A review of dipterocarps - Center for International Forestry Research

A Review of 

Dipterocarps 

Taxonomy, ecology and silviculture 

Editors 

Simmathiri Appanah 

Jennifer M. Turnbull

A Review of Dipterocarps: 

Taxonomy, ecology and silviculture 

Editors 

Simmathiri Appanah 

Jennifer M. Turnbull 

CIFOR ÃÃÃÃÃÃÃÃÃ 

FOREST RESEARCH INSTITUTE 

MALAYSIA

ã 1998 by Center for International Forestry Research 

All rights reserved. Published 1998. 

ISBN 979-8764-20-X 

Cover: Dipterocarp forest and logging operation in Central Kalimantan, Indonesia. 

(photos by Christian Cossalter) 

Center for International Forestry Research 

Bogor, Indonesia 

Mailing address: P.O. Box 6596 JKPWB, Jakarta 10065, Indonesia 

Tel.: +62 (251) 622622; Fax: +62 (251) 622100 

E-mail: cifor@cgiar.org 

Website: http://www.cgiar.org/cifor

Contents 

Authors 

Abbreviations 

Acknowledgements 

Foreword 

Introduction 

S. Appanah 

Chapter 1. Biogeography and Evolutionary Systematics of Dipterocarpaceae 

G. Maury-Lechon and L. Curtet 

Chapter 2. Conservation of Genetic Resources in the Dipterocarpaceae 

K.S. Bawa 

Chapter 3. Seed Physiology 

P.B. Tompsett 

Chapter 4. Seed Handling 

B. Krishnapillay and P.B. Tompsett 

Chapter 5. Seedling Ecology of Mixed-Dipterocarp Forest 

M.S. Ashton 

Chapter 6. Root Symbiosis and Nutrition 

S.S. Lee 

Chapter 7. Pests and Diseases of Dipterocarpaceae 

C. Elouard 

Chapter 8. Management of Natural Forests 

S. Appanah 

Chapter 9. Plantations 

G. Weinland 

Chapter 10. Non-Timber Forest Products from Dipterocarps 

M.P. Shiva & I. Jantan 

Scientific Index 

General Index 

v 

vii 

ix 

xi 

1 

5 

45 

57 

73 

89 

99 

115 

133 

151 

187 

199 

209

Authors 

S. Appanah 

Forest Research Institute Malaysia 

Kepong 

52109 Kuala Lumpur 

Malaysia 

M. S. Ashton 

School of Forestry and Environmental Studies 

Yale University 

Marsh Hall, 360 Prospect Street 

New Haven, CT 06511 

USA 

K. S. Bawa 

Department of Biology 

University of Massachusetts 

100 Morrissey Boulevard 

Boston MA 02125-3393 

USA 

L. Curtet 

Laboratoire de Biométrie, Génétique et Biologie 

des Populations 

Université Claude Bernard - LYON 1 

43, Boulevard du 11 Novembre 1918 

FR-69622 Villeurbanne Cedex 

France 

C. Elouard 

French Institute of Pondicherry 

11, St. Louis Street 

P.B. 33, Pondicherry 605001 

India 

I. Jantan 

Universiti Kebangsaan Malaysia 

50300 Jalan Raja Muda Abdul Aziz 

Kuala Lumpur 

Malaysia 

B. Krishnapillay 


Kepong 


Malaysia 

S. S. Lee 


Kepong 


Malaysia 

G. Maury-Lechon 

U.M.R. C.N.R.S. 5558 

Laboratoire de Biométrie, Génétique et Biologie 

des Populations 

Université Claude Bernard - LYON 1 

43, Boulevard du 11 Novembre 1918 

FR-69622 Villeurbanne Cedex 

France 

M. P. Shiva 

Centre of Minor Forest Products 

HIG-2, No. 8, Indirapuram 

Gen. Mahadev Singh Road 

P.O. Majra, Dehra Dun 248 171 

India 

P. B. Tompsett 

RBG Kew 

Wakehurst Place 

Ardingly, Haywards Heath 

Sussex, RH17 7TN 

United Kingdom 

G. Weinland 

Malaysian-German Sustainable Forest 

Management and Conservation Project 

GTZ 

Jalan Sultan Salahuddin 


Malaysia

Abbreviations 

ABA Abscisic acid 

ACOM Asian Conference on Mycorrhizae 

AFTSC ASEAN Forest Tree Seed Centre 

ASEAN Association of Southeast Asian 

Nations 

ASTAG Agriculture Division in the Asian 

Technical Department, World Bank 

(ceased January 1993) 

BHC Benzene hexachloride 

BIO-REFOR Biotechnology assisted 

Reforestation 

BIOTROP See SEAMEAO-BIOTROP 

CIFOR Center for International Forestry 

Research 

DABATTS Database of tropical tree seed 

research 

DENR Department of Environment and 

Natural Resources, Philippines 

DFID Department for International 

Development (United Kingdom) 

DNA Deoxyribonucleic acid 

EEC European Economic Community 

FAO Food and Agriculture Organization of 

the United Nations 

FD Forest Department of Peninsular 

Malaysia 

FORSPA Forestry Research Support Program 

for the Asia-Pacific 

FRIM Forest Research Institute Malaysia 

GTZ Deutsche Gesellschaft für 

Technische Zusammenarbeit 

IBPGR International Board for Plant Genetic 

Resources (now IPGRI) 

ICFRE Indian Council of Forestry Research 

and Education 

IIED International Institute for 

Environment and Development 

IPGRI International Plant Genetic 

Resources Institute 

ITTO International Tropical Timber 

Organisation 

IUCN The World Conservation Union 

IUFRO International Union of Forestry 

Research Organizations 

IWGD International Working Group on 

Dipterocarps 

JICA Japan International Cooperation 

Agency 

JIRCAS Japan International Research Centre 

LN Liquid nitrogen 

LSMC Lowest-safe moisture content 

MC Moisture content 

MP Melting point 

MTC Malaysian Timber Council 

MUS Malayan Uniform System 

NCT Non-crop trees 

NTFPs Non-timber forest products 

ODA Overseas Development Authority 

(United Kingdom) (now DFID) 

OLDA Orthodox with limited desiccation 

ability 

OTA Office of Technology Assessment 

PAR Photosynthetically active radiation 

PCARRD Philippine Council for Agriculture, 

Forestry and Natural Resources 

Research and Development 

PCT Potential final crop trees 

PEG Polyethylene glycol 

PSLS Philippine Selective Logging System 

RAPA Regional Office for Asia and Pacific 

(FAO) 

RAPD Random Amplified Polymorphic 

DNAs 

RIF Regeneration Improvement Fellings 

RIL Reduced Impact Logging 

ROSTSEA Regional Office for Science and 

Technology for South East Asia 

(UNESCO) 

SEAMEO- 

BIOTROP South-East Asian Regional Centre for 

Tropical Biology 

SMS Selective Management System 

SPDC Special Programme for Developing 

Countries (IUFRO)

SPINs Species Improvement Network 

TPI Tebangan Pilih Indonesia (Indonesian 

Selective Cutting System) 

TPTI Tebang Pilih Tanam Indonesia 

(Modified Indonesian Selective 

Cutting System) 

TROPENBOS The Tropenbos Foundation, 

Netherlands 

viii 

TSI Timber Stand Improvement 

UNESCO United Nations Educational, 

Scientific and Cultural Organization 

UPM Univesiti Pertanian Malaysia 

(Agriculture University of Malaysia) 

USDA United States Department of 

Agriculture 

VAM Vesicular arbuscular mycorrhizas


The dedication and enthusiasm of the authors have contributed to make this book what it is. Our special thanks go to 

Christian Cossalter of the Center for International Forestry Research for his major role at the start of the project 

deciding content, general structure and authorships and later in arranging external reviewers. His attention and support 

has freed us from the day-to-day problems of bringing such a book to completion and allowed us to concentrate 

on editorial tasks. We would also like to thank those who reviewed the various chapters: they are P.S. Ashton (Harvard 

Institute for International Development), Peter Becker (Universiti Brunei Darussalam), Tim Boyle (Center for International 

Forestry Research), P. Burgess, P. Moura-Costa (Innoprise), K.S.S. Nair (Kerala Forest Research Institute), 

F.E. Putz (Center for International Forestry Research), Manuel Ruiz-Perez (Center for International Forestry 

Research), Willie Smits (The International MOF TROPENBOS – Kalimantan Project), Paul B. Tompsett (Royal 

Botanic Gardens Kew), Ian Turner (National University of Singapore) and T.C. Whitmore (Cambridge University). 

