INTRODUCTION This book is intended to be read and enjoyed (in parts) by a wide range of people with a varying knowledge base. The book divides into 3 main sections starting with an overview of forest ecology. This is followed by a field guide to all the known from and ends with a guide to 33 significant wildlife sites. The intricate web of relationships in forest ecology is emphasised with numerous cross-references and ecological themes such as mimicry and seed dispersal. Some misconceptions are addressed and some new speculative ideas are introduced. This cannot possibly be comprehensive in the space available and numerous references are given to spur debate and further research. For all readers we hope this book will open some doors. Taxonomic changes There have been a large number of taxonomic changes since the publication of a Field Guide to the Mammals of Borneo by Payne et al (1985). Mammalian is moving from the traditional Biological Concept (BSC) to a Phylogenetic Species Concept (PSC) approach, resulting in the ‘discovery’ of many cryptic species. By definition cryptic species cannot be illustrated in a Field Guide, so this accounts for the 9 mammals not illustrated in this book. We have attempted to include the latest taxonomy but no doubt even this guide will soon be out of date so fast is DNA research developing (pp. 108, 168).

MATTERS FOR DEBATE CONCERNING BORNEO’S ECOLOGY 1 That if a is extinct or there is no fossil record it never occurred in Borneo (p. 18). 2 That extinct mammals are no longer relevant to forest ecology in Borneo (p. 38). 3 That Borneo has a ‘perhumid’ ever wet aseasonal (non seasonal) climate (p. 12). 4 That because the climate is aseasonal mammals breed throughout the year (p. 10). 5 That mass fruiting (masting) is an unusual phenomenon confined to Sunda Shelf islands (p. 44). 6 That small areas of protected forest e.g. Sepilok, Lambir, Matang can preserve biodiversity (p. 168). 7 Relative tree species abundance results from random events (Hubble’s Neutral Theory) (pp. 32, 46, 298). 8 That speciation in trees is largely the result of abiotic factors e.g. soils, water and sunlight (p. 46).

Speculation and untested hypotheses The book contains a considerable number of speculative ideas which I hope will stimulate further enquiry. Ecological themes often untreated in prior works on rainforest ecology include: 1 Mimicry and aposematic (danger warning) markings on many Bornean mammals (pp. 26, 157, 260). 2 Anachronistic fruit and the relevance of the size and shape of rainforest seeds (pp.34, 36, 38, 290, 296). 3 Changes of forest with altitude and recalcitrant and orthodox seeds (pp.45, 142, 212, 232, 237, 245). 4 Both the causes of and limits to speciation in tropical trees (p. 46). 5 Specialist langur-dispersed trees with sugar-free fruit and tiny seeds (pp. 176, 224). 6 Why small bark-eating are so numerous (p.195, 207). 7 Poisonous mammals and toxins sequestered from insects and plants (pp.118, 138, 150, 160, 272). Plant diversity in Borneo Bearing in mind the reader new to botany a few plant families e.g. the bananas, beans, dipterocarps, figs, oaks and palms are referred to repeatedly, whilst 90% of Borneo’s 15,000 plants and 210+ plant families are hardly mentioned. This is deliberate. This book is not intended to confuse with quantity and variety but to provide key examples of plant/mammal relationships such as pollination, seed dispersal, seed predation and herbivory, common to all Bornean plants. Sundaland Mammals As emphasized throughout the book, for most of the last 2.6 million years Borneo has been part of a much larger landmass known as Sundaland most of which has a similar climate and similar ecology. I have therefore freely quoted from the results of research into mammals elsewhere in Sundaland particularly the Malay Peninsula (Malaya) which is a zoo-geographical term including the political units of both West (Semananjung) and a portion of southern Thailand. Bats There is a revolution in bat taxonomy currently underway in Borneo and throughout SE Asia. As pointed out by Francis et al. (2010), SE Asia probably hosts double the species of bats currently listed. In the space and time available it was impossible to do justice to Borneo’s bats in this book. A specialist field guide to the bats of Borneo is long overdue. Errors and Omissions Any corrections, suggestions for improvements and significant new records should be emailed to [email protected]. An up-to-date list of Bornean mammals can be downloaded from Borneomammals.com. Sizes HB: Length from nose to base of tail. Where only one measurement is given this is an average, which may vary up to 15%.

