Ecology of Sulawesi

THE ECOLOGY OF SULAWESI

THE ECOLOGY OF INDONESIA SERIES

VOLUME IV

THE ECOLOGY OF INDONESIA SERIES

Volume IV: The Ecology of Sulawesi

Other titles in the Series

Volume I: The Ecology of Sumatra

Volume II: The Ecology of Java and Bali

Volume III: The Ecology of Kalimantan

Volume V: The Ecology of Nusa Tenggara and Maluku

Volume VI: The Ecology of Irian Jaya

Volume VII: The Ecology of the Indonesian Seas

Produced by

Environmental Management Development in

Indonesia Project, a cooperative project of the

Indonesian Ministry of the Environment

and

Dalhousie University, Halifax, Nova Scotia

under the sponsorship of the

Canadian International Development Agency

The Ecology of

Sulawesi

TONY WHITTEN

MUSLIMIN MUSTAFA

GREGORY S. HENDERSON

PERIPLUS

First Edition © 1987 Gadjah Mada University Press

First Periplus Edition © 2002

All rights reserved

Published by Periplus Editions (HK) Ltd.

ISBN: 978-1-4629-0507-2 (ebook)

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Table of Contents

Foreword ix

Acknowledgements x

Introduction xiv

Chapter 1 Physical, biological and human background 1

Geology 1

Geological history 1

Volcanoes 7

Minerals 11

Soils 14

Climate 16

Palaeoclimate 16

Present climate 20

Vegetation 29

Palaeovegetation 29

Present vegetation 30

Fauna 34

Palaeofauna 34

Present fauna 37

Endangered species 50

Biogeography 54

Background to biogeography 54

Wallace's Line 59

Biogeographical differences within Sulawesi 68

People of Sulawesi 72

Prehistory 72

Impacts of prehistoric man 79

History 82

Present-day people 88

Present state of natural ecosystems 91

Chapter 2 Seashores 105

Introduction 105

Physical conditions 105

Tides 105

Surface currents 108

Salinity 109

Temperature 110

Dissolved oxygen and nutrients 110

Sediment 111

Oxygen within the sediment 112

Bacteria 113

Adaptations of the fauna 113

Mangrove forest vegetation 116

Composition 117

Zonation 124

Biomass and productivity 131

Other coastal vegetation 136

Pes-caprae formation 136

Barringtonia formation 137

Rocky shores 140

Fauna of sediment beaches 140

Open area communities 140

Shorebirds 144

Turtles 151

Maleo birds 155

Seabirds 159

Invertebrates of mangrove forest 162

Terrestrial fauna of mangrove forest 174

Fauna of beach forests, particularly coconut crabs 175

Fauna of rocky shores 181

Introduction 181

Functions and products of mangrove forest 185

Tambak fishponds 187

Mangrove forest management 192

Chapter 3 Estuaries, seagrass meadows and coral reefs 195

Estuaries 195

Water characteristics 195

Fauna 196

Primary productivity 201

Seagrass meadows 201

Seagrasses 201

Reproduction 206

Biomass, productivity and decomposition 207

Effects of development 208

Dugongs 210

Coral reefs 212

Importance and species richness 212

Structure and formation 213

Reef invertebrates 219

Reef fish 232

Reef algae and herbivores 238

Productivity and plankton 241

Causes of coral death and reef destruction 241

Coral reef fisheries 243

Coral reef survey techniques 246

Chapter 4 Freshwater ecosystems 257

Introduction 257

Lakes and rivers 258

Physical features 262

Water inputs and outputs 264

Water chemistry 266

Plants 268

Macrophytes 268

Phytoplankton 283

Fungi, bacteria and blue-green algae 284

Fauna 285

Zooplankton 286

Macro-invertebrates 287

Schistosomiasis and echinostomiasis of the Lindu valley 293

Fish 298

Aquatic reptiles 301

Water birds 305

Malili lakes system 308

Physical patterns in lakes 310

Temperature 310

Oxygen 311

Nutrients and conductivity 312

Light penetration 314

Stability 315

Biotic patterns in lakes 316

Physical patterns in rivers 317

Discharge 317

Shear stress 319

Riverbed particle size 320

Temperature 321

Dissolved oxygen and mineral nutrients 321

Biotic patterns in rivers 323

Current 323

Substratum 324

Temperature and dissolved

oxygen 324

Mineral nutrients 325

Biotic factors 325

Energy flow 325

Benthos dynamics 328

Fisheries 330

Management of macrophytes 336

Impacts of development 340