Our warm thanks go also to Rosita Go and Meilinda Wan for secretarial assistance, Gideon Suharyanto for the 

layout, Paul Stapleton for the cover design, Patrick Robe for the scientific index and Michael Harrington for the 

general index. The photographs used in this book have been supplied by Christian Cossalter. 

The editors 

Simmathiri Appanah and Jennifer M. Turnbull

Foreword 

The Center for International Forestry Research (CIFOR) 

was established in 1993 at a time when there was a 

resurgence of interest in the sustainable management of 

the world’s tropical rain forests. At that time it was 

evident that a particular focus for CIFOR’s research 

should be in the moist tropical forests of Asia. Trees in 

the family Dipterocarpaceae, “the dipterocarps” are a 

major component of southeast Asia’s tropical forests. 

Their wood is pre-eminent in the international tropical 

timber trade and they play a key role in the economies 

of several countries. 

A considerable research effort had already been 

devoted to the management and utilisation of dipterocarp 

forests starting with the British in India last century and 

continuing throughout the 20th century, especially in 

Malaysia. A vast amount of information has been 

gathered, but unfortunately it has not been consolidated 

and no readily accessible compilation of results has been 

available. This has reduced the impact of the research 

and has almost certainly resulted in the duplication of 

efforts by national and international bodies. 

As a new international research centre it was 

appropriate that CIFOR should take the initiative and 

commission a general review of the current state of 

knowledge of dipterocarp taxonomy, ecology and 

silviculture, to identify gaps in this knowledge and to 

spell out priority areas for new research. This action 

accorded with the views of many members of the 

informal Round Table on Dipterocarps who had been 

meeting on a regular basis to share information on the 

family . A draft outline of the book was endorsed by the 

Fifth Round Table on Dipterocarps at its meeting in 

Chiang Mai, Thailand late in 1994. Since then, under the 

direction of Christian Cossalter at CIFOR and Dr S. 

Appannah at Forest Research Institute Malaysia (FRIM), 

13 authors have prepared and revised the 10 chapters of 

the book. With authors located in Asia, Europe and the 

United States this has been a major undertaking and the 

efforts of all concerned to bring this work to a successful 

conclusion are very much appreciated. 

I anticipate that this book will be especially 

beneficial to those planning research on dipterocarps in 

Asia. I hope it will assist university graduate and postgraduate 

researchers, and especially scientists in national 

and international organisations to re-orient their research 

to meet priority needs. The review should also be useful 

to forest managers in both public and private sectors who 

must make decisions based on whatever information is 

available to them and who have neither the time nor the 

resources to delve into the highly dispersed literature 

on dipterocarps. 

CIFOR is very grateful to many people for their 

assistance with this book; to all the contributing authors 

for their commitment and for their patience with demands 

made on them by the editors; to the reviewers who 

provided critical appraisals of the chapters and made 

valuable inputs; to the editors who brought all the 

contributions together and completed endless checking 

and cross-checking of the information: to the CIFOR 

Communications Group for typesetting and layout; and 

finally to the staff of the Forest Research Institute 

Malaysia and its Director General Dr. M.A.A. Razak for 

their unflagging support and cooperation in producing 

this book. I thank all who contributed in so many ways. 

Prof. Jeffrey Sayer 

Director General of CIFOR

Introduction 

S. Appanah 

As a family of plants, Dipterocarpaceae may perhaps hold 

the distinction of being the most well known trees in the 

tropics. This famed family of trees stand tall in some of 

the grandest forest formations the earth has ever 

witnessed. Their overwhelming presence has led us to 

call these vegetation zones dipterocarp forests. Currently 

the dipterocarps predominate the international tropical 

timber market, and therefore play an important role in 

the economy of many of the Southeast Asian countries 

(Poore 1989). The dipterocarps also constitute important 

timbers for domestic needs in the seasonal evergreen 

forests of Asia. In addition, these forests are sources of 

a variety of minor products on which many forest 

dwellers are directly dependent for their survival 

(Panayotou and Ashton 1992). Despite such eminence 

in the plant world, there has never been an attempt to 

assemble under one cover all the principal aspects of 

this exceptional family of trees. This is a serious lack 

which we hope to start redressing and thus pay fitting 

tribute to this great family of trees. 

A greater concern however belies this slim effort. 

The very existence of these trees and the forests they 

stand in is at stake today because of the unrelenting pace 

at which we are chopping down these forest giants and 

converting their forests to other forms of landuse (FAO 

1989). If present trends persist, not only will nations 

and people become impoverished, but mankind will stand 

to lose many species of plants and animals forever. These 

dipterocarp forests, especially those everwet formations 

of West Malesia, are among the richest worldwide in 

terms of flora and fauna (Whitmore 1975). 

Much of the knowledge on the species within the 

Dipterocarpaceae exists in a disparate form even though 

research on dipterocarps extends for about a century, 

almost since the beginning of tropical forestry in British 

India. Apart from some classical work on their taxonomy 

(e.g. Symington 1943) and silviculture (Troup 1921, 

Wyatt-Smith 1963), most other studies remain 

fragmented. A uniform and comparative body of 

information on dipterocarps did not develop. Studies 

equivalent to those on acacias or eucalypts in Australia 

never resulted (e.g. Jacobs 1981). This situation is the 

result of a number of factors including: 

1. The dipterocarps that comprise timber species are 

distributed over a very wide range throughout tropical 

Asia, covering several climatic zones and 

geographies. The number of species in each country 

varies from 1 to over 200 (Ashton 1982). 

Consequently the depth of interest differs from 

country to country. 

2. The historical emphasis upon forest management 

differs between countries, and this is reflected in 

differences in institutional strengths and development 

in research. While the dipterocarps are managed in 

some countries, in other locales they are simply 

exploited. A quick glance at the status of knowledge 

on the dipterocarps in the region confirms this 

unevenness. In some locations, the Indian continent 

for example, the knowledge on many aspects of 

dipterocarps is comprehensive. In others like Laos 

and Cambodia, it varies from fragmentary to cursory. 

3. Whatever scientific links that existed during the 

colonial period have broken down. In fact, the first 

forester brought in to attend to Malayan forest needs 

was from British India (Wyatt-Smith 1963). Today, 

scientific links between countries sharing the 

dipterocarps have become desultory. 

4. A considerable amount of information is sitting in 

national institutes either in unprocessed form in 

departmental files, or as internal reports, unpublished 

theses, etc. Some reports are written in the local 

language. Thus, a substantial wealth of knowledge is 

simply not available to the vast majority of scientists.

Introduction 

As a consequence, much of the knowledge on 

dipterocarps appears to be accessible only to specialists. 

The potential benefits of this family have not been fully 

recognised, and if the present situation is allowed to 

proceed, mankind may lose important opportunities. The 

following examples affirm this view. Few realise that the 

only moist tropical forests in the world where sustainable 

forest management has been demonstrably practiced are 

the dipterocarp forests (FAO 1989). The best silvicultural 

system that was ever formulated for a tropical forest is 

perhaps the Malayan Uniform System which is based on 

the exceptional regeneration properties of dipterocarps 

(Wyatt-Smith 1963). In fact dipterocarp forests are the 

envy of foresters and silviculturists toiling in the African 

and neotropical areas. However, these facts are seldom 

if ever highlighted. 

