Ecology of Kalimantan: Indonesian Borneo

THE ECOLOGY OF INDONESIA SERIES

Volume III: The Ecology of Kalimantan

Other titles in the Series

Volume I: The Ecology of Sumatra

Volume II: The Ecology of Java and Bali

Volume IV: The Ecology of Sulawesi

Volume V: The Ecology of Maluku and Nusa Tenggara

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

Copyright © 1996 Dalhousie University

All maps in this edition copyright © 1996 Periplus Editions (HK) Ltd

All rights reserved

Published by Periplus Editions (HK) Ltd.

ISBN: 978-1 4629-0505-8 (ebook)

Publisher: Eric Oey

Design: Pete Ivey

Prepress production: JWD Communications Ltd.

Copy editing: Sean Johannesen and Kathy McVittie

Distributors:

Indonesia:

C.V. Java Books, P.O. Box 55 JKCP

Jakarta 10510

Singapore and Malaysia:

Berkeley Books Pte. Ltd.

61 Tai Seng Avenue, #02-12

Singapore 534167

Tel: (65) 6280 1330

Fax: (65) 6280 6290

Email: inquiries@periplus.com.sg

Web site: www.periplus.com

The Netherlands:

Nilsson & Lamm B.V.

Postbus 195, 1380 AD Weesp,

Printed in the Republic of Singapore

Table of Contents

EMDI

Foreword

Acknowledgements

Abbreviations and Acronyms

Introduction

Chapter 1 The Island of Borneo

Geography

Geological History

Geology

Soils

Climate

Flora

Fauna

Biogeography

Biogeographical Units in Borneo

Human History in Borneo

Current Land Use

Chapter 2 Coastal Habitats

Offshore Islands and Rocky Islets

Rocky Shores

Beaches and Beach Forests

Estuaries and Mudflats

Seagrass Beds

Coral Reefs

Mangrove Forest

Mangrove Zonation

Mangrove Ecology

Mangrove Fauna

Mangrove Habitats as a Valuable Resource

Chapter 3 Freshwater Habitats

Swamp Forests

Peat Swamp Forest

Formation of Ombrogenous Peat Swamps

Ecology of Peat Swamp Forest

The Fauna of Peat Swamp Forests

Freshwater Swamp Forest

Fauna of Freshwater Swamp Forests

Rivers

Physical Patterns in Rivers

Water Chemistry

River Communities

Energy Flow in Rivers

Invertebrate Drift

Freshwater Fishes and Vertebrate Predators

Riverbank Communities

Kalimantan Lakes

Weedbeds and Swamps

The Kapuas Lakes - a Wetlands Ecosystem

Human-Made Lakes

HumanImpacts On Freshwater Ecosystems

Chapter 4 The Lowland Rainforest of Borneo

Lowland Dipterocarp Forest

Forest Structure

Stratification

Tree Form

Ground Layer

Climbers, Stranglers and Epiphytes

Forest Succession

Biomass And Productivity

Nutrient Cycling

Flowering, Fruiting And Leaf Production

Plant-Animal Interactions

Herbivory

Pollination

Seed Dispersal

Animal Communities

Stratification of Animal Communities

Competition and Niche Separation

Soil and Litter Communities

Forest Floor Community

Larger Herbivores

The Forest at Night

Chapter 5 Other Lowland Forest Formations

Heath Forest Or Kerangas

Forest Structure and Composition

Heath Forest Regeneration

Fauna

Ironwood Forests

Forest Over Ultrabasic Rocks

Gunung Silam: an Ultrabasic Mountain

Pleihari-Martapura Wildlife Reserve

Land Use on Ultrabasic Soils

Chapter 6 Limestone Habitats

Soils And Drainage

Limestone Flora

Limestone Forests

Lowland Scree Forest

Lowland Limestone Cliff Communities

Lowland Limestone Forest

Lower Montane Limestone Forest

Upper Montane Forest

The Fauna Of Limestone Hills

Caves

Cave Communities

Bats

Swiftlets

Roof Community

Floor Community

The Mulu Caves

Caves And Human History

Caves As A Resource

Conservation Of Caves And Limestone Formations

Chapter 7 Mountain Habitats

Climate

Mountain Soils

Productivity And Nutrient Cycling

Zonation Of Mountain Forests

Zonation Of Animals On Tropical Mountains

Mount Kinabalu - Summit Of Borneo

The Fauna of Mount Kinabalu

Bukit Raya

Effects Of Disturbance On Montane Habitats

Mountains As Centres For Biological Diversity

Chapter 8 Borneo Peoples - Migrations and Land Use

Penan: Harvesters Of The Forest

Dayak Groups

Dayak Migrations

Coastal Communities

Shifting Agriculture: Subsistence Farming

Shifting Agriculture: Ecologically Sound or Environmentally Damaging?

Harvesting The Forest

Omens And Augury

Transmigration And Resettlement

The Future

Chapter 9 Forest Resources

Forests For Timber

Effects of Logging on Forest Structure and Dynamics

The Sustainability of Timber Harvests

Other Forest Products

Rattans

Resins and Incense Wood

Illipe Nuts

Forests For Water And Soil Conservation

Forest Disturbance And Wildlife

Reforestation

Forest Fires On Borneo

Fire and Forest Ecosystems

Conservation Of Forests

Chapter 10 Wetland Resources

The Ecological Importance Of Wetland Habitats

Water Flow And Water Quality

Plant Harvests From Wetlands

Timber from Swamp Forests

Food Chains And Animal Resources

Fisheries In The Middle Mahakam Lakes

Aquaculture

Reclaiming Tidal Swamplands For Agriculture

Acid Sulphate Soils

Success and Failure in the Tidal Swamps

Peat Swamps for Agriculture

Peat For Fuel

Creation Of New Wetland Habitats

Rice Fields

Introducing Exotic Species

The Sungai Negara Swamps -A Converted Wetland Habitat

Wetlands For Conservation

Chapter 11 Coastal Resources

Estuarine And Coastal Fisheries

Harvests from the Sea

The Value Of Coral Reefs

Mangroves As A Resource

Mangrove Fisheries

Mangroves for Commercial Timber and Chipwood

Other Mangrove Products

Mangrove Services

Mangrove Management

Conversion to Tambak

Oil Spills And Coastal Ecosystems

Development And Coastal Resources

Chapter 12 Agriculture and Plantations

Agricultural Potential

Agroecosystems

Rice

Cassava

Fruit Trees

Pepper

Other Crops of Ladangs and Home Gardens

Plantations

Coconuts

Rubber

Oil palm

Cocoa

Timber as a Plantation Crop

Plantation Ecology

Pest Ecology and Control

Livestock

Genetic Resources

Future Trends In Agriculture In Kalimantan

Chapter 13 Development and the Environment

Mineral Resources

Gold

Coal

Limestone

Diamonds

Mining And The Environment

Petroleum

Oil Pollution

Industrial Developments

Industrial Effluents and Pollution

Forest Conversion

Forest Loss And Climatic Change

The Greenhouse Effect

Rehabilitating Critical Lands

Agroforestry On Critical Soils

The Urban Environment

Urban Ditches and Ponds

Houses

Indonesia's Expanding Population

Ecology Of Diseases

The Future

Chapter 14 Conservation: Protecting Natural Resources for the Future

Biodiversity

The Biological Importance Of Borneo

Species Extinctions

Endangered Species in Borneo

The Need To Conserve Biological Resources

Habitat Protection

Stabilisation of Hydrological Functions

Soil Protection

Stabilisation of Climate

Conservation of Renewable Harvestable Resources

Protection of Genetic Resources

Preservation of Breeding Stocks, Population Reservoirs and Biological Diversity

Maintenance of the Natural Balance of the Environment

Economic Benefits from Tourism

Environmental Monitors

Future Benefits

Genetic Resources

The Protected Areas System Within Borneo

Species Protection in Reserves

Will the Reserve System Alone Be Adequate to Protect Borneo's Biodiversity?

