and Highlands

The Santa Cruz Mountains

The Peninsular Ranges

The Santa Cruz Mountains are located in central and northern California, in the Southern Coast Ranges. The Santa Cruz long-toed salamander (Ambystoma macrodactylum croceum) is known only from the vicinity of a few isolated ponds within the Santa Cruz Mountains.

The Peninsular Ranges are a group of mountain ranges that stretch 1500 km from southern California to the southern tip of the Baja California Peninsula. Elevations range from 150 to 3300 m. The mountains are arid, with the western slopes of the more northerly ones dominated by montane chaparral and, higher up, by coniferous and mixed evergreen forests. The remaining areas are covered by desert and xeric scrub. The Baja California rock squirrel (Otospermophilus beecheyi atricapillus) is largely confined to areas near water holes within the Gigantas Sierra and Sierra de San Francisco, where it is threatened by hunting and habitat destruction.

The Santa Lucia Mountains The Santa Lucia Mountains are a rugged subrange located in coastal central California, at the southern end of the Southern Coast Ranges. The San Simeon slender salamander (Batrachoseps incognitus) and the lesser slender salamander (B. minor) are both confined to the Santa Lucia Mountains.

The Transverse Ranges The Transverse Ranges are a group of mountain ranges with an east–west orientation located in southern California. Most have a Mediterranean climate. Stebbins’ slender salamander (Batrachoseps stebbinsi) is confined to isolated areas of the Piute and Tehachapi Mountains.

The San Bernardino Mountains The San Bernardino Mountains are a high and rugged mountain range in southern California. Recreational development of the range began in the early twentieth century, when resorts were built around irrigation reservoirs. Since then, the mountains have been extensively engineered for transportation and water supply purposes, all of which have had a significant impact on habitats. The white-eared pocket mouse (Perognathus alticola) is divided into two subspecies separated by the San Gabriel Mountains. The San Bernardino white-eared pocket mouse (P. a. alticolus) is known only from the San Bernardino Mountains. Last collected in 1934, it may be extinct.

The San Gabriel Mountains The San Gabriel Mountains lie between the Los Angeles Basin and the Mojave Desert, and are within the Angeles National Forest. The San Gabriel slender salamander (Batrachoseps gabrieli) is confined to the San Gabriel Canyon system, typically between 1000 and 1500 m.

The Tehachapi Mountains The Tehachapi Mountains extend for approximately 65 km in southern California near the city of Los Angeles. They separate the San Joaquin Valley to the north-west with the Mojave Desert to the south-east, and as such serve as an important wildlife corridor. The Tehachapi white-eared pocket mouse (Perognathus alticola inexpectatus) is confined to the Tehachapi Mountains, where it is threatened by loss of habitat.

The Santa Rosa Mountains The Santa Rosa Mountains are a short mountain range located in southern California along the western side of the Coachella Valley. The sandstone night lizard (Xantusia gracilis) is confined to the Truckhaven Rocks area in the Anza-Borrego Desert State Park, on the south-eastern flanks of the Santa Rosa Mountains. The desert slender salamander (Batrachoseps major aridus) is known only from two localities on the east slope of the Santa Rosa Mountains.

The San Pedro Mártir Mountains The San Pedro Mártir Mountains (Sierra de San Pedro Mártir in Spanish) are located in north-western Mexico (Baja California). Mearns’ squirrel (Tamiasciurus mearnsi) is confined to the San Pedro Mártir Mountains.

The Laguna Mountains The Laguna Mountains (Sierra de la Laguna in Spanish) are located near the southern tip of the Baja California Peninsula (Baja Peninsula Sur). The peninsular mouse-eared bat (Myotis peninsularis) is confined to the southern part of the Baja California Peninsula, where it has been reported from all elevations but is relatively dependent upon caves for roosting.

The Colorado Plateau The Colorado Plateau is an area within western Colorado, north-western New Mexico, southern and eastern Utah, and northern Arizona. The majority is drained by the Colorado River and its tributaries, and much of the rest by the Rio Grande. Largely made up of high desert with scattered areas of forest, the Grand Canyon lies within its south-west corner. The Kaibab squirrel (Sciurus aberti kaibabensis) is a beautiful, tassel-eared form whose range lies entirely within the ponderosa pine forests of the Kaibab Plateau of Arizona. Fortunately, this area is protected within parts of Grand Canyon National Park and the Kaibab National Forest. In 1964 the total population was estimated at around 1000.