The general lack of comprehension about the family 

has led to a tide of opinion that it is not possible to 

manage tropical forests, an opinion strongly contested 

by those involved in dipterocarp forest management. Few 

realise that the apparent failures in establishing 

sustainable yields were more the result of changes in 

landuse patterns and economic restructuring than from 

an inherent inability of the forest to respond to 

appropriate silvicultural interventions (Appanah and 

Weinland 1990). To a degree, this lack of understanding 

has led us to exploit the forests somewhat carelessly 

without considering the wonderful opportunities they 

offer for practicing sustainable forestry. 

This ignorance of the qualities of dipterocarps has 

also led us to search elsewhere for usable tree species 

when interest in timber plantations for the moist tropics 

developed (e.g. Spears 1983). The general impression 

was that dipterocarps, as a group, are slow growing and 

planting material difficult to procure. Such overgeneralisations 

made us miss some important 

opportunities with dipterocarps. There are dipterocarps 

which make excellent plantation species (Appanah and 

Weinland 1993), and several have growth rates that are 

acceptable or superb for this purpose (Edwards and Mead 

1930). Few recognise the potential of dipterocarps with 

their mycorrhizal associations to grow under poorer soil 

conditions. Nor has attention been focused on the variety 

of dipterocarp species available that are adapted to a wide 

range of habitats and edaphic conditions making it 

possible to match species to specific conditions in 

plantations. 

Now that attempts to establish fast growing hardwood 

plantations based on exotic timber species in moist 

forests of Asia have met with many difficulties, there is 

a resurgence of interest in indigenous species for this 

purpose. Many of the species under consideration are 

dipterocarps (Anon. 1991). Throughout Southeast Asia, 

plans for planting dipterocarps are regularly announced 

while major reforestation activities are often based on 

the use of species from this family. Meanwhile, forest 

scientists and managers from all over the world are 

looking to dipterocarp forests to provide models for 

sustainable forest management for the moist tropics and 

ensure a steady supply of industrial wood in the future. 

Currently, numerous initiatives, both national and 

international, are underway to address the variety of 

issues related to dipterocarps and dipterocarp forests. 

These issues under investigation cover a very wide 

spectrum, from basic management issues (e.g. National 

Institutes, Food and Agriculture Organization (FAO), 

International Tropical Timber Organisation (ITTO), 

Department for International Development (DFID)), to 

producing quick field identification guides (DFID), and 

biodiversity (DFID), ecology and economics (National 

Science Foundation, Center for Tropical Forest Science), 

vegetative propagation (TROPENBOS, Japanese 

International Cooperation Agency), mycorrhiza 

(TROPENBOS, National Institute for Environmental 

Studies, European Commission), non-timber forest 

products, plantations (ITTO, TROPENBOS, Forestry 

Research Support Programme for Asia and the Pacific), 

and so on. In addition to the interest in planting 

dipterocarps, there is also a general surge of excitement 

over all other aspects of this family. Some major studies 

currently underway include sustainable management of 

dipterocarp forests (Sabah Forest Department/Deutsche 

Gesellschaft Fuer Technische Zusammenarbeit MbH) 

and carbon sequestration and reduced impact logging 

(Forest Absorbing Carbon-dioxide Emissions 

Foundation). 

While the above endeavours are laudable, and bear 

testimony to the value of dipterocarps, we view this 

proliferation of apparently uncoordinated initiatives with 

some concern. Undoubtedly, these undertakings are 

going to vastly increase our knowledge of the trees and 

the ecosystem, so that in the final analysis we get closer 

to our ultimate goal – the ability to manage these forests 

on a sustainable basis. But at what price in terms of 

2

Introduction 

efficient use of resources? Several issues require further 

reflection: 

1. While dipterocarps may seem to hold better prospects, 

one should not be trapped into the notion that 

they are the solution to our present problems. The 

difficulties encountered with planting of exotics are 

not limited to biological constraints (e.g. Evans 1982, 

Appanah and Weinland 1993). Management and economic 

issues played just as big a role in these difficulties. 

The same difficulties could be encountered 

with planting of dipterocarps. Therefore, past experiences 

should be analysed and/or new work started 

in areas like species trials, provenance testing, seed 

orchards, selection of plus trees, vegetative propagation, 

etc. 

2. There is a general lack of coordination between and 

among external agencies and international donors for 

most of the initiatives. While duplication of activity 

is common, experiences are rarely shared, leading 

to adoption of practices that have been proven to have 

no potential. Furthermore, if such duplication of research 

had been avoided, perhaps funds and resources 

could have been applied more optimally. 

3. The lack of a common and easily accessible body of 

information on dipterocarps has had unfortunate impact 

on the development of moist forest management 

techniques. Many a trial, effectively proven unworkable, 

is repeatedly tried out elsewhere in blissful ignorance, 

sometimes even in the same locale by a fresh 

generation of researchers and managers, while documentation 

of the previous experiences remained 

locked away in dusty filing cabinets. Lessons learned 

in the past have been misunderstood, forgotten or 

simply not recognised. One notorious example is the 

case of underplanting with dipterocarps. Despite 

ample proof that dipterocarps will need a reasonable 

amount of direct light for fast growth, even today 

hundreds (or even thousands) of hectares of exotic 

plantations have been underplanted with dipterocarps 

in several countries. Such trials are doomed to fail. 

4. Even the practice of silviculture has not been free of 

this repetition of mistakes. Here there appears to be 

a tendency to start at the bottom when it comes to 

research. Seldom research is initiated that follows 

through findings of previous researchers. A thorough 

understanding of past research seems to elude the 

next generation of scientists. Examples of such cases 

are disconcertingly numerous. For example, in the 

1930s the classic Departmental Improvement 

Fellings in Malaya were found incapable of releasing 

the bigger poles and residuals, unless the fellings 

were repeated several times at a high cost (Barnard 

1954). Instead, such fellings released the young regeneration. 

In the 1970s, the same approach under a 

different name, called Liberation Felling was adopted 

in Sarawak (see FAO 1981). The results were the 

same. However, the recognition that both these systems 

are the same in principle has not yet been appreciated 

by most forest scientists. 

5. Research on dipterocarps is still being carried out 

within the confines of narrow disciplines, and problem-oriented, 

multi-disciplinary approaches are indeed 

rare. Notable cases exist even within the same 

research institutions with their silviculturists and 

forest managers carrying out reforestation programs 

without the benefits of inputs from tree breeders and 

geneticists, while the latter appear more interested 

in theoretical, evolutionary issues. 

In conclusion, we can state that our efforts to manage 

dipterocarp forests is pitted with difficulties: missed 

opportunities, workable schemes arriving too late, and 

mistakes repeated time and again. There is no guarantee 

that this situation will not perpetuate unless we rethink 

our approach to the whole research and development 

question. Otherwise more mistakes will be made, more 

trials and management systems will fail, and the 

conclusions will point in the most negative direction – 

that it is not possible to manage tropical rain forest. This, 

we have to avoid. Time is also against us, considering 

the rate at which these forests are being logged. 

1. In the first instance, there is a need for thorough reviews 

of former research as well as application trials, 

both at country and regional levels. Agencies such 

as Center for International Forestry Research 

(CIFOR), FAO and Asian Development Bank are well 

placed to initiate these reviews. These, while pointing 

out the successful methods, should at the same 

time identify the unsolved problems and gaps in research 

for which urgent work is needed. 

2. Armed with these reviews, national and international 

agencies can approach donor agencies for funding. 

Agencies like the International Working Group on 

Dipterocarps could assist national and international 

institutions in identifying relevant projects. If several 

of these big projects are placed in one basket 

and handed to donor agencies, they could then select 

3

Introduction 

each project that is most needed for specific countries, 

and identify the specific groups that are in the 

best position to carry out the work. It is time 

dipterocarp forest scientists emulate the manner of 

astronomers. They are few in number, but collectively 

were able to put the billion dollar Hubble Space Telescope 

into space. 