Conservation Outside Protected Areas

Buffer Zones

The Value of Secondary Forests

Production Forests

Captive Breeding and Reintroductions

Research Needs

The Future

Appendices

Appendix 1. Plant genera and families, mentioned in text.

Appendix 2. Land birds found on offshore islands of Borneo.

Appendix 3. Distribution of Bornean montane birds.

Appendix 4. Distribution of Bornean snakes.

Appendix 5. Simple methods for inventory and monitoring of species.

Appendix 6. Useful forest plants used by Iban people.

Appendix 7. Field-sketch landscape profile, based on Field Rapid Rural Appraisal, for reforestation planning in South Kalimantan.

Appendix 8. Species Databases

Bibliography

Bibliography Addendum

Index

List of Boxes

Box 2.1. The dugong: mermaid or endangered marine mammal.

Box 2.2. Simplified classification of the major invertebrates occurring in the benthos of coral reefs.

Box 2.3. Feeding guilds of coral reef fishes.

Box 2.4. Mangrove species in Kalimantan.

Box 2.5. Mangrove forests in the Barito River estuary.

Box 3.1. Riverine habitats in Borneo: a classification of flowing waters.

Box 3.2. Ecomorphology of fish.

Box 3.3. Feeding guilds of fish.

Box 3.4. Rheophytes - riverine plant specialists.

Box 3.5. Watchful mothers - parental care in crocodiles.

Box 3.6. The freshwater fish of the Kapuas River system.

Box 4.1. Forest regeneration.

Box 4.2. Mass flowering and mast fruiting in dipterocarps.

Box 4.3. Dioecism and cross-pollination among scattered rainforest trees.

Box 4.4. Species richness in lowland rainforest.

Box 4.5. Fig eating and seed dispersal by rainforest birds.

Box 4.6. Niche separation in tropical squirrels.

Box 4.7. Niche separation in Sundaland primates.

Box 4.8. Gliding ? a rainforest adaptation.

Box 5.1. Mandor Nature Reserve.

Box 5.2. Myrmecotrophy: plants fed by ants.

Box 5.3. Plants as predators: pitcher plants.

Box 5.4. Agathis , a tropical conifer.

Box 5.5. White sands and blackwater rivers.

Box 5.6. Monospecific dominance in tropical rainforests.

Box 6.1. The Sangkulirang limestone formations.

Box 7.1. Rhododendrons.

Box 7.2. Orchids.

Box 7.3. The trig-oak, a missing link.

Box 8.1. Shifting agriculture.

Box 8.2. People and forests in East Kalimantan.

Box 8.3. Forest management by local villagers in West Kalimantan.

Box 8.4. Wild game meat in Sarawak.

Box 8.5. Traditional medicines of the Kenyah of East Kalimantan.

Box 8.6. Uses of forest plants in Sarawak.

Box 8.7. The cows of Long Segar: a lesson in development aid.

Box 9.1. Commercial forest management systems.

Box 9.2. The economic values of some forest species.

Box 9.3. Gaharu (aloe wood or incense wood).

Box 9.4. Valuation of a rainforest.

Box 9.5. The effects of logging.

Box 9.6. Forest functions.

Box 9.7. Logging and wildlife in the Sungai Tekam Forestry Concession, Pahang, West Malaysia.

Box 9.8. Watershed management: Riam Kanan, South Kalimantan.

Box 9.9. Some possible effects of burning on five ecological processes.

Box 9.10. Goods and services provided by tropical forests.

Box 10.1. Making aquatic weeds useful.

Box 10.2. Sago orchards of the Melanau.

Box 10.3. Melaleuca - the paper bark tree (gelam).

Box 10.4. Ikan siluk , the dragonfish of West Kalimantan.

Box 10.5. Wetland resources of economic value.

Box 10.6. Alabio ducks.

Box 10.7. Acid sulphate soils.

Box 10.8. Tidal swamplands and the Banjarese system of agriculture.

Box 10.9. Rice harvesting at Gambut and Kertak Hanyar, South Kalimantan.

Box 10.10. Threats to wetlands.

Box 11.1. Management implications of harvest, hunting and mining activities on estuaries.s

Box 11.2. Harvesting turtle eggs in the Berau turtle islands, East Kalimantan.

Box 11.3. Connections between coral reefs and neighbouring linked habitats.

Box 11.4. Products of mangrove ecosystems.

Box 11.5. Commercial and traditional uses of some Bornean mangrove species.

Box 11.6. Pollutants and their effects on coastal ecosystems.

Box 11.7. Coastal ecosystems in Kalimantan.

Box 12.1. Good agricultural practices.

Box 12.2. Irrigated rice fields in the Kerayan.

Box 12.3. Buginese pepper farmers.

Box 12.4. Cultivation of rattans.

Box 12.5. Some effects of grazing by domestic stock on five ecological processes.

Box. 13.1. Amdal: A guide to environmental assessment in Indonesia.

Box 13.2. Protection and wise utilisation of valuable ecosystems during development.

Box 13.3. The ozone layer - a global issue.

Box 13.4. Riam Kiwa - agriculture in a degraded valley.

Box 13.5. Simplified comparison of some system properties between a natural ecosystem and a man-made structure.

Box 13.6. Habitat manipulations that may be ecologically beneficial.

Box 13.7. Two visions for the future of Indonesia.

Box 14.1. Within-and between-habitat diversity. High diversity in the tropics could result from (a) more species occurring in each habitat; (b) more habitats, each containing the same number of species; or (c) a combination of both.

Box 14.2. Characteristics of species affecting their survival.

Box 14.3. Nonconsumptive benefits of conserving biological resources.

Box 14.4. Simplified scheme for assessing suitable protection category for protected habitats.

Box 14.5. Different categories of protected areas within Kalimantan.

Box 14.6. Linking conservation with development.

Box 14.7. Habitat fragmentation and loss of species: an Indonesian example.

Box 14.8. Benefits of buffer zones.

Box 14.9. Selection of crops for buffer zones.

Box 14.10. Habitat conservation within a large timber concession: Danum Valley, Sabah.

EMDI

The Environmental Management Development Project (EMDI) was designed to upgrade environmental management capabilities through institutional strengthening and human resource development. A joint project of the Ministry of State for Environment (LH), Jakarta, and the School for Resource and Environmental Studies, Dalhousie University, Halifax, Nova Scotia, EMDI supported LH's mandate to provide guidance and leadership to Indonesian agencies and organizations responsible for implementing environmental management and sustainable development. Linkages between Indonesian and Canadian organizations and individuals in the area of environmental management are also fostered.