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The Appalachian Mountains The Appalachian Mountains are an ancient chain, weathered down by time, located in eastern North America. The plains and hill country to the west, running from Pennsylvania to Alabama, supports a relatively small ecoregion featuring a remarkable convergence of forest habitats from mixed deciduous in the lowlands to spruce-fir, thus supporting a tremendous diversity of species. The Appalachians are home to some 30 endemic species of lungless salamanders that live on the forest floor, concealed in the leaf litter. These include a number of forms found only on a single isolated mountaintop. Two subspecies of northern flying squirrel (Glaucomys sabrinus) are found in the southern Appalachians, where they suffered significant declines during the twentieth century. The Virginia flying squirrel (G. s. fuscus) has recovered somewhat in recent years, although the Carolina flying squirrel (G. s. coloratus) remains threatened by loss of habitat.

Carolina. The South Mountains grey-cheeked salamander (P. meridianus) is confined to the South Mountains of western North Carolina. The Shenandoah salamander (P. shenandoah) is known only from three isolated mountains (Hawksbill, The Pinnacles, and Stony Man) in Shenandoah National Park, Virginia. The Big Levels salamander (P. sherando) is confined to a small area in the vicinity of Big Levels, Virginia. The red-legged salamander (P. shermani) is known from a few localities in North Carolina and Tennessee. The Peaks of Otter salamander (P. hubrichti) is known from the Blue Ridge Mountains north-east of Roanoke, Virginia. The Cheoah Bald salamander (P. cheoah) is confined to a small area in western North Carolina. Weller’s salamander (P. welleri) is found patchily in western North Carolina. The dwarf black-bellied salamander (Desmognathus folkertsi) is a semi-aquatic species confined to a small area of northern Georgia.

The Appalachian Plateau

The Ozark Mountains

The Appalachian Plateau is a series of rugged, dissected plateaus located on the western side of the Appalachian Mountains. The Pigeon Mountain salamander (Plethodon petraeus) is confined to Pigeon Mountain in north-western Georgia. The Tennessee cave salamander (Gyrinophilus palleucus) is known from around two dozen cave sites in Tennessee and Alabama. The Cumberland Plateau is located in east-central Tennessee. The laurel dace (Chrosomus saylori) is a type of rare freshwater minnow confined to a few small streams on the Cumberland Plateau.

The Ozarks are located in northern Arkansas, southern Missouri, north-eastern Oklahoma, and south-eastern Kansas. The spring grotto salamander (Eurycea nerea) is confined to the southern Ozark Plateau of Missouri and adjacent northern Arkansas. The bluestripe darter (Percina cymatotaenia) is confined to six streams in the Osage and Gasconade river drainages of the northern Ozarks, south-central Missouri. The Ozark cavefish (Amblyopsis rosae) is confined to a few caves and wells of the Springfield Plateau, in the Ozark Mountains.

The Ridge-and-Valley Appalachians The Ridge-and-Valley Appalachians are a belt situated between the Appalachian Plateau and the Blue Ridge Mountains. They are characterized by long, even ridges and continuous valleys. The Berry Cave salamander (Gyrinophilus gulolineatus) is confined to a few subterranean localities in eastern Tennessee.

The Blue Ridge Mountains The Blue Ridge Mountains straddle the eastern flank of the Appalachians and stretch from southernmost Georgia to Pennsylvania. Two major national parks – Shenandoah National Park in the northern section and the Great Smoky Mountains National Park in the south – are contained within the region, along with a number of smaller protected areas. The Janaluska salamander (Eurycea junaluska) is confined to south-western North Carolina and south-eastern Tennessee. Several species of woodland salamander (Plethodon) are endemic to the Blue Ridge Mountains, where they are threatened by loss of habitat. The Blue Ridge grey-cheeked salamander (P. amplus) is confined to a few locations in North

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The Ouachita Mountains The Ouachita Mountains are located in western Arkansas and south-eastern Oklahoma. The Fourche Mountain salamander (Plethodon fourchensis) is confined to the Fourche and Irons Fork subranges. The Kiamichi salamander (P. kiamichi) is confined to the Round and Kiamichi subranges. The Sequoyah salamander (P. sequoyah) is confined to the Ouachita Mountains. All are threatened by loss of habitat. The Kiamichi shiner (Notropis ortenburgeri) and the rocky shiner (N. suttkusi) are both confined to upland streams within the Ouachita Mountains.