3. Another possibility is to set up research centres 

exclusively devoted to research on dipterocarps. 

Interest has been expressed in setting up a Dry 

Dipterocarp Centre in Thailand and a Moist 

Dipterocarp Forest Centre in Kalimantan. 

For things to start moving in the right direction, it 

seems opportune to provide a general overview of what 

is already known about dipterocarps, and to identify the 

priority areas for further research, including what is 

needed to achieve the optimal use of dipterocarps. 

CIFOR has, therefore, undertaken to make this rapid 

overview of the family, from its systematics, ecology, 

management, end-uses, etc. This publication must be 

regarded as a first attempt to broadly cover several 

aspects of the dipterocarps. We take a broad look at the 

forests and the trees, and reexamine the way we manage 

them, and the opportunities awaiting their fullest 

development. Beyond that, we also touch on the research 

and development activities currently ongoing, and the 

future research and development needs. While the 

principal findings are stated, the document goes further 

to point out the important gaps in our knowledge and the 

kind of initiatives, both at international and national 

levels, that are needed. Finally, we hope this overview 

will form a precursor for a grander and more 

comprehensive coverage of this family of trees in the 

future. 

References 

Anonymous. 1991. Planting high quality timber trees in 

Peninsular Malaysia. Ministry of Primary Industries, 

Malaysia, Kuala Lumpur. 

Appanah, S. and Weinland, G. 1990. Will the management 

of hill dipterocarp forests, stand up? Journal of Tropical 

Forest Science 3: 140-158. 

Appanah, S. and Weinland, G. 1993. Planting quality 

timber trees in Peninsular Malaysia - a review. Malayan 

Forest Record no. 38. Forest Research Institute 

Malaysia, Kuala Lumpur. 221p. 

Ashton, P.S. 1982. Dipterocarpaceae. Flora Malesiana, 

Series I 92: 237-552. 

Barnard, R.C., 1954. A manual of Malayan silviculture 

for inland lowland forests. Part IV-Artificial 

regeneration. Research Pamphlet no. 14. Forest 

Research Institute Malaysia, Kepong. p.109-199 

Edwards, J.P. and Mead, J.P. 1930. Growth of Malayan 

forest trees, as shown by sample plot records, 1915- 

1928. Federated Malay States, Singapore. 151p. 

Evans, J. 1982. Plantation forestry in the tropics. Oxford 

University Press, Oxford. 472p. 

Food and Agriculture Organisation of the United Nations 

(FAO). 1981. Forestry Development Project Sarawak. 

Hill Forest Silviculture for Sarawak. FO:Mal/76/008. 

Working paper No. 4. FAO Rome. 


(FAO). 1989. Review of forest management systems 

of tropical Asia: case studies of natural forest 

management for timber production in India, Malaysia 

and the Philippines. FAO Forestry Paper no. 89. FAO, 

Rome. 

Jacobs, M.R. 1981. Eucalypts for planting. Forestry 

Series no. 11. FAO, Rome. 

Panayotou, T. and Ashton, P.S. 1992. Not by timber 

alone: economics and ecology for sustaining tropical 

forests. Island Press, Washington D.C. 282p. 

Poore, D. 1989. No timber without trees. Sustainability 

in the tropical forest. Earthscan Publications, London. 

Spears, J.S. 1983. Replenishing the world’s forests. 

Tropical reforestation: an achievable goal. 

Commonwealth Forestry Review 62: 201-217. 

Symington, C.F. 1943. Foresters’ manual of 

dipterocarps. Malayan Forest Record no. 16. Forest 

Department, Kuala Lumpur. 

Troup, R.S. 1921. Silviculture of Indian trees. Vol. 1. 

Forest Research Institute, Dehra Dun. 

Whitmore, T.C. 1975. Tropical rain forests of the Far 

East. Clarendon Press, Oxford. 

Wyatt-Smith, J. 1963. Manual of Malayan silviculture 

for inland forests. Vol. 1. Malayan Forest Record no. 

23. Forest Department, Kuala Lumpur. 

4

Biogeography and Evolutionary 

Systematics of Dipterocarpaceae 

G. Maury-Lechon and L. Curtet 

The history of Dipterocarpaceae botany, as understood 

in modern terms, started more than two centuries ago 

when Rumphius first mentioned the family in 1750. At 

that time dipterocarp forests were considered to be 

inexhaustible sources of wild products. The dipterocarps 

were thought to dominate extensively throughout 

southeast Asia. As soon as the high value of their products 

(camphor, resins, timber) was perceived funds were made 

available for botanists to conduct expeditions and 

laboratory research. A considerable amount of 

information has thereby been collected, and we now can 

recognise the valuable timber species in the forests and 

their natural distribution. The quality of market products 

thereby has become more uniform and predictable, thus 

favouring trade. At present, underestimated and 

unrestricted exploitation has encouraged excessive 

harvesting of dipterocarps and together with modern 

technologies and economics, has finally endangered the 

future of dipterocarp forests. 

As early as 1824 and 1868 de Candolle emphasised 

the importance of the number of stamens and their 

position in relation to petals to separate dipterocarp 

genera (Pentacme from Vateria, Petalandra from 

Hopea). These characters may affect the quantity of 

pollen produced and its availability for eventual 

pollinators. Similarly fruit and seed structures and shapes 

used in systematics also affect fruit-seed dispersal, 

germination and plant establishment. 

Present geographical distribution and the structures 

and functions of tropical plants are the results of past 

adaptations to environmental constraints. These features 

were produced in geological time under the influence 

of ancient climatic variation (Muller 1972, 1980). 

During the last decades, the intensification of human 

pressure on valuable trees has become the predominant 

factor of transformation for tropical forests (Maury- 

Lechon 1991). Excessive canopy openings provoke the 

Chapter 1 

rise of ambient temperature and desiccation. Faced with 

these new drastic conditions, past adaptations may no 

longer be suitable. If so, the definition of biological 

plasticity of well defined taxa according to their 

phylogenetic and ecological relations with the 

congeners will provide useful tools for forest managers 

(Maury-Lechon 1993). 

Such knowledge in systematics may have value in 

rehabilitation and sustainable management of forests. 

Understanding events such as pollination, fruit dispersal, 

seedling mycorrhization and survival, coupled with 

biogeographic distribution and evolutionary systematics 

may help to define lines of lesser phylogenetic resistance 

(Stebbins 1960, Maury-Lechon 1993). Such an approach 

provides the boundaries and physical limitations in which 

a species is able to survive and can be used to identify 

species most suitable for rehabilitation in the changing 

conditions that man has introduced into the environment. 

In this chapter, the present understanding of 

biogeography and evolutionary systematics of the family 

Dipterocarpaceae is reviewed and whenever possible 

there are attempts to link this knowledge to its use in the 

development sector. Finally, there are some notes on 

further research needs and expertise in the field. 

Presentation of the Family 

Dipterocarpaceae 

Taxonomy 

All Dipterocarpaceae species are arborescent and 

tropical (Fig. 1). The family type genus is the Asian 

Dipterocarpus Gaertn.f. Dipterocarps are trees with 

alternate entire leaves and pentamerous flowers. The 

family Dipterocarpaceae sensu stricto is homogeneous 

for only Asian plants while the Dipterocarpaceae sensu 

lato include three subfamilies: Dipterocarpoideae in 

Asia; Pakaraimoideae in South America; and

Biogeography and Evolutionary Systematics of Dipterocarpaceae 

Figure 1. Distribution of Dipterocarpaceae (adapted from Meher-Homji V.M. 1979). 

x 

x 

xxx 

x x x x 

Asian sub-division Dipterocarps 

Their presence in Seychelles and Andaman 

Marquesia 

African: 

Monotes 

Shorea robusta 

Fossils 

Doubtful fossils 

S. American Pakaraimaea 

New genus: Pseudomonotes tropenbosii 

Monotoideae in Africa and South America. The position 

of the African and South-American taxa relative to the 

Asian group varies with authors (Table 1). 