EMDI received generous funding from the Canadian International Development Agency (CIDA). CIDA provided Cdn$2.5 million to EMDI-1 (1983-86), Cdn$7.7 million to EMDI-2 (1986-89), and contributed Cdn$37.3 million to EMDI-3 (1989-95). Significant contributions, direct and in kind, were made by LH and Dalhousie University.

EMDI-3 emphasized spatial planning and regional environmental management, environmental impact assessment, environmental standards, hazardous and toxic substance management, marine and coastal environmental management, environmental information systems, and environmental law. The opportunity for further studies was offered through fellowships and internships for qualified individuals. The books in the Ecology of Indonesia series form a major part of the publications programme. Linkages with NGOs and the private sector were encouraged.

EMDI supported the University Consortium on the Environment comprising Gadjah Mada University, University of Indonesia, Bandung Institute of Technology, the University of Waterloo, and York University. Included in EMDI activities at Dalhousie University were research fellowships and exchanges for senior professionals in Indonesia and Canada, and assistance for Dalhousie graduate students undertaking thesis research in Indonesia.

For further information about the EMDI project, please contact:

Director

School for Resource and Environmental Studies

Dalhousie University

1312 Robie Street

Halifax, Nova Scotia

Canada B3H 3E2

Tel. 1-902-494-3632

Fax. 1-902-494-3728

Foreword

Indonesia covers only 1.3% of the earth's surface, yet it harbours some of the world's richest biodiversity: 10% of all flowering plants, 12% of the world's mammal species, 16% of all reptiles and amphibian species, 17% of the world's birds and a quarter of all marine and freshwater fishes. This incredible biological richness is not only an important part of our natural heritage, it provides the life support systems, natural harvests and environmental services on which the nation's health, livelihood and well-being depend. An estimated 40 million people (almost one-quarter) of Indonesia's population are dependent on biodiversity for subsistence through harvesting of coastal, freshwater and marine fisheries, collection of non-timber forest products or cultivation of indigenous fruits, vegetables, cereals and spices. Conservation of the country's natural ecosystems and the biodiversity and environmental services they provide is crucial to the sustainability of economic development in sectors as diverse as forestry, agriculture and fisheries; health care; science; industry and tourism.

Protection and wise utilisation of biological resources requires that we understand how ecosystems work and the interactions and interdependence between natural habitats and a healthy environment. The Ecology of Indonesia series, part of the CIDA-funded Environmental Management Development in Indonesia Project attempts to explain the principles of ecology in. the Indonesian context. This third volume in the series focuses on Kalimantan, part of Borneo, the world's fourth-largest island. Studies in Kalimantan provide valuable insights into ecological processes and the complex interactions between plants and animals in natural ecosystems. The book also considers human dependence on this island's biological resources and how human actions are changing and threatening the natural habitats on which so many Kalimantan peoples depend. As Kalimantan's vast tropical rainforests and coastal wetlands come under increasing pressure, this book reminds us that sustainable development depends on the maintenance of ecological processes and environmental functions.

The Ecology of Kalimantan was prepared as a collaborative effort between western and Indonesian scientists based at the environmental study centres in Kalimantan. The project provided training opportunities and resources for Kalimantan scientists and students to undertake ecological research in wetland and forest habitats and also established important literature collections and databases at KPSL-UNLAM, Banjarbaru, and PSL-UNTAN, Pontianak. I congratulate the authors and contributors on publication of this book and welcome this useful contribution to environmental literature of Indonesia.

Jakarta, September 1994

Sarwono Kusumaatmadja

State Minister of Environment

Republic of Indonesia

Acknowledgements

The Ecology of Kalimantan is the third volume in The Ecology of Indonesia series prepared as part of the Environmental Management Development in Indonesia (EMDI) Project. EMDI is a joint project implemented by the Indonesian Ministry of State for Environment (LH) and the School for Resources and Environmental Studies (SRES) at Dalhousie University, Halifax, Nova Scotia, Canada. The project is funded by the Canadian International Development Agency (CIDA).

This book was prepared originally at the environmental study centre, KPSL-UNLAM, at Banjarbaru, South Kalimantan between 1988 and 1990. After delays in publication Kathy MacKinnon undertook to revise and update the text prior to this publication. The authors are grateful to the Rector of UNLAM, Dr. Supardi, and to Ir Hazairin Noor and other staff at the KPSL for their assistance and support with fieldwork and research. Kathy and Richard Eaton, then CUSO volunteers at the KPSL, helped with research, fieldwork and reference collection, and established and catalogued the KPSL library. Kevin Teather and the staffs of PSL-UNTAN, Pontianak, PSL-UNPAR, Palangkaraya and PSL-UNMUL, Samarinda undertook fieldwork and data collection for the book. Wim Giesen and Olivier Klepper of the Asian Wetland Bureau collaborated with the KPSL staff to survey the Sungai Negara wetlands.

The authors also wish to acknowledge the support of Dr. Koesnadi Hardjosoemantri (former Rector of Gadjah Mada University, Yogyakarta) Arthur Hanson, Shirley Conover, George Greene, Gerry Glazier, John Patterson, Barbara Patton and Diane Blachford (SRES, Halifax and EMDI, Jakarta). Special thanks are due to Ani Kartikasari, who helped with all stages of the book from compiling the original references to final revisions. Ani took full responsibility for the production of the Indonesian version, which was translated by Professor Gembong Tjitrosoepomo (Yogyakarta) Agus Widyantoro and herself. The whole book was reviewed by Kuswata Kartawinata, Mien Rifai, John MacKinnon and Tony Whitten. S.C. Chin, George Greene, Lesley Potter, Carla Konsten, Bob Maher, Kevin Teather and Andrew Vayda all read and commented on one or more chapters. Great thanks are due to these people for their constructive criticisms; any faults that remain are the responsibility of the authors.