Miscellaneous Mountains and Highlands The Santa Catalina Mountains are located in the southwestern United States (southern Arizona). The Santa Catalina grey squirrel (Sciurus arizonensis catalinae) is confined to the Santa Catalina Mountains, where it is highly threatened by habitat destruction. The Manzano Mountains are located in the south-western United States (central New Mexico). The Manzano cottontail rabbit (Sylvilagus cognatus) is confined to high elevation conifer forests in the Manzano Mountains.

Lowland Forests

The Inyo Mountains are a short mountain range east of the Sierra Nevadas in eastern California. They separate Owens Valley to the west and the Saline Valley to the east. The Inyo Mountains salamander (Batrachoseps campi) is known only from scattered localities in the Inyo Mountains, where it lives along small, permanent desert springs and seeps. The Panamint Range is a short, rugged mountain range in the northern Mojave Desert of east-central California, and forms the western wall of Death Valley. Being a ‘sky island’ habitat, with more precipitation and temperature variation than the desert floor and hills, it is home to a number of endemic species. The Panamint alligator lizard (Elgaria panamintina) is a rare and secretive species known from comparatively few museum specimens since its discovery in 1958. The Spring Mountains are located in the south-western United States (southern Nevada). Palmer’s chipmunk (Neotamias palmeri) is confined to the Spring Mountains. The Edwards Plateau is located in the south-western United States (west-central Texas). Essentially a limestone uplift with numerous caves, the landscape is mostly savanna scattered with a few trees. The golden-cheeked warbler (Setophaga chrysoparia) occurs only in a small part of the Edwards Plateau in south-central Texas. The population is small and declining as a result of bush eradication, in spite of the fact that the bird is protected by law.

Lowland Forests During the last few centuries North America has witnessed one of the most violent exploitation of forests that has ever been seen. We are still so close to these drastic alterations that we can hardly appreciate their full significance. The rapid decrease of forests and the habitat changes in them have had serious effects not only on the wildlife but also on entire ecosystems. Many species have been wiped out or are on the verge of extinction. Fortunately, an increasing public as well as official governmental consciousness of the value of unspoiled habitats, rich in wildlife, has done much to ameliorate the situation in Canada and the United States during the last few decades. The ivory-billed woodpecker (Campephilus principalis) is traditionally divided into two subspecies. The nominate form, the northern ivory-billed woodpecker (C. p. principalis), is (or was) one of North America’s most impressive birds. Historically it occurred in low densities throughout the bottomland swamp forests of the south-eastern United States from the Carolinas, Georgia, and Florida west to Texas, and in the Mississippi Valley north as far as Illinois, where it was highly dependent upon dead and dying trees. Loss of habitat and shooting reduced its numbers, and DDT poisoning may also have been implicated in the disappearance. The last confirmed record was from north-eastern Louisiana in 1944, although in 1950 two ivory-bills were reportedly seen in northern Florida, and in 1960, six. In 1967 a trained ornithologist claimed to have seen it in the Big Thicket country of south-

eastern Texas, estimating the population at between 5 and 10 pairs. In 2004, it was reportedly rediscovered in the Big Woods region of eastern Arkansas. A thorough investigation of the available evidence, which included a number of sound recordings and a short, poor-quality video, were inconclusive, and subsequent searches of the area have revealed nothing more. There were also unconfirmed reports made by researchers along the Choctawhatchee River in Florida between 2005 and 2007, as well as from Louisiana between 2006 and 2008. In addition, in the years between the last confirmed sightings in the 1960s and the purported ones in the 2000s there were more than 20 others that are at least plausible. There are still large areas of undisturbed habitat available, not least of which being the coastal mangrove and inland hammock forests of south Florida, but unless or until hard evidence is obtained this subspecies must now be considered extinct. The other subspecies, the Cuban ivory-billed woodpecker (C. p. bairdii), may yet survive in south-eastern Cuba. Two subspecies of Carolina parakeet (Conuropsis carolinensis), a small endemic conure, formerly occurred in oldgrowth riverine and swamp forests of the eastern, midwestern, and plains states of America, making them the northernmost of all parrots. They seem to have shown a strong preference for deep cypress swamps, but this was not enough to save them from being shot indiscriminately and taken captive in large numbers. It is probable that habitat destruction and disease, although to a lesser degree, also played a role in their ultimate extermination. The birds’ range collapsed from east to west with settlement and clearing of the eastern and southern deciduous forests. The nominate form (C. c. carolinensis) was more a bird of the south-eastern coastal areas, ranging from the Carolinas down through Florida and as far west as southern Louisiana. It was rarely reported outside of Florida after the 1860s, and by the turn of the century was largely confined to swamp forests in the central part of the state. The last-known wild specimens, a flock of 13 birds, were seen (and a few killed) at Lake Okeechobee 1904. ‘Incas’, the last captive specimen, died at the Cincinnati Zoo on 21 February 1918, in the same cage as had ‘Martha’, the last passenger pigeon. The place where the aviary once stood now serves as a monument for both, and is listed as a national historic site. It has been claimed that a small number of parrots may have held on near Lake Okeechobee until the late 1920s, but this cannot now be verified. Reports after this from the Okefenokee Swamp of Georgia and the Santee River swamps of South Carolina are dubious. The Louisiana parakeet (C. c. ludovicianus) appears to have lived more inland that the nominate form, being found generally west of the Appalachians from New York to the Rocky Mountains and south as far as Texas. It was already becoming extremely rare by the mid-nineteenth century. The last confirmed sighting in the wild was in 1910, with total extinction occurring around 1912. Bachman’s warbler (Vermivora bachmanii) was last reported in 1988 and is most likely extinct. It is known to have