Consequently the family contains either 15, 16 or 

19 genera (Table 2) and 470 to 580 or more species 

(plus the newly found South American taxon, the 

monospecific genus called Pseudomonotes 

tropenbosii which has been attributed to the 

Monotoideae by its authors (Londoño et al. 1995, 

x 

x x 

x 

x x 

x x 

x x 

x 

Ã 

+ 

X X X 

S 

F 

D 

▲ 

 

Ã 

Morton 1995). During the past decade the numbers have 

reduced with the increase in collections and systematic 

expertise. However, uncertainties remain in Asia and 

Africa, underlining the necessity of an exhaustive and 

detailed review. 

Diversity of opinions also exists for generic 

divisions, especially with the genus Shorea and the group 

of genera Vatica and Cotylelobium. A synthetic 

classification is thus needed. It could be produced from 

6


Table 1. Recent content of Dipterocarpaceae family. 

Families Sub families Genera 

Maguire et al. 1977, Maguire and Ashton 1980 

Dipterocarpaceae Monotoideae Monotes 

Marquesia 

Pakaraimoideae Pakaraimaea 

Dipterocarpoideae see table 2 

Maury 1978, Maury-Lechon 1979a, b* 

Monotaceae Monotoideae Monotes 

Marquesia 

Pakaraimoideae Pakaraimaea 

Dipterocarpaceae see table 2 see table 2 

Kostermans 1978, 1985, 1989 

Monotaceae Monotes 

Marquesia 

Pakaraimaea 

Dipterocarpaceae see table 2 see table 2 

Londoño et al. 1995 

Monotoideae Pseudomonotes 

Monotes 

Marquesia 

* presented 1977, no formal status for taxonomic ranks, emphasis on 

greater affinities among taxa. 

the data now available, and the collaboration of still active 

workers, to define a solution acceptable to all in the 

laboratory, herbaria and field and the timber markets. 

First, however, more collections are needed of what 

appear to be key characters, in order to test their validity, 

particularly among species currently difficult to assign 

to supraspecific groupings. 

Botany 

Pakaraimaea are relatively small trees or sometimes 

even shrubs with alternate leaves (Table 3), conduplicate 

in aestivation, triangular stipules tomentulose outside 

and glabrous within, early fugaceous, glabrescent 

petioles, inflorescences axillary, racemi-paniculate, 

flowers 5-merous, petals shorter than sepals, neither 

connate at the base nor forming a cup and not winged at 

all, all 5 sepals become ampliate and none alate, calyx 

persistent, anthers deeply basi-versatile, connective 

conspicuously projected as an apical appendage, pollen 

grains tricolporate, exine 4-layered, ovary 5-locular 

(rarely 4), each loculus 2-ovulate (rarely 4), fruit with 5 

aliform short sepals, capsule at length dehiscent or 

splitting along dorsal line of carpel, wood, leaves and 

ovary devoid of resin or secretory canals, wood rays 

dominantly biseriate. No economic use is known 

(Maguire et al. 1977, Maguire and Steyermark 1981). 

Monotoideae are of three genera, Monotes, 

Pseudomonotes and Marquesia, and are trees or shrubs 

(Table 3). They have alternate leaves presenting an extrafloral 

nectary at the base of the midrib above (Verdcourt 

1989), small caducous stipules papyraceous, 

inflorescences in simple panicles, flowers 5-merous, 5 

sepals equally accrescent, petals longer than sepals and 

variously pubescent, calyx persistent, anthers basiversatile 

with apical connective-appendage scarcely to 

somewhat developed, pollen grains tricolporate, exine 

4-layered; ovary 1 to 3 locular (rarely 2, 4 or 5) with 

generally 2 ovules in each locule (rarely 4) except in 

Pseudomonotes (1 only), wood, ovary and commonly 

leaves without resin ducts, fruit sepals aliform and neither 

connate at the base nor forming a cupule, wood rays 

dominantly uniseriate. 

In Marquesia, trees are tall to medium-sized and 

buttressed, leaves evergreen and acuminate, nerves 

prominent with tertiary venation densely reticulate, 

indumentum of simple hairs and minute spherical glands 

on nerves and venation; flowers are small in terminal and 

axillary panicles; ovary 3-locular becoming 1-locular 

above parietal placentation, 6 ovules; fruit is ovoid with 

5 wings derived from the accrescent calyx, often 1seeded 

and apically 2, 3 or 4-dehiscent. 

Monotes are shrubs to medium-sized trees without 

buttresses, with leaves mostly rounded or retuse at apex, 

rarely acuminate, with more or less rounded extra-floral 

nectary at the base of the midrib above and sometimes 

additional ones in lower nerve-axils, with very varied 

indumentum and small spherical glands sparse or dense 

on both surfaces which often make the blades viscid, 

flowers in axillary small or compound panicles, ovary 

ovoid and hairy completely divided in 1, 2 or 3 

(sometimes 4: Maury 1970b, or 5: Verdcourt 1989) 

locules with 2 ovules in each locule, fruit subglobose 

presenting 5 equal minutely hairy wings derived from 

accrescent calyx, fruit normally 1-seeded and 

indehiscent (often 2, sometimes 3 or 4, rarely 5; in Maury 

1970b). 

Pseudomonotes trees are 25-30 m tall with a 70-80 

cm diameter, with poorly developed buttresses. This 

species forms entire alternate leaves conduplicate in 

7


Table 2. Recent (1994) genera, sections and sub-sections related to Dipterocarpaceae and authors. (Londoño et al. 

1995: new genus Pseudomonotes included, into Monotoideae sensu Maguire et al.) 

Ashton 

1964, 68, 77, 80, 82 

Meijer and Wood 

1964, 76 

Maury 1978 

Maury-Lechon 1979a, b 

s.: section; s.s.: sub-section; s.g.: sub-genus; subgr.: sub-group. 

Kostermans 

1978, 81a, b, c, 82a, b, 83, 

84, 85, 87, 88, 92 

1 Hopea 1 Hopea 1 Hopea 1 Hopea 

s.Hopea s.Hopea 

s.s.Hopea s.s.Hopea 

s.s.Pierrea s.s.Pierrea 

s.Dryobalanoides s.Dryobalanoides 

s.s.Dryobalanoides s.s.Dryobalanoides 

s.s.Sphaerocarpae s.s.Sphaerocarpae 

2 Neobalanocarpus not yet created 2 Neobalanocarpus 

2 Balanocarpus heimii 3 Balanocarpus 

3 Shorea 2 Shorea 3 Shorea 4 Shorea 

s.Shorea s.Shoreae Shorea including 

s.s.Shoreae s.g.Eushorea= Shorea s. Barbatae Pentacme genus 

s.s.Barbata (1992: p.60) 

s.Richetioides 4 Richetia 

s.s.Richetioides s.g.Richetia s.Richetioides 

s.s.Polyandrae s.Maximae 

s.Anthoshorea s.g.Anthoshorea 5 Anthoshorea 

s.g.Rubroshorea 6 Rubroshorea 

s.Mutica subgr.Parvifolia s.Muticae 

s.s.Mutica s.s.Muticae 

s.s.Auriculatae s.s.Auriculatae 

s.Ovalis subgr.Ovalis s.Ovalis 

s.Neohopea s.Rubellae 

s.Rubella s.Neohopeae 

s.Brachypterae s.Brachypterae 

s.s.Brachypterae subgr.Pauciflora s.s.Brachypterae 

s.s.Smithiana subgr.Smithiana s.s.Smithianeae 

s.Pachycarpae subgr.Pinanga s.Pachycarpae 

s.Doona 7 Doona 5 Doona 

s.Pentacme 8 Pentacme 

4 Parashorea 3 Parashorea 9 Parashorea 6 Parashorea 

5 Dryobalanops 4 Dryobalanops 10 Dryobalanops 7 Dryobalanops 

6 Dipterocarpus 5 Dipterocarpus 11 Dipterocarpus 8 Dipterocarpus 

7 Anisoptera 6 Anisoptera 12 Anisoptera 9 Anisoptera 

s.Anisoptera s.Pilosae s.Anisoptera 

s.Glabrae s.Glabrae s.Glabrae 

8 Upuna 7 Upuna 13 Upuna 10 Upuna 

9 Cotylelobium 8 Cotylelobium 14 Cotylelobium 

10 Vatica 9 Vatica 15 Sunaptea 11 Sunaptea (+Coty.) 

s.Sunaptea s.g.Synaptea 16 Vatica 12 Vatica 

s.Vatica s.g.Isauxis s.Vatica 

(s.Pachynocarpus 1964) s.g.Pachynocarpus s.Pachynocarpus 

11 Stemonoporus 17 Stemonoporus 13 Stemonoporus 

12 Vateria 18 Vateria 14 Vateria 

13 Vateriopsis 

14 Monotes 

15 Marquesia 

16 Pakaraimaea 

19 Vateriopsis 15 Vateriopsis 

8


Table 3. Affinities between Dipterocarpaceae sensu lato and other close angiosperm families. 