A volume like this is necessarily a compilation of much information drawn from other sources. The authors are grateful to the many scientists and researchers who generously shared their research data and provided copies of publications. We would also like to thank all those who provided us with hospitality, reference materials, encouragement and assistance with field visits. Special thanks are due to Abdurraman, Pak Aeon of Camera Foto, Lamri Ali and the staff of Sabah Parks, Wim Andriesse, Patrick Andau, Ir Asfihani, Max van Balgooy, Sonny Barnas, Chip Barber, Liz Bennett, Dr. Boeadi, Jean-Marie Bompard, Henk van Bremen, Nick Brown, Arie Budiman, Jules Caldecott, Ir Chaeruddin, Leo Chai, Paul Chai, Cecilia and Danny Chew, Lucas Chin, Carol Colfer, Mark Collins, Alain Compost, Rokhmin Dahuri, Rene Dekker, Michael Dove, Djasmani Hisbi, John Dransfield, Julian Dring, Ros and Ian Edwards, Mochtar Effendi, Clara van Eyck-Bas, Birute Galdikas, Wim Giesen, Ron Greenberg of USAID, Colin Groves, Jerry Harrison, Heruyono, Paul and Coby Hillegers, Jeremy Holloway, Derek Holmes, L.B. Holthuis, Indra, Robert Inger, Alan Irving and staff from Kaltim Primacoal, Clive Jermy, Tim Jessup, Andy Johns, Kuswata Kartawinata, Mike Kavanagh, Victor King, Olivier Klepper, KOMPAS Borneo, Carla Konsten, Jan de Korte, the late Doc Kostermans, Maurice Kottelat, Drs. Koesoemanto of GAMA Press, Jan Krikken, Tony and Anthea Lamb, Danna Leaman, Mark Leighton, Cornelius Lintu, Richard Luxmoore, John MacKinnon, Clive Marsh, Ian McKelvie, Jeff McNeely, Willem Meijer, John Mitani, Margaret Mockler, Peter Neame, Philip Ngau, Hazairin Noor, Riwai Noor, Christine Padoch, Jim Paine, Junaidi Payne, Nancy Peluso, Ron Petocz, Roger and Miranda Pratt, Herman Prayitno, John Proctor, Karen Phillipps, Gusti Rahmat, R. Rajanathan, Ir. Rifani, Ans and Herman Rijksen, Widodo S. Ramono, Anoma and Charles Santiapillai, Toga Siallagan, Marcel Silvius and other staff of AWB, Karta Sirang, Willy Smits, Dr. Soetikno, Belinda Stewart-Cox, Nigel Stork, Anwar Sullivan, Effently Sumardja and staff from PHPA both in Bogor and Kalimantan, Sunardi, Stephen Sutton, Akira Suzuki, Syamsuni Arman, Tan Fui Lian, Andrew Vayda, Jan Vermeulen, Noel Vietmeyer, Ed de Vogel, David Wall, Dick Watling, Tim Whitmore, Tony and Jane Whitten, Kumpiadi Widen, Jan Wind, Richard and Cory Wink, Nengah Wirawan, Eric Wong and the staff of Mount Kinabalu National Park, WWF Bogor, WWF Malaysia, Yayasan Sabah and the staff of Danum Valley research centre. The authors also wish to record their thanks to the staff and librarians of Bogor Herbarium, Bogor Zoological Museum, Brunei Museum, the British Museum (Natural History), Kew Gardens, Leiden Herbarium and Zoological Museum, Sarawak Museum, Sabah Museum and Sabah Forestry Research Centre and the University Malaysia campus in Kota Kinabalu.

Katarina Panji and Rosalind Edwards typed and revised the text with care and much patience. Ismed Inono and Zain Noktah in Kalimantan and David MacKinnon in England prepared the artwork. James and Andrew MacKinnon assisted with the bibliography and proof reading. Kathy McVittie copy edited the English text and prepared the index. Ani Kartikasari and Anoma Santiapillai patiently checked the Indonesian translation. This has been a truly collaborative effort; the authors are grateful for the friendship, support, collaboration, encouragement and sheer hard work of the many people who made it possible.

Abbreviations and Acronyms

Introduction

Kalimantan is the name given to the Indonesian portion of the great island of Borneo, the third largest island in the world after Greenland and New Guinea. Kalimantan covers 73% of Borneo's land mass. The four provinces, Kalimantan Barat (West Kalimantan), Kalimantan Tengah (Central Kalimantan), Kalimantan Selatan (South Kalimantan) and Kalimantan Timur (East Kalimantan) together have a total area of 539, 460 km². This is 28% of the total land mass of Indonesia, with East Kalimantan alone accounting for 10% of the republic. The north of Borneo comprises the Malaysian states of Sarawak and Sabah and the small independent Sultanate of Brunei Darussalam (fig. I.1). Present-day political boundaries are a reflection of past colonial interests.

Despite its large area, Kalimantan supports just 5% of the total population of Indonesia, 9.1 million people (in 1990) at an average density of 17 people per square kilometre. Nevertheless, Kalimantan has played a key role in Indonesia's economic development and is a major earner of foreign revenue. In 1987 just one province, East Kalimantan, accounted for 21% of Indonesia's export revenues. This wealth is not due to industrial production nor to Kalimantan's agriculture or plantations but to the island's rich reserves of natural resources: forests, oil, gas, coal and other minerals. The exploitation of these natural gifts is the major development on Kalimantan. Although this exploitation began much earlier, the pace of development increased rapidly during the 1970s and 1980s to make Kalimantan a key area in the national development process. As development continues and gains momentum, several questions need to be asked. Is exploitation of Kalimantan's natural resources wise in the long-term? Is the exploitation sustainable? Or will it lead to environmental damage, degradation of the soils and forests, and pollution of waterways? What can be done to mitigate such potential damage?

Why is such a large area so underpopulated? Kalimantan's vast open spaces would appear to have massive potential for agricultural development and industrial plantations. The island seems to have great potential for large-scale human resettlement or transmigration from overcrowded Java. For a government planner from Java, this vast green island of low-lying undulating land, with no dry season and plenty of sunshine, must seem full of agricultural promise. Yet agronomists agree that the soils of Kalimantan are very poor, very fragile and very difficult to develop for agricultural use.

Figure I.1. State and provincial boundaries on Borneo.

Kalimantan can be developed, but only within rather tight ecological constraints and only with great care. Already large areas of degraded lands, poorly managed logging operations and failed agricultural schemes scar the Kalimantan landscape. Extensive white sandy padang and red laterised heaths are abandoned where great forests once grew. Each year a great sea of alang-alang grassland curls dry and burns. The forests are not allowed time to regenerate, and the sea of grassland expands, providing little other than fodder for cattle. Fires are set to clear ladang (agricultural fields) and to stimulate new grazing for cattle. Each year, like a ravenous herbivore, the fires nibble away at the forest boundaries. In the drought year of 1982-83 huge fires swept across the forests of East Kalimantan, and drought and fire together caused damage to 3.6 million hectares (36, 000 km²) of forest, an area the size of Belgium. At the same time fires set by agriculturalists raged out of control in Sabah, affecting another million hectares of forest and leading to considerable financial loss. Again the cause for such massive resource loss was poor primary development and careless forestry practice.

River pollution, from opening up the forests, from untreated industrial discharges, from domestic sewage and from illegal methods of gold extraction, is making many waterways dangerous for human use and is causing loss of fish stocks. Clearance of mangroves for tambak (fish pond) development can result in consequent loss of valuable offshore fisheries of shrimp and milkfish. Without careful planning, development of Kalimantan may bring short-term economic gain that creates long-term environmental damage.

Global climate is warming, a result of the greenhouse effect caused by an increase in the amount of carbon dioxide in the upper atmosphere. Indonesia would be one of the countries most affected if sea level were to rise by even a few metres. It would lose a huge area of prime agricultural land, and millions of coastal families would have to be moved to higher ground. Most of the greenhouse gases result from the burning of fossil fuels, but destruction of rainforests also plays a key role, as burning ladang and forests release carbon dioxide into the atmosphere. At the same time, as the area of forest shrinks, the forests become less effective as carbon sinks that remove carbon dioxide from the atmosphere and bind the carbon into productive biomass. As the largest expanse of rainforest in Southeast Asia, Kalimantan's forests are of environmental significance on a global as well as a national scale.

With its great mineral and forest reserves, Kalimantan does have great development potential, but development must be carefully planned with full understanding of the ecology of the land units being developed and of their inter-relationships with surrounding habitats. These constraints can be ignored by developers only at great peril.