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The Nearctic Realm Figure 9.3 Ivory-billed woodpecker (Campephilus principalis), hand-coloured engraving. Male on the left, females at the top and right. (Credit: Arthur A. Allen and Paul P. Kellogg)

bred in Missouri, Arkansas, Kentucky, Alabama, and South Carolina, with additional non-breeding records from various other south-eastern U.S. states. The species wintered in Cuba and, occasionally, Florida. Loss of its riverine swampland habitat appears to have been the cause of the decline. The Pine Barrens tree frog (Dryophytes andersonii) is found sporadically across the coastal plains of the eastern

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United States, with populations in New Jersey, the Carolinas, and western Florida and adjacent Alabama. It is also known in Georgia from an old record of a single specimen. Although common where found and not currently considered to be seriously threatened, the disjunct nature of its distribution suggests a formerly much more widespread range.

Lowland Forests

Figure 9.4 A live captive Carolina parakeet (C. c. carolinensis) photographed around 1900. (Credit: Robert W. Shufeldt.)

Lowland Boreal Forests North America’s boreal forests are located just south of the tundra, covering most of inland Canada, Alaska, and parts of the northern continental United States. They are strikingly similar to those of the Eurasian Region, with the same general pattern of interspersed bogs, lakes, and rivers. Much of the biodiversity is also shared, although the forests of the boreal Nearctic Realm are much richer in species. The dramatic and tragic history of the plains bison (Bison bison bison) is well known and detailed elsewhere in this book, but the near-extinction and recovery of the wood bison (B. b. athabascae) is also of considerable interest. This northern subspecies – the largest terrestrial animal in North America – historically also had a wide range, extending throughout the boreal forests of Alaska, Yukon, western Northwest Territories, north-eastern British Columbia, northern Alberta, and north-western Saskatchewan. When the plains bison had become almost extinct through wanton exploitation, men turned to the wood bison. It could not withstand the tremendous persecution, and the number dwindled rapidly. It was practically extinct south of the Peace River by 1875, and by 1891 only about 300 remained in an area south of Great Slave Lake. In 1903–04 only 24 individuals were observed, and three years later, 33. In 1922 the Canadian government set aside a protected area, Wood Buffalo National Park, which included the entire habitat of the remaining herd. By 1929 the population had increased to about 1500, but at that time a serious mistake was made in the introduction into the reserve of no less than 6673 plains bison. As might have been expected, the two subspecies interbred freely and the wood

Figure 9.5 A living Bachman’s warbler photographed in 1958. (Credit: Jerry A. Payne.) (A black and white version of this figure will appear in some formats. For the colour version, please refer to the plate section.)

bison disappeared as a pure breed. Fortunately, in 1957 another small herd of some 200, genetically pure animals was discovered in the north-western part of Wood Buffalo National Park, where they had been isolated by swamps. By 1965 only about 100 were left, however, and so 18 were transplanted to Fort Providence north of the Mackenzie River in 1963, in order to establish an independent herd. In 1965 another 43 were captured, destined this time for Elk Island National Park in Alberta. All three of these populations have since prospered with careful management, and additional reintroductions have since taken place elsewhere in Alberta as well as in the Northwest Territories, Yukon, British Columbia, and Manitoba. In 2006 an outherd was established in north-eastern Siberia, where the related steppe bison (B. priscus) died out over 6000 years ago, and there are currently plans to establish populations in Alaska. All told, about 7000 wood bison now live in the wild. The caribou (Rangifer tarandus), previously discussed in this volume, is also found in Nearctic boreal forests. The

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The Nearctic Realm

woodland caribou (R. t. caribou), the largest subspecies, occurs across southern Canada and the north-western United States, where it is nevertheless considered vulnerable. Osborn’s caribou (R. t. osborni) is confined to an area of British Colombia.