BOTANICAL CHARACTERS 

inflorescence paniculate (compound raceme) + + ± 

racemi-paniculate (+) + + + 

cyme appearance (+) 

bisexual flower + + + + + + + + + 

unisexual flower - - - + (-) (-) 

5-merous perianth + + + ± ± ± - ± 

bud-flower sepals imbricate + - + + - + + + 

valvate + + - - + - - - 

open-flower sepals imbricate + - + 

valvate + + - 

contorted corolla ± ± - ± - + + 

persistent sepals and calyx + + + + + 

fruit-sepals imbricate + + ± ± + + + 

valvate + + + + + + 

centrifugal stamens + + + + + + 

hypogynous stamens numerous + + + + + 

many + + + + 

2-celled anthers generally dehiscing longitudinally ± + + + 

subversatile anthers + + + + 

connectival appendage ± - - + 

pollen tricolporate - + + + + 

tricolpate + 

exine pollen 2-3 layers + 

4 layers - + + + 

ovary (2)-3-locular + + 

(2)-3-(5)-locular + 

4-5-locular + 

superior + + + 

semi-inferior (+) (+) 

generally 2 ovules/ cell + (+) + 

placentation axile + + + + 

aliform fruit sepals + + + - 

short-sepal fruit calyx + - 

possibility of peltate scales on the twig + 

seeds with scanty endosperm + + 

Dipterocarpoideae 

Dipterocarpaceae* 

Monotoideae 

sensu lato 

Pakaraimoideae 

Guttiferae 

Theaceae 

Tiliaceae 

Elaeocarpaceae 

Ochnaceae 

Sarcolaenaceae 

9


Table 3. (continued) Affinities between Dipterocarpaceae sensu lato and other close angiosperm families. 

BOTANICAL CHARACTERS 

leaves opposite - - - + 

alternate + + + + + + + + 

leaf venation prominent pinnate + + + + 

vertically transcurrent + + + + 

entirely transcurrent and + + + + + + 

presence of columns of sclerenchyme - 

indistinct leaf venation + + ± 

dentate leaves ± 

paired basal leaf nerves - - + 

stipule ± ± ± - - + + + + 

hypodermis (+papillose lower epiderms) - + 

1-2 layered hypodermis ± + + 

hair (various within a section) stellate ± - - 

tufted ± - + 

glandules ± - - 

complex indumentum + + + + 

geniculate petiole + + + + 

indumentum + complex anatomy petiole (Malvales type) + + + 

rays uniseriate - + - - 

biseriate - + 

multiseriate + - - - 

mixed uni/multiseriate + + 

presence of resins + + + - 

intercellular resin canals + - - + - 

mucilage canals in cortex and cells in the epidermis + + + - + 

elongate medullary mucilage cells + + - + + 

arrangement of bast fibres * into outwardly tapering wedges + + + + 

pith and primary cortex with indumentum + + + + 

anomocytic stomata + + + + 

complex petiolar vascular supply ± - - - 

chromosomes n=7 + + (+) (+) (+) - - 

n=11 + - - - - 

±: present and other possibilities; 

+: present; 

-: absent; 

Dipterocarpoideae 

Dipterocarpaceae* 

Monotoideae 

sensu lato 

Pakaraimoideae 

Guttiferae 

Theaceae 

Tiliaceae 

Elaeocarpaceae 

Ochnaceae 

(-) or (+): exceptions; 

* : adapted from Ashton 1982, Maury-Lechon 1979, and other 

works (see in text: Classification). 

Sarcolaenaceae 

10


vernation, oblong-ovate and chartaceous, with a vestigial 

gland on the midrib at the base of the blade, triangular, 

glabrous and caducous stipules. Inflorescences are 

axillary, subcymose, with bisexual 5-merous flowers, 

showing a glabrous calyx with 5 lobes which form a 

shallow cup at the base, a glabrous corolla with contorted 

petals, the petals longer than sepals, the stamens 

numerous, cyclic, hypogynous, the anthers basi-versatile, 

the connective broad and very expanded, continued into 

a triangular appendage one-fourth to one-half as long as 

the body of the anther, the pollen grains tricolporate, 

rarely tetracolporate, sometimes trisyncolpate, exine 

minutely reticulate to foveolate, columellate, 

tectateperforate, the ovary glabrous, 3-locular, one ovule 

per loculus. The fruit is a dry nut, glabrous with a woody 

pericarp, a persistent calyx with 5-winged accrescent 

sepals, thinly papyraceous, and 1 seed per fruit. As in 

African monotoids the wood anatomy of 

Pseudomonotes shows solitary vessels (occasionally in 

radial pairs), rays mainly uniseriate with infrequent 

biseriate portions, heterocellular rays, resinous contents 

present in vessel, rays and parenchyma cells, and presence 

of secretory cavities in the pith. No economic use is 

known but the local name (Nonuya Indians) means (in 

Spanish) ‘arbol de madera astillosa’, thus wood is 

probably used by native people. 

Pseudomonotes, Monotes and Marquesia may share 

solitary vessels or vessels in radial pairs, simple 

perforation plates, resinous content present in the 

vessels, rays and parenchyma cells, wood rays, presence 

of secretory cavities in the pith, lack of resin canals, 

single gland on the upper surface of the lamina at the 

base of the midrib, basi-versatile anthers and tricolporate 

pollen grains. Pseudomonotes differs from the Asian 

dipterocarps in the absence of fasciculate trichomes, 

multiserate rays, wood, ovary and leaves resin canals and 

tricolpate grains, and having one ovule per locule with 

nearly basal placentation. 

Dipterocarpoideae, the Asian dipterocarps are small 

or large, resinous, usually evergreen trees, often 

buttressed and usually developing scaly or fissured bark 

on large trees. Some or most parts present a tomentum, 

with alternate simple leaves, margin entire or sinuate, 

not crenate, penninerved, with a more or less geniculate 

petiole, stipules paired, large or small, persistent or 

fugaceous and leaving small to amplexical scars, 

inflorescence in panicles with racemose branches usually 

11 

with flowers secund, i.e. turned to one side, except in 

Upuna (cymose appearance perhaps due to reduction of 

a panicle of the Shorea type, and an even stronger 

reduction in some Stemonoporus and Dipterocarpus 

rotundifolius, whose flowers are solitary; in Kostermans 

1985). Extra-floral nectaries were recently found in 

many genera (Ashton, personal communication). In the 

5-merous flower, petals are longer than sepals and 

variously pubescent, calyx persistent with 0, 2, 3 or 5 

sepals enlarged into wing-like lobes in fruit, either free 

down to the base, forming a cup or a tube more or less 

enclosing the fruit, adnate to or free from it; when free 

to the base they are mostly imbricate. The basifixed erect 

anthers bear mainly 2 pollen sacs (rarely 4) on the 

connective terminated by a short or prominent 

appendage. Pollen grains are tricolpate with a 2 or 3layered 

exine. The ovary is superior or semi-inferior, 3 

(rarely 2) locular, each loculus contains 2 ovules. The 

fruit is loculicidally indehiscent, or at length splitting 

irregularly, or opening at staminal pore at germination, 

normally 1-seeded (sometimes 2, exceptionally up to 

12 or 18), with woody pericarp and persistent more or 

less aliform sepals. The stipules are often conspicuously 

large. Wood, ovary and leaves contain resin secretory 

canals. Wood rays are multiseriate (Maguire et al. 