Humankind generally has two options: to harvest products from a natural or modified natural productive system, or to replace a natural system with an artificial one. In either case environmental factors such as soil type, slope and climate place strict limits on what types of development are possible. This book aims to outline the ecology, and potential for development, of the various natural land systems of Kalimantan: mountains, lowland forests, special forest types, peat swamps, mangroves, coastal habitats and other landforms.

The book takes a long-term view of development. The rapid harvesting of timber, the stripping of land for underlying minerals, the clearing of hills for two fast crops while the soils are still fertile, these are all highly profitable activities in the short-term, but can leave the land stripped, useless and unproductive. This type of development is hardly desirable. If development is to be useful to the nation it must be sustainable. Development should be seen as an investment for the future, establishing long-term productivity by adoption of harvesting techniques that can continue to give good yields, year after year, without depleting the resource base on which they depend.

The need for sustainable development was the message emphasised in the World Conservation Strategy (UNEP/IUCN/WWF 1980). This message is the theme of this book. The World Conservation Strategy (WCS) identified three main global objectives, all highly relevant to the development of Kalimantan:

maintenance of essential ecological processes and life-support systems;

preservation of genetic diversity; and

sustainable utilisation of species and ecosystems.

Essential ecological processes are those that are governed or moderated by ecosystems and are essential for food production, health and other aspects of human survival and sustained development. Life-support systems are the main ecosystems involved. The most threatened life-support systems are agricultural systems, forests, wetlands and coastal systems.

Kalimantan lacks Java's rich volcanic soils, the source of that island's fertility and agricultural productivity. Agricultural systems in Kalimantan are constrained by low fertility; soil erosion; mineral stress; pernicious weed invasion; failure of irrigation systems; increased flood hazard; salinisation; chemical pollution; loss of natural control agents for insects and other pests; and industrial and urban expansion. The traditional style of shifting agriculture, with short periods of cultivation and long fallow, is well suited to Kalimantan's poor soils, but is being replaced by less appropriate forms of land use.

Forests are being threatened by overexploitation, poor harvesting methods, limited regeneration, lack of reforestation technology, and losses to shifting cultivation and to fire. Kalimantan's important wetlands are threatened by changed rates of water flow, increased erosion and sedimentation, pollution, and drainage for other developments. The coastal areas are threatened by clearance of mangrove and other coastal vegetation, causing loss of fisheries and increasing the danger of marine erosion and storm damage.

The conservation of natural habitats and the preservation of genetic diversity are both an insurance and an investment. They are necessary to sustain and improve agriculture, forestry and fisheries production; to keep open future options; as a buffer against harmful environmental change; and as the raw material for much scientific and industrial innovation. Conservation and protection of biodiversity are matters of prudent management as well as of moral principle.

Much of Borneo is not suitable for sawah (wet rice cultivation), yet the island's productive forests prove that it has high potential for plant growth. Making the best use of the land in the future will certainly depend on finding more suitable crops and appropriate combinations of species to grow. Species that occur naturally in Kalimantan-timber trees, fruits, rattans, medicinal plants-should all be considered for their potential as plantation and agricultural crops. Only a small number of the world's plants have been developed on a commercial scale, yet many more are useful to human communities. The native peoples of Kalimantan use many species for food, medicines and as handicraft and construction materials. It is vital that the widest possible range of plant and animal species is protected as a reservoir for potential new crops and breeding material.

The sustainable utilisation of species and ecosystems is particularly important in an island where such a large proportion of the population depends heavily on direct harvesting from nature (hunting, fishing, collecting firewood and other plant materials) and where forestry is such an important sector of the local and national economy. Development should follow the goal of achieving maximum sustainable yield rather than overharvesting natural resources to extinction.

Sustainable utilisation is analogous to spending interest whilst keeping the capital. A society that insists that all utilisation of living resources be sustainable ensures that it will benefit from those resources virtually indefinitely. Unfortunately, current levels of utilisation of many natural resources in Kalimantan are not sustainable, and some controls will need to be introduced.

This book starts by examining each of the major ecosystems in Kalimantan and the inter-relationships of some of the component species. An attempt has been made to identify the limitations on direct harvesting of products and the vulnerability of different ecosystems to overexploitation and modification. A chapter on Bornean peoples describes their traditional attitudes to, and relationships with, their forest environment. There are many lessons to be learned about sustainable use of natural resources from the island's indigenous peoples. The chapter traces the changing patterns of land use, emphasising some of the problems and dilemmas that now confront the developer and land-use planner.

The book examines the potential of different land units for the development of artificial production systems such as agricultural cropping, plantations, and agroforestry, and the development options open for forests, wetlands and coastal resources. Later chapters outline the ecological consequences that should be considered when planning mineral exploitation and industrial development, and the need to conserve habitats and species on Borneo, which is a global centre for biodiversity.

Although the book is entitled The Ecology of Kalimantan, political boundaries on Borneo bear no relation to biogeographical boundaries and plant and animal distributions. For this reason, many topics such as climate, vegetation and faunal characteristics cover the whole island. Throughout the book considerable reference is made to studies conducted outside Indonesian Borneo, especially where findings are directly relevant to the Kalimantan situation. For the sake of completeness, special Bornean features such as the biology of Sabah's Mount Kinabalu (Borneo's highest mountain) are included. Nevertheless, the text is generally oriented towards Kalimantan, and Indonesian research is quoted wherever appropriate.

This book is a companion volume to The Ecology of Sumatra and The Ecology of Sulawesi (Whitten et al. 1987a, 1987b) which are readily available within Indonesia. Since these volumes are comprehensive in their coverage of basic ecological principles and case studies, coverage of similar topics in this book has been condensed, to avoid duplication and to allow greater emphasis on the wise use of natural resources for sustainable development.

This book was prepared originally between 1988 and 1990 as part of the EMDI project to promote institutional development in the Kalimantan environmental study centres (PSL). The authors worked at the environmental study centre (KPSL-UNLAM) of Universitas Lambung Mangkurat at Banjarbaru, South Kalimantan, and were assisted by collaborative teams at the environmental study centres at Pontianak, Palangkaraya and Samarinda. Changes in project design and objectives have delayed publication, but the manuscript has been revised and updated for this edition.

Chapter One

The Island of Borneo

Borneo straddles the equator between latitudes 7° N and 4° S. It is the third largest island in the world (after Greenland and New Guinea) and the largest land mass in the Sundaic Region, more than five times the size of Java (fig. 1.1). Borneo lies in a region of constant rainfall and of high temperatures throughout the year, ideal conditions for maximum plant growth. As a result, the island has some of the most luxuriant tropical habitats on earth, and contains the largest expanse of tropical rainforests in the Indomalayan realm. The island is rich in biodiversity and is a fascinating place to study natural ecosystems.

The bulk of Borneo (539, 460 km² or 73%) lies within Indonesian territory and is known as Kalimantan; the rest of the island consists of the states of Sarawak and Sabah (together forming East Malaysia) and the oil-rich, independent sultanate of Brunei Darussalam. Together these political states form one geographic unit and share a wealth of biological resources and diverse tropical habitats. Although this book focuses primarily on Kalimantan it begins with a general description of the whole island of Borneo. Geography, geology, climate and human land use have shaped the island's natural ecosystems to create the present Bornean landscape.