Industrial, domestic and agricultural pollution 340

Poisons, bombs and electric shocks 341

Forest clearance 342

Chapter 5 Lowland forests 343

Diversity 343

Structure and components 346

Characteristics 346

Layering 347

Basal area and biomass 350

Roots 350

Buttresses and trunks 351

Climbing and creeping plants 353

Epiphytes and epiphylls 356

Dynamics 359

Succession and growth cycle 359

Litterfall, nutrient cycling and productivity 365

Flower, fruit and leaf production 366

Herbivory 369

Seed dispersal and predation 379

Composition 385

Composition of mature-phase forest 385

Composition of pioneer- and building-phase forest 394

Animal communities 400

Soil and litter communities 400

Forest floor community 404

Lower and upper canopy communities 415

Comparison of mature- and pioneer-phase faunas 433

The effects of opening forest 434

General effects 434

Wider implications of forest conversion 437

Effects of selective logging on the forest 438

Effects of selective logging on soil and hydrology 440

Effects of selective logging on the fauna 442

Chapter 6 Specific lowland forest types 445

Introduction 445

Peatswamp forest 445

Formation and location 445

Vegetation 446

Fauna 449

Freshwater swamp forest 449

Physical conditions 449

Vegetation 450

Riverine forest 454

Vegetation 454

Fauna 457

Forest on ultrabasic soils 457

Soils 457

Vegetation 460

Fauna 466

Forest on limestone 468

Physical conditions 468

Soils 472

Hydrology 472

Vegetation 474

Effects of disturbance 478

Fauna 479

Monsoon forest 480

Vegetation 480

Fauna 486

The Palu valley: past and future 487

Chapter 7 Mountains 489

Introduction 489

Climate 489

Temperature 489

Relative humidity 491

Clouds 492

Rainfall 492

Ultra-violet radiation 493

Soils 494

Vegetation 495

Structure 495

Zonation 501

Characteristic plants 503

Selected mountains 511

Pollination and dispersal 515

Biomass and productivity 517

Mineral cycling 518

Volcanoes 520

Animals and their zonation 523

Invertebrates 523

Birds 526

Mammals 528

Effects of disturbance 533

Chapter 8 Caves 535

Introduction 535

Cave formation 539

Temperature, humidity and carbon dioxide 543

Characteristic animals and food chains 543

Effects of darkness 544

Echo-location 546

Roof community 549

Swiftlets 553

Floor community 553

Differences within and between caves 561

Effects of disturbance 561

Chapter 9 Agroecosystems 567

Characteristics 567

Swidden, shifting and intensive agriculture 570

Pest ecology and control 573

Rice fields 577

Introduction 577

Rice fields as an ecosystem 579

Insect pests on rice 581

Rats and their control 583

Rice field birds 585

Coconut and clove 588

Coconut 589

Clove 596

Cotton 598

Chapter 10 Urban ecology 601

Introduction 601

Gardens and streets 602

Vegetation 602

Birds 608

Bats 612

Walls 614

Ditches 617

Houses 620

Spiders and ants 620

Geckos 620

Possible urban ecology studies 621

Chapter 11 Resources and the future 625

Forestry 625

Watershed management 628

Implications of Island Biogeographic theory 631

Extinction and conservation 633

Genetic resources 637

Future scenarios 638

Drawing the line 638

Appendices 641

  A. Interpretation of soil analysis data 641

  B. Plant family abbreviations used in the text 642

  C. Key to the trees of mangrove and estuarine areas 644

  D. Key to seagrasses 646

  E. Keys to the submerged and floating freshwater macrophytes 648

  F. Key to tree ferns 652

  G. Key to termites 654

  H. Key to mudskippers 656

  I. Key to toads and frogs 658

  J. Key to house geckos 661

  K. Key to parrots 662

  L. Key to bat families 663

  M. Key to fruit bats 664

  N. Data sheet for coral reef monitoring 666

Notes 669

Bibliography 681

Index 727

Foreword to

the first edition (1987)

Indonesia, with its large population and vast and varied natural resources, must strive for economic development at the same time as protecting and enhancing the environment. Development must involve a harmonious relationship between Man and God, Man and his fellow man, and Man and Nature.