1977). 

Ecology 

Monotes grows in deciduous formations, and most 

Marquesia species form dry deciduous forests or 

savanna woodlands. One species, M. excelsa, grows in 

Gabonese rain forest and resembles the Malaysian rain 

forest dipterocarps. Pseudomonotes is found in wet, 

evergreen rain forest and Pakaraimaea in evergreen 

associations. 

Pakaraimaea dipterocarpacea may dominate in dry 

seasonal evergreen forests on a variety of topographical 

situations, at altitudes of 450 to 600 m, on weakly 

ferralitic sandstones. The tallest tree recorded is 20 m 

with a diameter of 50 cm. Older or damaged trees freely 

coppice from the base as do some savanna dipterocarps 

in Asian seasonal regions. 

Pseudomonotes tropenbosii develops at 200-300 m, 

on clayey to sandy sediments, on summits of hills and 

along shoulders of slopes. These trees constitute the 

most ecologically important species in the rain forest a 

few kilometres south of Araracuara (Colombia).


Asian dipterocarps deeply imprint the forest ecology 

and economy of the places where they grow. They 

constitute prominent elements of the lowland rain forest 

(Whitmore 1988) and are also well represented in the 

understorey. As a family they dominate the emergent 

stratum. Most belong to the mature phase of primary 

forest, which contains most of the entire genetic stock 

(Jacobs 1988). All species can colonise secondary 

forests during the succession phases provided there is a 

seed source; seed dispersal is limited, except among 

water dispersed species. However, none seems presently 

confined to secondary formations. Certain dipterocarps 

of the seasonal regions dominate the fire-climax 

deciduous forests of northeast India and Indo-Burma. 

In Asia, dipterocarps occupy a large variety of habitats 

(Symington 1943, Wyatt-Smith 1963) from coastal to 

inland, riverine to swampy and to dry land, undulating to 

level terrain, ridges, slopes, valley bottoms, soils deeply 

weathered to shallow, well-drained to poorly drained, and 

rich to poor in nutrients. In Peninsular Malaysia the 

altitudinal zonation of their main habitat types ranges 

from 0-300 m (low-undulating dipterocarp forest), 300- 

750 m (hill dipterocarp forest), and 750-1200 m (upper 

dipterocarp forest). Zonation however, differs in Borneo 

and Sri Lanka. The freshwater swamps, especially in drier 

parts, are rich in species (Corner 1978, in Jacobs 1988) 

while true peat-swamp is relatively poor. The dipterocarp 

flora is also poor on limestone and riverine fringes. 

Asian dipterocarps are limited altitudinally 

(Symington 1943) by climatic conditions, and the 

conjunction of altitude and other natural barriers, such 

as large rivers and watersheds, have obstructed the 

distribution of species in Borneo. For example, the 

northwest and northeast of Kalimantan, Sarawak, Brunei 

and Sabah are much richer in species than the rest of 

Kalimantan. The everwet areas are also richer in species 

than the seasonal ones as shown in Sri Lanka by the 

concentration of species in the southwest quarter, or in 

the Thai-Malaysian transition belt, or from Java to the 

Lesser Sundas (Jacobs 1988). 

Distribution of Dipterocarps and 

Related Taxa 

The present distribution patterns of dipterocarps are 

thought to reflect routes of colonisation and past climatic 

conditions (Fig. 1). Living dipterocarps sensu lato are 

spread over the tropical belt of three continents of Asia, 

Africa and South America. They occupy several 

12 

phytogeographical zones that mainly conform to climatic 

and ecological factors. However, in southeast Asia, 

Wallace’s line where it runs east of the Philippines and 

between Borneo and Celebes, is a major phytogeographic 

boundary for dipterocarps. It cannot be explained in terms 

of climatic differences but requires the intervention of 

continental shelf drift. 

Phytogeographical Regions of Living Taxa 

The South American region (Fig. 1, Table 4) corresponds 

to Guyana, Venezuela and the part of Colombian Amazon 

which overlies the Guyana shield. 

The African region (Fig. 1, Table 4) includes a 

continental area and an insular part in Madagascar. The 

former is in two disjunct areas (Aubreville 1976): a) a 

narrow strip in the northern hemisphere from Mali on 

the west, to Sudan on the east, neither reaching the 

Atlantic nor the Indian Ocean; and b) in the southern 

hemisphere the Monotes-Marquesia area covers a semidry 

region between the two oceans, south of the 

Congolese rain forest, most of which is essentially 

central and does not reach the Atlantic or Indian Oceans. 

The Asian region (Fig. 1, Table 4) corresponds to the 

Indo-Malesian area, which concentrates a high number 

of genera and species in the equatorial forests. This area 

is limited northward by the Himalayan foothills, then 

approximately by the borders of Assam, Arunachal 

Pradesh (India), Burma, Laos and Vietnam, and 

penetrating into south China including Hainan Island. On 

the extreme southwest the large belt of Asian 

dipterocarps reaches the Seychelles (1 sp. Vateriopsis 

seychellarum), and covers India and Sri Lanka. Its eastern 

border corresponds to New Guinea. The Sundalands 

delimit the most southern part. No dipterocarp species 

is found in Australia. 

Five main phytogeographical regions are classically 

recognised within this distribution area: 1) Malesia: 

Peninsular Malaysia, Sumatra, Java, Lesser Sunda Islands, 

Borneo, the Philippines, Celebes, the Moluccas, New 

Guinea and the Bismarks. The northern frontier of 

Peninsular Malaysia delimits this part; 2) Mainland 

southeast Asia: Burma, Thailand, Cambodia, Laos, 

Vietnam and south China (Smitinand 1980, Smitinand et 

al. 1980, 1990); 3) south Asia: India, Andaman islands, 

Bangladesh, Nepal; 4) Sri-Lanka; and 5) Seychelles. In 

these Asian phytogeographical areas each dipterocarp 

group manifests a more or less distinctive pattern of 

variation at the species level (Ashton 1982).


Table 4. Phytogeographical regions and distribution of numbers of genera and species. 

Area Country 

Number of genera Number of species 

area country area country 

*: numbers in Ashton’s 1982 publication; 

°: Shaw’s numbers in Jacobs 1981; 

Philippines**: east of Wallace’s line only 3 genera and 13 species. 

Authors 

Malesia 10 465* *Ashton 1982 

Malaya 14 155* 168 Symington 1943 

Borneo 13 267* 276 " 

Sumatra ? 106* *Ashton 1982 

Philippines** 11 50* 52 " W. Wallace’s line 

Sulawesi 4 7 *Ashton 1982 

Moluccas 3 6 *Ashton 1982 

New Guinea 

Mainland 

3 15 " E. Wallace’s line 

Southeast Asia 8 76 Smitinand 1980 

Smitinand et al.1990 

Burma 6 33 " 

Thailand 8 66 " 

Laos 6 20 " 

Cambodia 6 28 " 

Vietnam 6 36 " 

China 5 24 Huang 1987 

11 Xu and Yu 1982 

Sri Lanka 7 44-45 Ashton 1977 

South Asia: 9 58 Kostermans1992 

India+Andamans 5 (6) 31 Tewary 1984 

North India 4 10 Jacobs 1981 

South India 5 14 " 

Andamans 2 8 " 

Seychelles 1 1 Parkinson 1932 

Africa 37* * Ashton 1982 

plus Madagascar 3 49° ° Shaw 1973 

Africa 2 48 Shaw 1973 

36 Ashton 1982 

≈30 Verdcourt 1989 

Madagascar 1 1 

South America 1 1 Maguire et al.1977 

Maguire and Ashton 1980 

NB: - Number of genera in China assumes China’s view of the 

China-India border, not accepted by India or internationally 

(Shorea probably does not occur in China). 