GEOGRAPHY

The first-time visitor to Borneo is often surprised at the flatness of the island, with vast areas of low coastal and river plains, especially in the south. Over half the island lies below 150 m in altitude, and water can be tidal up to 100 km inland. Borneo has no active volcanoes, but its main mountain ranges are igneous in origin. The mountain chains run down the island's centre like an inverted trident from north to south, with the three spurs diverging in the south (fig. 1.2). Borneo's highest mountain is Mount Kinabalu in Sabah. At 4, 101 m Kinabalu is the highest peak in Southeast Asia and the highest mountain between the Himalayas and the snowcapped peaks of Irian Jaya. It consists of a granitic plug forced up by volcanic pressures, and it is still rising. Few of the other Bornean mountain peaks exceed 2, 000 m.

Figure 1.1. Borneo compared with other major islands: areas are given in km².

The Iran Mountains, between East Kalimantan and East Malaysia, rise in the north to 2, 160 m at Mount Harden (Harun) near the Sabah border. A western spur of the central Iran mountain chain forms the Kapuas Hulu range along the border between Sarawak and West Kalimantan, rising to Mount Lawit (1, 767 m) and Mount Cemaru (1, 681 m). From the central mountains around Mount Cemaru, the Miiller Mountains (highest point Mount Liangpran 2, 240 m) and Schwaner Mountains (Bukit Raya 2, 278 m) run southwest along the border between West and Central Kalimantan. To the southeast a lower offshoot, the Meratus Mountains, (highest point Gunung Besar 1, 892 m) separates Central and East Kalimantan and extends southwards along the coast. These are all secondary mountain chains with an average height of 1, 000-1, 500 m with only occasional peaks rising above 2, 000 m. In the Iran range, Gunung Makita (2, 987 m) near Longnawan and Gunung Siho (2, 550 m) near Longsaan, both on the Sarawak border, are the highest peaks in Indonesian Borneo, followed by the massif of Gunung Mantam (2, 467 m) west of Tanjung Redeb, East Kalimantan (fig. 1.2). None of the higher Kalimantan mountains are volcanic in origin, in contrast to the volcanic cones that dominate the other Sunda islands and Sulawesi to the east.

Figure 1.2. Map of Borneo showing main mountain ranges, main rivers and lake systems.

Borneo is dissected by great rivers which run from the interior heart lands to the coast and provide the main routes of transportation and communication. The island boasts Indonesia's three longest rivers: the Kapuas (1, 143 km), the Barito (900 km) and the Mahakam (775 km). The Kapuas (as long as the Rhine in Europe) flows west from Mount Cemaru, drains the greater part of West Kalimantan, and is navigable by small steamers 500600 km upriver from Pontianak. Also flowing westwards are the Rajang (Sarawak's longest river) and the Baram. The great Barito originates in the Miiller Mountains and flows southwards, draining the southern swamplands and emptying into the sea near Banjarmasin. Smaller, but historically important, the Kahayan also drains to the south coast. The Kayan and the Mahakam flow from the mountainous interior of the island to the east coast. Sabah's two main rivers, the Segama and the Kinabatangan, also drain eastwards.

Several major river systems have extensive lake systems in their inland basins and lowland reaches. The Mahakam, Barito, Negara, and Kapuas (Kalimantan) and Baram (Sarawak) all have oxbow and seasonal lakes in their lowland floodplains. In the south, Lake Belajau drains into the Seruyan River. The extensive Kapuas, Negara and Mahakam lakes are important inland fisheries (chapters 3 and 10). With much of the lowland plain poorly drained and swampy, overland travel is difficult, and rivers are the main highways into the interior. Human settlements in Borneo are concentrated around the coast and the main rivers and lake systems. Perhaps this explains why the island has been so poorly explored and the lack of reliable maps and topographical data for much of interior Kalimantan.

GEOLOGICAL HISTORY

The biogeography of the whole Indonesian archipelago and the distribution of its soils, plants and animals have been determined by the area's geological and climatic history. This is a story of plate tectonics and continental drift, climatic events and changing sea levels, a story that is still unfolding (Audley-Charles 1981, 1987; Burrett et al. 1991). As recently as 25 million years ago, very recent on the geological time scale (fig. 1.3), the Indonesian archipelago as we know it today simply did not exist, but the story began much earlier than that.

Geologists now recognise that the continental land masses are by no means permanent and that the earth is in a dynamic state. The outer solid part of the earth, the crust, is quite thin, like the rind of an orange. There are two kinds of crust: oceanic and continental. Oceanic crust is usually young (0-200 million years), thin (5-15 km) and composed mostly of dense volcanic rock. Continental crust often has a core of older rocks (200-3, 500 million years), is thicker (20-50 km) and is less dense than oceanic crust, composed of rocks such as sandstones and granites (RePPProT 1990). Western Indonesia, comprised of much of Kalimantan, Sumatra and west and central Java, is composed predominantly of continental crust, as is much of the shallow sea floor between these islands. Below the earth's crust is a zone where the rock is hotter and more plastic. Continental and oceanic plates float on the fluid, underlying material.

Turbulent convection currents rising from the planet's molten core carry the continents and oceanic plates about, creating zones of weakness and disturbance. Where two plates move apart, liquid rock wells up to fill the gap; where two plates collide, one subducts to dive beneath the other, creating deep trenches and crumpled mountain ranges.

Figure 1.3. Geological time scale showing the appearance of major life forms and the occurrence of major geological events.

On a human time scale, plate movements are very slow, only a few centimetres per year, but over a period of 60 million years (the time since the dinosaurs became extinct) a plate drifting only 1 cm per year would have moved 600 km. The Indonesian region is dominated by three major plates, the southeast Asian plate, the Indo-Australian plate and the Pacific plate, as well as several smaller platelets that have sheared off Irian Jaya (Katilil989).

The process is still continuing today. Some of the Indonesian plates move as much as 13 cm per year (RePPProT 1990). Australia is moving northwards and at its present rate of progress will crash through most of Indonesia during the next 50 million years. At the same time the Pacific plates are moving south and west to meet the Australian plate. New Guinea, Java, Sumatra and the islands of Nusa Tenggara lie on the points of impact and are regularly shaken by earth tremors and volcanic eruptions caused by the colliding plates.

Until about 280 million years ago (280 Ma), the earth's land mass formed a single continent, Pangea. During the Triassic, about 250 Ma, Pangea rifted into two supercontinents, Laurasia (including present- day North America, Europe and much of Asia) and Gondwanaland (including present-day South America, Africa, India, Australia, Antarctica and the rest of Asia). The two supercontinents were separated by the Tethys Ocean.