This book has been written as one important element in the incorporation of environmental thinking into development activities on Sulawesi. It provides important ecological information which will assist government planning agencies and project developments in including ecological considerations in development activities. The Ecology of Sulawesi is part of a series of books on the ecology of Indonesia. The first book in the series, The Ecology of Sumatra, was published in 1984 and is now in its third printing. A continuing demand for books in the series is evident.

The Ecology of Sulawesi has been written with the involvement of scientists from the Centre for Resource and Environmental Studies at Hasanuddin University, Ujung Pandang. The books in this Ecology of series serve not only to provide basic information on the environment. The process of preparation of the books also allows for training and technology transfer in environmental research between the authors and the junior scientists at the university environmental study centres.

We hope that scientists will be encouraged by this book to pursue further research on the ecological conditions in Sulawesi, and to apply the results of this continuing research to solutions to the environmental challenges posed by development in Sulawesi.

Emil Salim

Minister of State for Population and Environment

Republic of Indonesia

Acknowledgements

to the first edition (1987)

This book was produced within the Environmental Management Development in Indonesia (EMDI) Project, implemented by the Indonesian Ministry of State for Population and Environment (KLH) and the School for Resource and Environmental Studies (SRES) at Dalhousie University, Halifax, Nova Scotia, Canada. The project is funded by the Canadian International Development Agency (CIDA). The commitment to the publication of The Ecology of Sulawesi of those people involved in the administration of the project is much appreciated and special thanks are due to Koesnadi Hardjasoemantri, Sjafran Sjamsuddin (KLH, Jakarta), Arthur Hanson, Geoffrey Hainsworth, and George Greene (SRES, Halifax). The book has been born out of the experience of producing The Ecology of Sumatra which was financed by the United Nations Development Programme (UNDP) between 1982 and 1984.

The Rectors of the four state universities on Sulawesi namely Fachruddin, (Hasanuddin University, Ujung Pandang), Mattulada (Tadu-lako University, Palu), W.J. Waworoento (Sam Ratulangi University, Manado) and Eddy Agussalim Mokodompit (Haluoleo University, Kendari) and their staffs have cooperated by supplying information and have supported the effort to further the ecological understanding of Sulawesi.

The entire English text has been critically reviewed and examined by Jane Whitten (Bogor), Anoma Santiapillai (Bogor) and Gembong Tjitrosoepomo (Yogyakarta), the third of whom made the translation for the Indonesian version, Ekologi Sulawesi. Major portions were reviewed by Peter Bellwood (Canberra), Chris Bennett (Manado), David Bulbeck (Ujung Pandang), S.C. Chin (Kuala Lumpur), James Davie (Rock-hampton), David Dudgeon (Hong Kong), Ian Glover (London), Atmadja Hardjamulia (Bogor), Duncan Parish (Kuala Lumpur), Nicholas Polunin (Port Moresby), and Nengah Wirawan (Ujung Pandang). Great thanks are due to these people for their constructive criticisms, but it must be stressed that the mistakes remaining in the text are entirely the responsibility of the authors.

The Executive Director of Gadjah Mada University Press, H.J. Koesoe-manto†, has always been willing to advise, listen, and cooperate and we owe him a considerable debt of gratitude.

We also express gratitude to the Royal Entomological Society of London for permission to visit their Project Wallace research site in the Bogani Nani Wartabone (formerly Dumoga-Bone) National Park, Bolaang Mongondow, in 1985.