6 Conservation Those interested in the continued preservation of diversity in Borneo need to work in partnership with governments, logging concessions, oil-palm companies and local NGOs, always bearing in mind that projects will only succeed where they are fully endorsed by and of benefit to the local inhabitants both in the short and long term. In particular working to implement FSC certification in all logging concessions and the standards supported by the Roundtable on Sustainable Palm Oil in all oil-palm plantations should be a baseline target. However, only the protection of large areas of virgin forest can preserve diversity. IUCN Red List The Red List designations are not used in this book because taxonomy is moving so fast that the Red List is outdated almost as soon as it is published. Because legally protected in Borneo are usually listed based on the Red List, this is causing major problems in legal enforcement against poachers. One solution would be for the Bornean states to enact laws that legally protect all mammals and only list a few exceptions such as pigs. Borneo’s most threatened large mammals (2016) Total species of land mammals (247) of which (100) are bats and (63) are endemic to Borneo. Mammals almost extinct in Borneo; Sumatran Rhino, Langur, Kutai Langur, Abbot’s Gibbon. Mammals seriously threatened with extinction in Borneo, Hose’s Langur, Banteng. Mammals with collapsing populations due to human activities, Proboscis Monkey, Orangutan, all gibbons, all langurs, Tufted Ground , Banded Civet, Pangolin. Common Names Should be simple and self explanatory. Mostly I have used the names in Medway (1978) and Payne et al. (1985). I have only changed names where it appeared necessary to make memory easier or for ease of understanding. The English names of some new splits from HBMW (2014) have been replaced with geographic names to simplify matters for the reader, e.g. Sarawak Langur (p. 171), Grey Langur (p. 164). ABBREVIATIONS AND MALAY PLACE NAMES (see also Glossary p. 398) KYA Thousands of years ago LGM Last Glacial Maximum M YA Millions of years ago YBP Years Before Present Bukit Hill Kg Kampong or village Sg Sungai or River Gng Gunung or Mountain CONVERSION FACTORS 100 mm 3.93" 1 ha 10,000 sq m 2.59 sq km 1 sq mile 1m 3.28' 100 ha 1 sq km 1 sq km 247.1 acres 10 km 6.2 miles 1 ha 2.47 acres 1 sq mile 259 ha

KEY TO DISTRIBUTION MAPS The distribution maps show the original mammal distributions based on preferred altitude and original , not the fine detail of current fragmented distributions.

Throughout Endemic resident Coastal Lowland Lowland and Borneo throughout hills

Hills and Mountains Hills and slopes Rivers and Local distribution mountains alluvial habitat

7 HOSES’S CIVET Hemigale hosei Illustration by Joseph Smit of the type specimen collected by Hose on Gng Dulit in NW Sarawak, PZSL 1892 (pp. 242 and 270).

THE TOP TEN BORNEAN MAMMALS AND WHERE TO SEE THEM (Map inside back cover) Bornean Orangutan Sepilok, Kinabatangan, Danum, Tabin, Semengoh, Tg Puting p. 188 Proboscis Monkey Bako, Bay, Labuk Bay, Klias p. 174 Clouded Leopard Tabin, Danum, Kinabatangan, Deramakot p. 276 Sumatran Rhino Tabin p. 294 Pygmy Elephant , Tabin, Danum, Deramakot p. 288 North Borneo Gibbon Danum, Tabin, Deramakot p. 184 Red Langur Sepilok, Gomantong, Danum, Tawau Hills, Sabangau p. 162 Hose’s Langur Ulu Temburong, Brunei p. 164 Sun Bear Danum, Sepilok, Sg Wain p. 246 Bearded Pig Bako, Danum, Kinabatangan, Tabin p. 300