From this stage the early geological history of Indonesia is unclear. Until recently it was believed that the two halves of the Malesian archipelago were derived totally from different supercontinents and had been separate from the Triassic (250 Ma) until a mid-Miocene collision (15 Ma). It was thought that the western half (the Malay Peninsula, Sumatra, Java, Borneo and western Sulawesi) was derived from Laurasia, while the eastern islands, including the rest of Sulawesi, were derived from Gondwanaland much later (Audley-Charles 1981). According to this model, much of what is now mainland Southeast Asia, including Kalimantan, was on the southern side of the large continental mass of Laurasia (Sengor 1988). By 260 Ma this southern margin was the site of a subduction zone and volcanic arc (RePPProT 1990). Old volcanic rocks in Kalimantan date from this period and are associated with limestones in South Kalimantan, indicating that fringing coral reefs were present as they are today around volcanic islands in Nusa Tenggara.

To the south, separated from Laurasia by the Tethys Ocean, lay the continental mass of Gondwanaland. The northern flank of Gondwanaland was unstable, and fragments split off and were transported northwards, where they eventually collided with the southern flank of Laurasia between 230 Ma and 200 Ma. The older or basement rocks of much of Sumatra and Peninsular Malaysia and the surrounding continental crust were welded onto Borneo to create the block of continental crust known as Sunda-land, about 180 Ma. As a result of the collision a large amount of granite was intruded in the Thailand-Malaysia-Bangka-Belitung-West Kalimantan zone; economic tin deposits are associated with much of this granite (RePPProT1990).

Between 200 Ma and 150 Ma, further fragmentation of Gondwanaland occurred, and Africa, India, Australia and Antarctica (until then a single land mass) started to separate. During the Mesozoic, tectonic movements caused the Tethys Ocean to move northwards accompanied by subduction and arc formation at the southern side of Sundaland. The late Mesozoic granites of Kalimantan date from this time (RePPProT 1990). The most dramatic result of the breakup was the separation and rapid movement northward of India, which collided with the Laurasian plate. This massive collision destroyed the Tethys Ocean and caused the uplift of the Himalayas.

In the light of more recent paleontological and geological discoveries, an alternative theory has been proposed. This suggests that most of Southeast Asia, Sumatra and Borneo were not part of Laurasia but separated from Gondwanaland much later, in the mid-Jurassic and Cretaceous (Audley-Charles 1987; Burrett et al. 1991). Veevers (1988) recognised three main stages of rifting on Australia's northwest shelf: Cambrian (equated with the rifting of North China), late Carboniferous to early Permian (rifting of Shan Thai), and the late Jurassic. The Jurassic-Cretaceous event rifted the many Gondwana fragments now found in Indonesia (Burrett et al. 1990). These continental fragments rifted from Australia-New Guinea (the northern margin of east Gondwanaland) and provided stepping-stones of dry land during the Tertiary. Parts of Sumatra and southern Kalimantan probably rifted from Australia in the early Cretaceous. Northern Borneo had probably rifted even earlier. These and other Southeast Asian blocks became relatively isolated within the Tethys Ocean between Gondwanaland and the Asian mainland for perhaps as long as 60 million years, but from the late Cretaceous (100 Ma) onwards they provided an archipelago of islands that could have permitted land plant dispersal between the Australasian and Asian land masses (fig. 1.4).

About 90 Ma the continental plate of Australia and New Guinea broke away from Antarctica and began its rapid drift northwards. During the Paleocene (60 Ma) northern Borneo and southern Kalimantan were quite separate, but they probably collided in the Eocene, 50 Ma (Pieters and Supriatna 1990). By the Oligocene (30 Ma) Borneo was part of a land mass that also included western Sulawesi (Burrett et al. 1991). By 20 Ma the Australian plate was beginning to interfere with the volcanic arc on the southern edge of Sundaland. Fragments of the continental plate sheared off and were forced westward, accommodated by subduction of the ocean floor beneath Sundaland. The plate fragments derived from the area of Irian Jaya have collided progressively with eastern Sundaland to cause trapping and then obduction of the sea floor to form ophiolites in the Meratus Mountains of Kalimantan, in east Sulawesi and in Halmahera (Moluccas).

Figure 1.4a. Reconstruction for Paleocene 60 Ma. NKAL, North Kalimantan.

Figure 1.4b. Reconstruction for Oligocene 30 Ma. South Kalimantan and northern Borneo already joined.

Figure 1.4c. Reconstruction for Late Miocene 10 Ma.

Figure 1.4. The changing locations of the components of Southeast Asia since the first rifting from Gondwanaland.

Source: Burrett et al. 1991

The last 15 million years have been a time of dramatic geological activity in the Indonesian archipelago, marked by the uplift of sedimentary rocks from the ocean bed and the creation of new volcanic islands. Tectonic movements, resulting from the northwards and westwards movements of the major plates, led to the uplift of new land from the sea bed and the upthrust of the Bornean mountain blocks. During the later Tertiary, erosion of these mountain chains created thick sedimentary deposits. In Borneo the thickest and most economically important of these Tertiary sedimentary sequences were deposited in deltas, such as the Mahakam delta in East Kalimantan. Biological material washed into the Mahakam basin was changed by heat and pressure into coal and oil deposits.

During the Tertiary and Quaternary, the Indonesian archipelago experienced periods of lowered sea levels and more seasonal climates than those recorded today. Sea level changes during the Tertiary resulted from tectonic activity. The expansion of the oceans due to sea floor spreading resulted in marked changes in ocean volumes. The mid-Oligocene onset of Antarctic glaciation also caused a marked lowering of global sea levels (fig. 1.5).

The late Cenozoic was a period of repeated alternations of glacial and interglacial periods. The major climatic cycle, with a period of about 100, 000 years, was probably initiated by astronomical forces, originating from small changes in the ellipticity of the earth's orbit. During the Quaternary the resulting changes in insolation were enhanced by changes in the carbon dioxide (CO2) concentration in the atmosphere. Since concentration of atmospheric CO2 causes the greenhouse effect (chapter 13), lowered CO2 at glacial maxima may have had a profound influence on world temperatures (Morley and Flenley 1987). Simultaneously the increase in albedo (whiteness) of ice-covered surfaces causes decreased absorption of sunlight in glacial regions, explaining why temperate climates fluctuated more than tropical ones in the Quaternary.

Changes in sea level during the Quaternary can be calculated with some precision from oxygen isotope data. The maximum lowering of the sea level in the Quaternary was about 200 m, which would have exposed the Sunda platform and the Sahul Shelf. Submarine channels on both shelves indicate the courses of rivers when these continental platforms were above sea level (Verstappen 1975). In the Malesian region sea level reached its minimum, and land areas were most exposed, in the middle Pleistocene. Later, in the Holocene, sea levels were slightly above present levels (fig. 1.6; Tjia et al. 1984). Reduction in sea levels provided land linkages from mainland Southeast Asia to the islands of the Sunda Shelf.

Figure 1.5. Hydrological cycles (a) in warm conditions, and (b) in cold conditions, when sea level falls as water is retained in the ice cap.

There is considerable evidence for more strongly seasonal climates in both the late Tertiary and the Quaternary. Studies of peat swamps and analysis of the pollen record indicate that the climate of South Kalimantan was much more seasonal then than today (Morley 1982).

