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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 mammals known from Borneo 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 taxonomy is moving from the traditional Biological Species 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 We have attempted to include the latest taxonomy but no doubt even this guide will soon be out of date so fast is DNA research developing (pp. 108, 168).

INTRODUCTION

MATTERS FOR DEBATE CONCERNING BORNEO’S ECOLOGY1 That if a mammal 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 squirrels 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 Malaysia (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 info@borneomammals.com. 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%.

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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 animals 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, Sarawak 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 Squirrel, 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 primate splits from HBMW (2014) have been replaced with geographic names to simplify matters for the reader, e.g. Sarawak Langur (p. 171), Sabah Grey Langur (p. 164).

ABBREVIATIONS AND MALAY PLACE NAMES (see also Glossary p. 398)

KYA Thousands of years ago LGM Last Glacial Maximum

MYA Millions of years ago YBP Years Before Present

Bukit Hill Kg Kampong or village

Sg Sungai or River Gng Gunung or Mountain

Throughout Borneo

Endemic resident throughout

Coastal Lowland Lowland and hills

Hills and mountains

Mountains Hills and slopes Rivers and alluvial habitat

Local distribution

CONVERSION FACTORS

100 mm 3.93" 1 ha 10,000 sq m 2.59 sq km 1 sq mile1m 3.28' 100 ha 1 sq km 1 sq km 247.1 acres10 km 6.2 miles 1 ha 2.47 acres 1 sq mile 259 ha

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

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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, Brunei Bay, Labuk Bay, Klias p. 174

Clouded Leopard Tabin, Danum, Kinabatangan, Deramakot p. 276

Sumatran Rhino Tabin p. 294

Pygmy Elephant Kinabatangan River, 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

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).

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