8 SIXTY-THREE ENDEMIC MAMMALS AND WHERE TO SEE THEM

SUNDA TAILLESS FRUIT BAT Megaerops ecaudatus Tasek Merimbun, Brunei p. 94 BORNEAN FRUIT BAT Aethalops aequalis Kinabalu, Gng Penrissen p. 92 DAYAK ROUNDLEAF BAT Hipposideros dyacorum Danum, Mulu, Matang p. 108 RED-BROWN PIPISTRELLE Hypsugo kitcheneri Sepilok, Ulu Temburong p. 118 VORDERMAN’S PIPISTRELLE Hypsugo vordermanni Samunsam, Tg Puting p. 118 GOMANTONG MYOTIS Myotis gomantongensis Gomantong p. 130 BORNEAN WHISKERED MYOTIS Myotis borneoensis Sepilok, Lambir p. 132 BLACK ater Kiau Gap, Kinabalu p. 138 HOSE’S PIGMY SHREW Suncus hosei Sepilok, Ulu Ulu p. 138 BORNEAN SHREW foetida Tabin, Matang p. 138 KINABALU SHREW Crocidura kinabauensis Kinabalu Park HQ p. 138 BORNEAN WATER SHREW Chimarrogale phaeura Kinabalu streams p. 138 SMOOTH-TAILED TREESHREW melanura Park HQ p. 146 montana Summit trail, Kinabalu p. 146 STRIPED TREESHREW Tupaia dorsalis Matang, Gng Penrissen p. 146 Tupaia gracilis Danum, Tabin, Mulu p. 144 PLAIN TREESHREW Tupaia longipes Danum, Tabin, Mulu p. 144 PAINTED TREESHREW Tupaia picta Lambir, Tasek Merimbun p. 148 SPLENDID TREESHREW Tupaia splendidula Tg Puting, Sabangau p. 148 KAYAN SLOW LORIS Nycticebus kayan Malai Basin, Kelabit Hghlds p. 160 BORNEAN SLOW LORIS Nycticebus borneanus Sabangau p. 160 RED LANGUR Presbytis rubicunda Sepilok, Sabangau p. 162 HOSE`S LANGUR Presbytis hosei Ulu Temburong p. 164 SABAH GREY LANGUR Presbytis sabana Tabin p. 164 KUTAI LANGUR Presbytis canicrus Wehea Community Forest p. 164 WHITE-FRONTED LANGUR Presbytis frontata Lanjak Entimau, Sg Wain p. 164 SARAWAK LANGUR Presbytis chrysomelas Samunsam, Maludam p. 170 NATUNA LANGUR Presbytis natunae Bunguran Besar, Natunas p. 168 PROBOSCIS MONKEY Nasalis larvatus Bako, Kinabatangan p. 174 ABBOTT’S GIBBON Hylobates abbotti Samunsam p. 184 NORTH BORNEO GIBBON Hylobates funereus Danum, Tabin p. 184 MÜLLER’S GIBBON Hylobates muelleri Sg Wain p. 184 WHITE-BEARDED GIBBON Hylobates alibarbis Tg Puting, Sabangau p. 184 BORNEAN ORANGUTAN Pongo pygmaeus Sepilok, Tg Puting p. 193 KINABALU SQUIRREL baluensis Kinabalu Park HQ p. 195 EAR-SPOT SQUIRREL Callosciurus adamsi Sepilok p. 202 BROOKE`S SQUIRREL brookei Kinabalu Park HQ p. 200 JENTINK`S SQUIRREL Sundasciurus jentinki Kinabalu Summit Trail p. 200 BORNEAN BLACK-BANDED SQUIRREL Callosciurus orestes Kinabalu Park HQ p. 204 BORNEAN MT GROUND SQUIRREL Dremomys everetti Kinabalu Summit Trail p. 212 FOUR-STRIPED GROUND SQUIRREL hosei Tawau Hills p. 212 WHITEHEAD`S PIGMY SQUIRREL whiteheadi Kinabalu Park HQ p. 206 BORNEAN PIGMY SQUIRREL Exilisciurus exilis Danum, Matang,Tg Puting p. 206 SCULPTOR SQUIRREL Glyphotes simus Crocker Rng, Malaiau Basin p. 208 TUFTED GROUND SQUIRREL Rheithrosciurus macrotis Malaiu Basin, Tawau Hills p. 214 HOSE`S PIGMY FLYING SQUIRREL Petaurillus hosei Sepilok, Tasek Merimbun p. 216 EMILY’S FLYING SQUIRREL Petaurillus emiliae Mulu p. 216 KINABALU baluensis Panar Laban, Kinabalu p. 230 BORNEO GIANT RAT Sundamys infraluteus Kinabalu Park HQ p. 230 MOUNTAIN Maxomys alticola Kinabalu Park HQ p. 230 LONG-TAILED MOUNTAIN RAT rapit Kinabalu p. 232 BAEODON MAXOMYS Maxomys baeodon Sepilok, Danum, Maliau p. 236 CHESTNUT-BELLIED MAXOMYS Maxomys ochraceiventer Maliau, Gng Penrissen p. 236 LARGE SUNDA TREE-MOUSE Chiropodomys major Sepilok, Matang p. 240 GREY-BELLIED SUNDA TREE-MOUSE Chiropodomys muroides Kinabalu, Poring p. 240 BORNEAN SUNDA TREE-MOUSE Chiropodomys pusillus Poring, Mulu p. 240 EMMON’S TREE-RAT Pithecheirops otion Danum DVFC p. 242 RANEE MOUSE Haeromys margarettae Sepilok, Gng Penrissen p. 242 BORNEAN PORCUPINE Thecurus crassispinis Danum. Tabin, Tawau Hills p. 244 BORNEAN STRIPED PALM CIVET Arctogalidia stigmatica Poring, Danum, Tabin p. 264 HOSE`S CIVET Hemigalus hosei Pulong Tau, Crocker Range p. 270 BAY CAT Felis badia Danum. Tabin, Tawau Hills p. 282 BORNEAN YELLOW MUNTJAC Muntiacus atherodes Danum. Tabin, Tawau Hills p. 310

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