The more seasonal climates of the late Tertiary and mid-Pleistocene created a savanna corridor between Southeast Asia and Borneo which allowed migration of savanna plants and animals across the area from Thailand to Java. Cooler climatic conditions in Asia during the glacial periods encouraged animals to move southwards along the land bridges over a period of half a million years, and successive waves of animal, and human, immi grants were able to reach the Greater Sunda islands from Asia. During glacial maxima, cooler temperatures lowered montane vegetation zones, providing stepping-stones for the migration of mountain and temperate taxa. During the Pleistocene, glacial periods were longer than the interglacial periods, and for the greater part of the last two million years most of the Sunda Shelf was exposed above sea level (fig. 1.7) but everwet habitats were less extensive than today. Late Tertiary and Quaternary paleoclimates thus help to explain many disjunct plant and animal distributions in the Indonesian archipelago (Morley and Flenley 1987).

Figure 1.6. Changes in sea level on the Sunda Shelf during the last 6, 000 years. (After Tjia 1980.)

At the times of lowest sea level during the Pleistocene, all the islands on the Sunda Shelf—Java, Borneo, Sumatra, Bali and Palawan—were thus connected by land bridges. Connections were formed and broken several times. Java and Palawan, at the extremities of the shelf, were connected less often and less recently than either Borneo or Sumatra, and this is reflected in their respective distinctiveness.

Figure 1.7. A generalised paleogeographical reconstruction of the Sunda-Sahul region during one of the Quaternary glacial maxima. A savanna corridor may have extended across Sundaland in the middle Pleistocene. Everwet refuges may have occurred in montane areas. (After Morley and Flenley 1987.)

Even at the times of lowest sea levels in the late Cenozoic, Borneo was not connected to Sulawesi. Thick sediments in the Makassar straits indicate that Borneo and Sulawesi have been separate for at least 25 Ma (Audley-Charles 1981) and possibly longer. This long isolation is reflected in differences in flora (Balgooy 1987), lepidopterans (Holloway 1987) and mammals (Musser 1987) between Borneo and Sulawesi. Sulawesi's flora and fauna show much stronger similarities to those of the Philippines and Lesser Sundas than to those of Borneo and the other Sunda islands.

The shallow continental shelves of the South China Sea and Java Sea are incised with several ancient river channels, including three major rivers, between Borneo and Sumatra: the Anambas, the North Sunda (with the proto-Kapuas as a tributary), and the proto-Lupar valleys (Haile 1973; fig. 1.8). Two large, parallel rivers run along the bed of the Java Sea between Java and Borneo, towards the Straits of Makassar (Verstappen 1975). Similarities in freshwater fish between western Borneo and Sumatra indicate that the rivers of these islands were once linked; the Musi of Sumatra and the Kapuas of Borneo were once part of the North Sunda river system. Some of these large rivers probably served as barriers to floral and faunal distribution; dipterocarps show some breaks in distribution at the Lupar River (Ashton 1972). Similarly, the major rivers between Borneo and Java probably slowed faunal dispersals between the two islands.

Figure 1.8. The Sunda Shelf showing present coastlines (unshaded), the area of Sundaland exposed at times of lowest sea level (dark shade) during the last glacial about 12, 000 years ago, and past and present river systems. (After Tjia 1980.)

The climate of the region was probably only 2°-4°C cooler in the Quaternary and Pleistocene than it is today, but conditions were often more arid (Morley and Flenley 1987). Elsewhere climatic changes have been cited as a cause of species extinctions, but of 200 large mammal extinctions worldwide in the Pleistocene only 11 occurred in Southeast Asia (Medway 1972b). Of the 32, 000-year-old fauna recorded from excavations at Niah only one species, the giant pangolin, is truly extinct (Harrisson 1961), although other mammals, such as the tapir and Javan rhino, have been locally extirpated in Borneo, probably by overhunting. The lesser gymnure Hylomys suillus and ferret badger Melogale orientalis recorded from the Niah excavations are now found only on the higher slopes of Mount Kinabalu, which lends support to the theory of a cooler climate at Niah in the Late Pleistocene.

GEOLOGY

Publication of systematic geological maps is less advanced for Kalimantan than for any other part of Indonesia (RePPProT 1990). Table 1.1 gives a simplified overview of major geological events in Borneo. Figure 1.9 illustrates the geology of Kalimantan. There are four main geological units represented in Kalimantan: rocks associated with plate margins; basement rocks; younger consolidated and nonconsolidated rocks; and alluvium and young superficial deposits.

The Kalimantan basement complex in west and central Kalimantan (including the Schwaner Mountains) represents the largest continental basement outcropping in Indonesia. Basement rocks, at the bottom of the stratigraphic sequence, are generally older than the overlying rocks. They are usually metamorphosed by heat. Typical products of metamorphism are marbles (from alteration of limestones), green schists from volcanic rocks, and gneiss from sandstone or granite. Areas of metamorphic rock or basement are typical of continental crust, often invaded by younger intrusive rocks. The Kalimantan basement complex consists of schist and gneiss intruded by granites of Paleozoic, Mesozoic and Tertiary age to form a widespread crystalline terrain.

Rocks associated with plate margins in Kalimantan include ophiolites (derived from oceanic crust) and melange. Fragments of oceanic floor are found on land in several parts of Borneo. Characteristically they are composed of dark, dense igneous rocks of basic and ultrabasic type with a granite component. Thin oceanic siliceous sediments (chert) and carbonate sediments may also be present. This suite of rocks is called ophiolite. Such ophiolites are created by plate collisions when oceanic crust, instead of subducting, is trapped by plate tectonic movements and forced onto the adjacent plate margin where it remains exposed. This obduction process is often accompanied by crumpling and fracture of the rocks. The ophiolite complexes of Pulau Laut and the Meratus Mountains originated in this way.

Melange is a mixture of broken fragments of different rock types and sizes in a sheared clayey matrix, indicating very strong compression. Fragments can be very small (a few centimetres) to large (hundreds of metres or more). Melange is often associated with the subduction process. It is a combination of material scraped off the downgoing oceanic plate with sediments derived from the nearby land mass or volcanic arc. The whole mass is squeezed and sheared due to the underthrust of the downgoing plate. Rocks formed in this way are associated with the large upthrust slabs of ophiolite in the Meratus Mountains.

Table 1.1. Summary of geological time scale and events relevant to Borneo over the last 350 million years.

Figure 1.9. Geology of Kalimantan.

Source: RePPProT 1990

The large area of melange in the centre of Borneo, straddling the border between Kalimantan and Malaysia, is less well understood. This is a zone of broken rocks often including fragments of ophiolite, but its width and extended geological age (late Mesozoic to older Tertiary) are difficult to explain in terms of simple plate tectonics (Williams et al. 1989).

Much of Kalimantan consists of consolidated and semiconsolidated rocks, including Quaternary limestones (in the Sangkulirang peninsula and Meratus range), volcanic rocks, and Tertiary sediments. Although Borneo has no active volcanoes, as do Sumatra and Java, it does have substantial areas of old volcanic rocks in southwest and eastern Kalimantan. These are a legacy of Indonesia's geological history, which has included many periods of volcanic activity from 300 Ma to the present day. Volcanic rocks are formed when magma from deep in the earth reaches the surface. Where magma cools and consolidates below the earth's surface, intrusive rocks such as granodiorite are formed. Where the old Kalimantan volcanic rocks have been eroded, stocks of gold-bearing intrusives, originally below the volcanoes, have been exposed. The interaction of magma with groundwater beneath volcanoes is an important part of the process