In addition, many people have provided considerable assistance (often more than they realize) by sending reports, papers, unpublished information and other forms of information, by advising, cajoling, identifying specimens, by making helpful suggestions, or by helping the authors in the field. They are: Abdul Rachman Abudi, Amran Achmad (Ujung Pandang), Mohamad Amir (Bogor), Dick R. Askew (Manchester), Michael Audley-Charles (London), Andy Austin (Adelaide), Max van Balgooy (Leiden), Henry S. Barlow (Kuala Lumpur), Wim Bergmans (Amsterdam), David Bishop (Chesterhill and NSW), Roger Blackith (Dublin), Peter Bloks (Leiden), Boeadi (Bogor), Hans A.J. in den Bosch (Leiden), W. Boudewijn (Ujung Pandang), Martin Brendell (London), Francois Brouquisse (Tolouse), Sean Brown (London), Arie Budiman (Bogor), Elisa Bung'alo (Lumuk), Burhan (Ujung, Pandang), Roger Butlin (Norwich), Diane Cal-abrese (Carlisle, Penn.), Ray Catchpole (Kendari), Ailsa Clark (London), Lynn Clayton (Oxford), Nigel Collar (Cambridge), Mark Collins (Cambridge), David Coyle (Watampone), Wempy Dahong (Ujung Pandang), Rokhmin Dahuri (Bogor), Sengli Damanik (Medan), Anthony Davis (Nagercoil), Louis Deharveng (Tolouse), Rene Dekker (Dumoga), Peter Dinwiddie (Swindon), Henry Disney (Cambridge), Machfudz Djajasasmita (Bogor), John Dransfield (Kew), Julian Dring (London), Jans Duffels (Amsterdam), Lance Durden (Nashville), Rusly Durio (Ujung Pandang), Siegfied Eck (Dresden), Chris Escott (Saskatoon), C.H. Fernando, (Waterloo), Theodore Flemming (Coral Gables, FL), Ben Gaskell (London), S.S. Gasong (Palu), E. Gittenberger (Leiden), Emily Glover (London), Michael Green (Cambridge), Penny Greenslade (Canberra), Steven Greenwood (Oxford), Colin Groves (Canberra), James Guiry (Soroako), Surastopo Hadisumarno (Yogyakarta), Gavin Hainsworth (Vancouver), Tony Harman (Canterbury), Hanna Hardjono (Ujung Pandang), A.M. Hashi (Palopo), Loky Herlambang and the staff of the Nusantara Diving Centre (Manado), John E. Hill (London), Bert Hoeksema (Leiden), Ian Hodkinson (Liverpool), Jeremy Hollowax (London), Derek Holmes (Jakarta), L.B. Holthuis (Leiden), Marinus Hoogmoed (Leiden), Geoff Hope (Canberra), Hans Huijbregts (Leiden), Jaffre (Palopo), Paula Jenkins (London), Clive Jermy (London), Ahdul Rachman Kadir (Ujung Pandang), John Katili (Jakarta), Peter Kevan (Guelph), Ashley Kirk-Spriggs (Cardiff), David Kistner (Chico), Roger Kitching (Armidale), Robby V.T. Ko (Bogor), Jan de Korte (Amsterdam), Maurice Kottelat (Courrendlin), Jan Krikken (Leiden), Jaroslav Klapste (Melbourne), Bill Knight (London), Andrew Lack (Swansea), David de Lauberfels (Syracuse), Philippe Le Clerc (Tolouse), Cecile Lomer (Bogor), C.H.C. Lyal (Auckland), Colin McCarthy (London), Ron and Patty McCullogh (Palu), Odilia Maessen (Halifax, N.S.), Ayub Mahmud (Malili), Syafii Manan (Bogor), Adrian, G. Marshall (Aberdeen). Joe Marshall (Washington), G.A. Matthews (Ascot), John Miksic (Yogakarta), Andrew Mitchell (London), Willem Moka (Ujung Pandang), Robert Molenaar (Manado), Hans Moll (Leiden), Kate Monk (Safat), Barry P. Moore (Canberra), Evelyn Mundy (Bogor), Guy Musser (New York), Safiruddin Natsir (Ujung Pandang), Baharuddin Nurkin (Ujung Pandang), Grace O'Donovan (Dublin), Sharifuddin Andi Omar (Ujung Pandang), H.S. Padeato (Manado), S.W. Padiato (Luwuk), Ray-mondus Palete (Dumoga), J.L. Panelewan (Manado), P.E.A. Pangalila (Manado), Duncan Parish (Kuala Lumpur), Michael Pearce (London), Totok Prawitosari (Ujung Pandang), T. Racheli (Rome), Musaka Rachmat (Ujung Pandang), Anthony Reid (Canberra), Christopher Rees (York), Mien Rifai (Bogor), Willem Rodenburg (Koudekerk a/d Rijn), Frank G. Rozendaal (Bilthoven), Didi Rukmana (Ujung Pandang), A. Sabani (Palopo), Peter Sane (Kendari), Sjahril T. Selamat (Ujung Pandang), Victoria Selmier (San Francisco), Dan Sembel (Manado), Siahaan (Luwuk), Alison Skene (Bogor), the late C.G.G.J. van Steenis (Leiden), Nigel Stork (London), J.S.W.D. Subroto (Manado), N. Sulaiman (Jakarta), Mulyadi Susanto (Kendari), Stephen Sutton (Leeds), Antius Tolesa (Luwuk), Michael Tremble (Albuquerque), John Uttley (Durham), Charlotte Ver-meulen (Dumoga), Ed de Vogel (Leiden), Michael Wade (Bogor), B. Wahyu (Soroako), Sarah Warren (Yale), Dick Watling (Suva), Chris Watts (Adelaide), Alice Wells (Adelaide), Chris Wemmer (Washington), Michael Walters (Tring), Alwyne Wheeler (London), Tim Whitmore (Oxford), Willem de Wilde (Leiden), Johanna Wilson (Bogor), Mike Wilson (London) and John Winter (Townsville).