Lowland Broadleaf and Mixed Forests About 40 per cent of the United States mainland was originally covered by virgin deciduous forest. In 1600 they were still intact except for some local tree removal in the north-eastern states and in Virginia. Today, primeval deciduous forests cover no more than a tiny fraction of this, chiefly in the eastern United States and southern Canada. The red wolf (Canis rufus) is a controversial form, variously considered to be either a species in its own right (with three subspecies), a subspecies of the grey wolf (C. lupus), or merely a grey wolf/coyote hybrid. Historically these animals were found widely throughout the woodlands of the eastern United States from the Atlantic to the Gulf of Mexico, but were nearly driven to extinction by the mid-nineteenth century due to human persecution, habitat destruction, and extensive hybridization with coyotes (C. latrans). By the late 1960s they survived only in small numbers in the coastal prairies and marshes of western Louisiana and eastern Texas. Fourteen of these survivors were taken into captivity in the 1970s to establish a breeding programme, which has been a success. The red wolf was officially declared extinct in the wild in 1980. In 1987 captives were released into the Alligator River National Wildlife Refuge in North Carolina. A second release, since reversed, took place two years later in the Great Smoky Mountains National Park. In all, 63 animals were introduced to the wild between 1987 and 1994. The population rose to as many as 100–120 in 2012, but declined to just 35 by 2018 as a result of widespread shooting by local hunters, who frequently mistake the wolves for coyotes. In early colonial days the eastern elk (Cervus canadensis canadensis) ranged throughout the vast forests of southern and eastern Canada and the eastern United States as far west as the Mississipi. It was everywhere common, but always much hunted. Gradually it declined and was at last exterminated. In 1851 John James Audubon noted that a few elk could still be found in the Allegheny Mountains, but that they were virtually gone from the remainder of their range. In Pennsylvania the last one was killed in 1877. A few populations held on till much later in the western parts of the range, where human pressure was less severe. It was found in Minnesota north of Lake Superior as late as 1885, and still existed in Wisconsin in 1892, but (reports from northern Ontario as recently as the 1980s notwithstanding) it was almost certainly extinct by the end of the nineteenth century. The wood turtle (Glyptemys insculpta) is a semi-aquatic species still found over a wide area of the north-eastern United States and parts of southern Canada. It has declined greatly in number due to loss of habitat and overcollection for the international pet trade.

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Lowland Coniferous Forests Areas of temperate coniferous forest once covered much of North America, mainly in the northern areas. Of special note are the Pacific temperate rainforests that lie along the northwestern coast of North America from Prince William Sound in Alaska, through western British Colombia, Washington, Oregon, and northern California. Unique in being rainforests dominated by conifers, this extraordinary habitat is produced by a combination of maritime moist climate, mild temperature, high rainfall, and the huge mass of organic matter accumulated and produced by the forests themselves. Altogether the region contains a quarter of the world’s remaining temperate rainforests, and is rich in animal life. Sadly, they are highly threatened by logging activity. Kirtland’s warbler (Dendroica kirtlandii) is a small songbird that very nearly went extinct during the mid-twentieth century. While it may have once been much more widespread, by the 1960s its breeding range had been reduced to a small area of central Michigan, from where the 1000 or so birds that made up the total population migrated to the Bahamas and the Turks and Caicos Islands for the winter. The species has extremely specialized habitat requirements for breeding, namely large areas of dense, young jack pine historically created by forest fires. A stand of growing trees satisfies these warblers for only 10–15 years, after which they move to another area where the trees are smaller. Intensive conservation measures including controlled burning and timber harvesting now produce the exact habitat that the birds need, and as a result they have been able to expand their breeding range into parts of southern Ontario and Wisconsin. The species is now considered to be out of immediate danger.