The principal typist has been Lisbet† who has worked with great diligence, care and commitment. The majority of the figures were drawn by Syamsul, to whom our considerable thanks are due.

Finally, and mainly, we thank Almighty God for His grace and caring love shown throughout the preparation of this book, without which it would never have been completed.

† Deceased

Figure I.1. Main towns, island groups and islands of Sulawesi.

Introduction

We arrived in Sulawesi in 1991, our bags overstuffed with the gear of ecologists—binoculars, notebooks, two dictionaries and one extremely heavy reference book, The Ecology of Sulawesi. The Ecology of Sulawesi provided a summary of the current knowledge of the island at that time. There was no argument about the thoroughness of the book; if a topic wasn't mentioned in The Ecology of Sulawesi, it probably hadn't been studied. At the time, there wasn't even a field guide to the birds, and our first identifications of endemic mynas relied on a small, black and white sketch in chapter one. Because few other references were available, The Ecology of Sulawesi became our bible. Over a four-year period of constant use by visitors, students, and biologists—not the least ourselves—our copy developed a broken binding and dog-eared, underlined pages—the highest compliment to the authors.

Much has changed in the past 13 years since the first edition of The Ecology of Sulawesi was published. There have been discoveries that we never imagined, and losses beyond what only the worst of pessimists would have dreamed in 1987. Unfortunately, funding is not yet available for a full revision that would incorporate all these changes into a new edition of The Ecology of Sulawesi. This second edition of The Ecology of Sulazvesi, however, will still be invaluable in any library. The updated bibliography should provide readers with numerous sources for more recent information about the island. Our first edition copy (now rebound) never stays on the shelf long and remains a primary reference when we, our staff and students are writing manuscripts.

Surely the most dramatic discovery of the last 13 years was the completely unexpected find of a coelocanth fish in a Manado market (Erd-mann et al. 1998). Indeed, it was heralded by some as 'the zoological find of the century'. Only one other population of this 'living fossil' (Forey 1998), located 1,000 kms off the coast of east Africa, has ever been discovered. Unfortunately, the excitement of discovery was marred by politics of taxonomy and currently controversy rages over whether or not the Manado population is a separate species. But the arguments certainly do not take away from the sheer excitement of such a discovery.

Perhaps the most satisfying find, especially for the authors of The Ecology of Sulawesi, was the rediscovery of the endemic Cerulean paradise flycatcher on Sulawesi's satellite island of Sangihe (Wardill and Riley 1999). The caption of plate 4 of the first edition states that the Cerulean paradise flycatcher bird is probably extinct. The flycatcher became become a poster bird for extinction when plate 4 appeared on the cover of the first issue of the journal for Conservation Biology and the species was declared 'almost certainly' extinct (Whitten et al. 1987). First described in 1873, the bird was known only from one museum specimen and was last sighted in 1981. In 1998 however, members of a small non-governmental organization (NGO), Action Sampiri, spotted the flycatcher while conducting ornithological surveys of the island. Although uncommon and restricted to an isolated mountaintop, the bird survives and provides a rare spark of optimism.

Figure I.2. Division of Sulawesi into provinces and counties.

The Sangihe-Talaud Islands have proven fertile grounds for other ornithological discoveries. Frank Lambert and colleagues uncovered three species new to science, including two rails and an owl (Lambert 1998a, b), verified the continued existence of the Sangihe shrike-thrush (Rozendaal and Lambert 1999), and uncovered two distinct scops owl species from Sangihe and Siau (Lambert and Rasmussen 1998). Similarly, surveys for mammals on Sangihe-Talaud have produced the first records this century of an endemic rat, a distinct form of bear cuscus, and a the Talaud fly ing fox—the latter found in a small restaurant, presumably headed for the pot.

Back on the mainland, biologists and other modern-day explorers have been making similar discoveries, including taxa as diverse as fungi (Rogers et al. 1987), freshwater crabs (Ng 1993), turtles (McCord et al. 1995; Platt et al. in press), rodents (Musser 1991; Musser and Holden 1991), snakes and lizards (Iskandar 1999a; Lazell 1987; Bosch and Ineich 1994), insects (Conde 1992a, b, 1994; van Tol 1987, 1994; Monk and Butlin 1990) and plants (Dransfield 1989, 1992) as well as co-evolved relationships between ants and trees (Maschwitz and Fiala 1995). Parks and protected areas have revealed a number of secrets, underscoring their importance for species protection. In the late 1980s and early 1990s, expeditions to Lore Lindu National Park revealed two new tarsier species (Musser and Dagosto 1987; Nemitz et al. 1991). More recently, a Ninox owl, previously known only from a single museum specimen (Rasmussen 1999), and the Matinan flycatcher were recorded in Gunung Ambang Nature Reserve (Lee and Riley in press). Most recently, a Heinrich's nightjar was observed in Panua Nature Reserve and unexpectedly large populations of anoa and babirusa have been found in Bogani Nani Wartabone (formerly known as Dumoga-Bone) National Park (Lee pers. comm.). The Sulawesi palm civet, thought to be tumbling towards the vortex of extinction, was photographed by a team from The Nature Conservancy in the forests of Lori Lindu National Park (Indonesian Observer 2000).

In addition to species discoveries and rediscoveries, the 1990s experienced an explosion of medium- and long-term research projects. These projects focused primarily on large vertebrates, especially Sulawesi endemics. Sulawesi's seven species of endemic macaques received the lion's share of attention with research covering taxonomy (Bynum 1999; Evans et al. 1999; Watanabe et al. 1991), conservation status (Bynum 1999; O'Brien and Kinnaird 1996), ecology (Lee 1997; Matsumura 1993, 1996, 1998; O'Brien and Kinnaird 1997; Rosenbaum et al. 1998), and the dramatic scale of hunting (Alvard 2000; Lee 2000a, b; O'Brien and Kinnaird 1996, 2000). The previously unknown natural history of the rare babirusa was unfolded by Clayton (1996) and Clayton and MacDonald (1999) during her intensive studies at a salt lick in Paguyaman forest, North Sulawesi. First-ever studies were published on the ecology of bear cuscus (Dwiyaherni et al. 1999), endemic forest kingfishers (Sunarto 1999), and hornbills (O'Brien 1996; Kinnaird and O'Brien 1993, 1999a; Kinnaird et al. 1996; Suryadi et al. 1994, 1996). Dekker and colleagues (Argeloo 1994; Dekker 1990; Dekker et al. in press) provided the first detailed information on the status and distribution of Sulawesi's oddest avian endemic— the Maleo. Gursky (1994, 1998) has greatly improved our knowledge of tar-sier behavior and ecology, Alvard and Winarni (1999) assessed bird communities in relation to habitat disturbance in Morowali Nature Reserve, and Bynum (1999) developed new habitat monitoring techniques for use in Lore Lindu National Park.