South-eastern Conifer Forests The south-eastern conifer forests are located in Louisiana, Mississippi, Alabama, Florida, and Georgia. It represents the largest area of coniferous forest east of the Mississippi River. The so-called Florida panther is now generally thought to be an isolated, remnant population of North American cougar (Puma concolor couguar), although many scientists continue to recognize it as a distinct subspecies in its own right. As late as the end of the nineteenth century cougars were still common throughout the Florida peninsula as well as in adjacent areas of Georgia, Alabama, and Louisiana. However, constant persecution led to a general decrease in numbers and local extermination until it was gone everywhere except in south Florida, where it lives primarily in protected areas such as Everglades National Park, the Florida Panther National Wildlife Refuge, the Picayune Strand State Forest and the Big Cypress National Preserve. In the 1970s the total wild population had declined to around 20, but this has increased to as estimated 230 by 2017. The gopher tortoise (Gopherus polyphemus) is an important keystone species owing to the fact that hundreds of others use its burrows. Long hunted for food across the south-eastern

Lowland Grasslands, Savannas, and Shrublands

United States, today the main threat is habitat destruction and capture for the pet trade. The Louisiana pine snake (Pituophis ruthveni) historically occurred in parts of west-central Louisiana and extreme eastcentral Texas. It has been reduced to a few isolated areas due to loss of habitat and other factors. The Rim Rock centipede snake (Tantilla oolitica) is confined to southernmost Florida, where it is threatened by habitat destruction. The dusky gopher frog (Lithobates sevosus) historically occurred from eastern Louisiana and southern Mississippi to the Mobile River delta in Alabama. Owing to fungal disease and other factors it has declined catastrophically, and by the beginning of the present century was known only from Glen’s Pond in Desoto National Forest, Mississippi. It has since been reported from two additional sites in Mississippi, although it is unclear whether these represent stable populations. Ainsworth’s salamander (Plethodon ainsworthi) is known only from two specimens collected in Jasper County, Mississippi in 1964. It is most likely extinct. The Red Hills salamander (Phaeognathus hubrichti) is confined to the Red Hills regions of south-central Alabama, where it inhabits forested ravines. It is threatened by habitat destruction. Two other species of salamander of the genus Ambystoma are endemic to pine flatwoods habitat in the south-eastern coastal plain. The reticulated flatwoods salamander (A. bishopi) is restricted to Eglin Air Force Base in north-western Florida, while the frosted flatwoods salamander (A. cingulatum) occurs in fragmented populations in South Carolina, Georgia, and Florida.

of Mexico. This type of habitat burns easily, almost exploding into flame, and fires have long played an important role in forming its development. The southern California kit fox (Vulpes macrotis macrotis) was extensively hunted for its fur as well as poisoned during campaigns against coyotes. The last known individual was trapped on the San Jacinto plain in 1903. Stephens’ kangaroo rat (Dipodomys stephensi) is found in the San Jacinto Valley and adjacent areas of southern California, with the largest known population on the Warner Ranch near Lake Henshaw. The Morro Bay kangaroo rat (D. heermanni morroensis) is confined to the south side of Morro Bay, southern California.

Lowland Grasslands, Savannas, and Shrublands

Florida sand pine scrub is an arid subtropical forest region found throughout Florida. It occurs on coastal and inland sand ridges, and is characterized by an evergreen plant community dominated by sand pine shrubs and dwarf oaks. The Florida black wolf (Canis lupus floridanus) was endemic to Florida, where it became extinct in 1934 due to habitat destruction and hunting. The Florida deermouse (Podomys floridanus) remains fairly widespread but is everywhere under threat by habitat destruction. The Florida scrub jay (Aphelocoma coerulescens) has declined significantly since the mid-nineteenth century, and most rapidly since 1950. It is now confined to a few fragmented populations. The Florida sand skink (Plestiodon reynoldsi) is confined to high sandy ridges in central Florida.

A wide variety of temperate and subtropical grassland types once covered a broad belt of central North America, with outlying regions in north-western, south-western, and southern United States and northern Mexico. The black-footed ferret (Mustela nigripes) may have once been the rarest mammal in North America, and indeed very nearly went extinct by the late 1980s. This large weasel is entirely dependent upon prairie-dog towns, which provide it both with prey and den sites. Thus, its former range coincided with that of the ‘prairie dogs’ (Cynomys), that is, the grasslands steppes between southern Canada and northern Mexico. Destruction of the prairie dogs as agricultural pests by means of poisoning and the resulting elimination of prairie-dog burrows, as well as of the original grasslands themselves, was the cause of this species’ decline. By 1970 it had already become extremely rare. In 1981 a small, remnant population of