Sadly, all this ground-breaking research and discovery is occurring against a background of destruction. Since the publication of the first edition of The Ecology of Sulawesi, deforestation has continued unchecked. In spite of the efforts of international donor agencies (e.g., United States Agency for International Development [USAID], the World Bank, United Nations Development Program [UNDP], Asian Development Bank [ADB]) and the dedication of many NGOs, habitat loss on Sulawesi has actually accelerated. In a shocking report by Derek Holmes (2000) for the World Bank and the Ministry of Forestry and Estate Crops, Holmes shows that between 1985 and 1997 Sulawesi lost 20% of its natural forest cover. This figure does not account for forest quality, and land classified as forest could contain logged or burned forest, areas of reduced value for biodiversity conservation. Most startling, Holmes states that lowland dry forest, the most valuable type of Indonesian forest for logging and biodiversity conservation is 'essentially defunct as a viable resource in Sulawesi'. This should have been a wake-up call to the Indonesian government but instead the list of causes of forest destruction grew longer and more complex.

The leading cause of deforestation has been large-scale logging by a few conglomerates and the use of timber concessions for political patronage— huge tracts of state-owned forests were held in the 1980s and 1990s by family and business associates of former president Suharto (Richardson 2000). Ironically, central government policy has encouraged deforestation. Logging concessions, sugarcane plantations, and increasingly, oil palm plantations are not regulated and as Walton and Holmes (2000) state, 'perverse incentives exist that make it more lucrative to clear forested land for plantations than to plant open and unproductive land'. As timber exports shifted from logs to plywood and then to pulp and paper, more log processing plants were established until processing capacity exceeded sustainable yields. Finally, poor enforcement has allowed excesses to go unchecked. Illegal logging has become rampant, even in national parks, on a scale that exceeds the volume of legal logging. Authorities look the other way while the government loses tax revenue at the rate of roughly $500 million each year (Walton and Holmes 2000).

An increasingly important agent of deforestation on Sulawesi is the influx of new migrants into forested areas. In the late 1990s, as Indonesia plunged into economic crisis, unemployed urbanites returned to the countryside in need of land. Long-term residents also cleared new land to expand production of low-value vegetable crops and high-value cash crops (e.g., coffee, cocoa). As we write, new immigrants are arriving from areas of ethnic strife in the Moluccas. Thousands of displaced and land-hungry refugees are arriving daily; many observers report that there are now more than 200,000 refugees in North Sulawesi alone. Unless the problem is resolved soon, these people will resettle and require more land to support themselves and their families.

Forests that are not being cut are not necessarily safe. One of the most alarming discoveries of the 1990s is the degree to which people of Sulawesi are destroying their wildlife heritage (Lee 1997; O'Brien and Kinnaird 1996, 2000). Researchers showed that a number of endemic mammals, including the babirusa, anoa, and crested black macaque were on the edge of extinction as a result of over-hunting for subsistence and commercial bushmeat trade. Further evidence came from modeling exercises by Clayton and colleagues (1997) who showed that under current practices, babirusa populations will be reduced to very small and probably nonviable population numbers. Lee (1997, 2000a, b, c) conducted an extensive analysis of commercial and subsistence hunting and concluded that hunting for markets was having the greatest impact on local populations of wildlife, and Alvard (2000) showed a lesser, but still considerable, impact on wildlife through subsistence hunting by the Wana people of Morowali Nature Reserve. Overcollecting of sulphur-crested cockatoos for the pet trade has almost eliminated the species from Sulawesi except for a few remnant birds near Palu (PHPA/LIPI/BirdLife International-IP 1998). Finally, the breakdown of traditional customs and harvest regimes has played a major role in the on-going decline of the Maleo. Although more breeding sites have been discovered since the classic work of Dekker (1990), most are in danger from over-collecting and many sites are no longer active.

Sulawesi has a diverse system of parks and nature reserves that were set up to provide sanctuary for the island's biodiversity, free from hunting, and habitat destruction. The island boasts three national parks, including Lore Lindu National Park, a World Heritage Site, two marine parks, nineteen nature reserves, and an assortment of tourist parks and wildlife refuges. These areas cover most major habitats, contain populations of most endemic species, and several are large—all good qualities for reserves design. But in today's climate, the subject of reserve design is largely one of academic debate. Sulawesi's parks are not safe and although most of the protected areas have staff, the areas function as little more than paper parks. The question of what percentage of species will become extinct in 50, 500 or 5,000 years' time is not of great relevance when it is by no means certain how much of its reserves will be intact in even 25 years time. In Sulawesi today the major priority is simply to maintain the integrity of reserves against legal and illegal forms of habitat disturbance and resource exploitation.

Hunting and habitat destruction continue unchecked throughout Sulawesi's protected area system. Development projects such as road building, new transmigration schemes and dam development are being permitted in the parks, and in most cases supported by local government officials. Mining, both illegal and legal, continue to plague areas such as Bogani Nani Wartabone National Park. As we write, more than 4,000 illegal miners scour the hillsides of the park in search of gold, poisoning themselves and the environment with the mercury they use to separate the precious metal (Sydney Morning Herald 2000). An additional 5,000 illegal miners—supported by the military, local police and government officials— have invaded the Perth-based Aurora Gold's claim on the outskirts of Manado, similarly poisoning themselves and the surrounding waters of Bunaken Marine National Park (Sydney Morning Herald 2000). Logging operations encroach on reserve boundaries in many areas. Fires set by plantation companies and local people ravaged reserve land in 1994-95 and again in 1997-98. Illegal fishing nets, set in protected waters of Tangkoko drowned thousands of marine mammals and sea turtles, and killed manta rays and other pelagic fish. In spite of legal protection, hunters, rattan collectors and other forest product collectors roam freely throughout many reserves in North and Central Sulawesi, undaunted by apathetic, or in some cases, participating enforcement officials.

All of the abuses described have been exacerbated by the prolonged economic crisis, the collapse of central authority and the proposal to decentralize natural resource management. In the confusion and uncertainty, local natural resource managers are more reluctant than ever to take action. As local people move into protected areas and even logging concessions, they find that there is little resistance by the government. In addition, military and land speculators are getting into the act on the belief that if the land is cleared, they will be able to keep it. In Lore Lindu National Park, park management turned over approximately 2,000 ha of park land to local people for illegal coffee and cacao plantations, rather than confront the issue (Indonesian Observer 2000). As management breaks down, guards feel less responsible, are demoralized and stop working. In Tangkoko Nature Reserve in 1997, guards watched, but failed to report the forest fires that damaged more than half of the reserve.

Although the problems of Sulawesi's forests, parks and protected areas are severe, they are not insurmountable. Today, a growing number of organizations and individuals are committed to the conservation and rational management of Sulawesi's natural resources. The Natural Resource Management and Coastal Resource Management Programs of USAID have been working with local stakeholders including the government, local NGOs, and communities to develop policy and implement plans for sustainable use of marine and terrestrial resources. They were responsible for developing the management plan and basic infrastructure for Bunaken Marine National Park, one of Indonesia's most exquisite and popular marine sanctuaries. Conservation International and Biological Conservation Network have been working on the Togian Islands developing community-based management of coral reef tourism and protection of local forests. Sahabat Morowali, an Indonesian NGO, focuses on the conservation issues of Morowali National Park. They are especially concerned with involving the resident Wana people in management decisions concerning the park and have developed ecotourism ventures. The Nature Conservancy concentrates its efforts in Lore Lindu National Park. They have provided resources and training to the Directorate of Nature Protection and Conservation (PKA) staff, and are trying to develop alternative, environmentally friendly income sources aimed at reducing pressure on the park. Group Sampiri, a local NGO based on the work of Action Sampiri, is carrying out a rural awareness project on Sangihe-Talaud. Our organization, the Wildlife Conservation Society, has been active in Sulawesi since 1991 carrying out wildlife research and training of conservationists in protected areas such as Tangkoko, Gunung Ambang, and Bogani Nani Wartabone in North Sulawesi. The Wildlife Conservation Society conducts in-depth ecological research and baseline surveys on a variety of endangered species, conducts conservation education campaigns in North Sulawesi and trains park guards and provincial officials in conservation enforcement. At present, we are carrying out an island-wide field survey of protected areas and wildlife, and combining these surveys with training and management assistance.

In the final analysis, all the hard work of foreign and domestic conservation organizations will pale unless Indonesia as a nation—including its government, military, police forces, local NGOs, universities and communities—begin to show a serious commitment to conservation on Sulawesi. Unless the talk stops and the action begins, we will lose everything that made Sulawesi a special place in Alfred Russell Wallace's heart, as well as our own. And the next edition of The Ecology of Sulawesi could very easily become a history book.

M.F. Kinnaird and T.G. O'Brien

September, 2000

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