- Symington included undescribed entities, most of which were 

later absorbed in described entities by Ashton, which explains 

the difference between numbers. 

13


Present Distribution in the Phytogeographical 

Regions 

The South American region possesses now two 

monospecific genera related to Dipterocarpaceae sensu 

lato, and belonging to the two different non-Asian main 

groups: Pseudomonotes attributed by its authors to 

Monotoideae, and Pakaraimaea in Pakaraimoideae 

sensu Maguire et al. (1977). 

Pseudomonotes tropenbosii appears to be confined 

to a small area in the southwesternmost limit of the 

Guyana Highland and the superposed Roraima Formation 

sediments in Amazonian Colombia (Fig. 1: nov. gen.). In 

spite of being found near the distribution area of 

Pakaraimaea, Pseudomonotes has stronger affinities 

with the African species. Such affinities recall the remote 

Gondwanan connection between Africa and South 

America. 

Pakaraimaea dipterocarpacea contains two 

subspecies: P. dipterocarpacea ssp. dipterocarpacea in 

Imbaimadai savannas, Pakaraima Mountains, Guyana, and 

P. dipterocarpacea ssp. nitida in Gran Sabana and 

Guaiquinima, Venezuela (Maguire and Steyermark 

1981). The new genus Pseudomonotes from Colombia 

(Fig. 1) in most respects seems to be a Monotoideae 

(sensu Maguire and Ashton in Maguire et al. 1977), not 

a Pakaraimoideae (Londoño et al. 1995). 

African dipterocarps need a reassessment to reduce 

over-estimations in the Angolan flora. All Monotes 

(about 26 instead of 32 (Verdcourt 1989)) and 

Marquesia (3 or 4 species) grow in the southern 

hemisphere. Only Monotes kerstingii occurs in both 

hemispheres (Fig. 1). It occurs in the northern 

hemisphere as an isolated species in a narrow strip, and 

in the southern hemisphere in the main distribution area 

of the Monoitoideae. Some species exist through 

Katanga, Zambia and Mozambique up to the Indian Ocean. 

Only one species (Monotes madagascariensis) reaches 

south Madagascar. 

Marquesia may form monospecific open forests 

along the fringe of the Congolese rain forest, at the limit 

of Zaire, Angola and northern Zambia. 

The numbers of genera and species in Asia (Table 4: 

* indicates Ashton’s 1982 numbers) show much greater 

diversity compared to Africa and South America. As 

expected the higher numbers clearly occur in the everwet 

regions. The same trend exists from the Malesian region 

(10 or 14 genera, 465* species) and particularly from 

Borneo (13 genera, 267* species) and Peninsular 

14 

Malaysia (14 genera, 155* species), westwards to 

mainland southeast Asia (8 genera, 76 species) to Sri 

Lanka (7 or 9 genera, 44-58 species), India (5 or 6 genera, 

31 species) and the Seychelles (1 genus, 1 species). The 

same situation appears eastwards inside the Malesian 

region from Borneo to Peninsular Malaysia or to the 

Philippines (11 genera, 50* species) and from Malesia 

to China (5 genera, 11 or 24 species). The number of 

taxa strongly decrease on the east side of the Wallace’s 

line in the Philippines (3/11* genera, 13/50* species) 

and New Guinea (3* genera, 15* species). 

Particular needs for a new synthesis concern the 

Chinese taxa (Yunnan, South China, Hainan Island), using 

both the published literature (Wang et al. 1981, Tao and 

Tong 1982, Tao and Zhang 1983, Tao and Dunaiqiu 1984, 

Huang 1987, Zhu and Wang 1992), the on-going works 

(Yang Yong Kang 1994 personnal communication) and 

new collections to be done. Dipterocarps in New Guinea 

and the Philippines have been identified (Revilla 1976) 

but some biological aspects have to be specified. 

Cotylelobium Pierre and Pentacme, are the only 

Asian dipterocarps with a present disjunct distribution 

area (Table 5). Cotylelobium grows in Sri Lanka, 

mainland Southeast Asia and Sundaland, both under 

seasonal and aseasonal evergreen forests. Pentacme 

develops in mainland Southeast Asia and also in the 

Philippines and Papua-New Guinea. Only five genera 

develop east of Wallace’s line (Ashton 1979a) if 

Pentacme is not merged into Shorea genus: 

Dipterocarpus, Vatica (including Sunaptea), Hopea 

(section Hopeae) and Shorea (Anthoshorea and 

Brachypterae groups) and Pentacme. Apart from 

Pentacme, the other four genera presently exist in India 

and occur in Indian fossils (Table 5) as does the genus 

Anisoptera (presently extinct). Anthoshorea Heim 

extends from India to east of Wallace’s line. Section 

Shorea Ashton is mainly centered in southeast Asia but 

is well represented in Sri Lanka; it contains two fireresistant 

species in the Indian and Indo-Burmese dry 

dipterocarp forests. The other genera or sections have 

more restricted areas (Table 5). The south Asian endemic 

taxa are Vateria genus represented in south India and Sri 

Lanka, and Stemonoporus and Doona confined to Sri 

Lanka. In the southeast part Upuna is endemic in Borneo. 

Two genera, Vatica (sensu Kostermans) and Hopea, 

show the largest distribution from India to east of 

Wallace’s line. This is an important fact.


Table 5. Distribution of living and fossil dipterocarp genera or section. 

Taxa S.Am Afri Mada Seyc India Sri-L Chin Burm InCh Thai Mal Born Indo Phil N.Gu 

Pakaraimaea O 

Marquesia O 

Monotes O O 

Vateriopsis O 

Vateria O* O 

Stemonoporus O 

Doona O 

Balanocarpus K. O O 

Vatica Kosterm. O* O O O O O O O O* O O 

Dipterocarpus * (?) O* O O* O* O O O O* O 

Anisoptera * O O O O O O O O 

Anthoshorea O O O? O O O O O O O 

s.Shorea (*°) O*° O O? O*° O*° O O O O*° O 

s.Hopea O* O O O O O O O O O O 

s.Dryobalanoides O O O O O O O O 

Parashorea O O O O O O O O 

Pentacme *(?) O O O (O) O O 

Sunaptea O O O O O O O O O 

Cotylelobium O O O O O 

Neobalanocarpus (O) O 

Dryobalanops * * O O* O* 

s.Richetioides (O) O O O O 

s.Rubroshoreae (O) O O O O 

s.Brachypterae O O O O 

s.Pachycarpae O O 

s.Rubellae O O 

s.Neohopea O 

Upuna O 

S.Am: South America; Afri: Africa; Mada: Madagascar; Seyc: Seychelles; Sri-L: Sri-Lanka; Burm: Burma; Chin: China; InCh: Indo-China; 

Thai: Thailand; Mal: Peninsular Malaysia; Born: Borneo; Indo: Sumatra, Java and other Indonesian islands but Borneo; Phil: Philippines; 

N.Gu: New Guinea; (O): extreme geographic position (Langkawi island for Malaysia, extreme S-W Thailand); O: living species; *: fossils; 

O*: living species and fossils; s.Shorea(*°): both section Shorea, and Shorea sensu lato for fossils; O*°: both section Shorea and 

Shorea sensu lato when precise taxonomic level not specified, particularly for fossils. 

Potential Taxa for Differentiation 

The preceding facts suggest that Dipterocarpus, Vatica, 

Hopea section Hopeae and Shorea (sections 

Anthoshorea and Shorea) could be the main 

Dipterocarpoideae taxa from which new forms could 

arise by diversification during periods of isolation of 

Indian and East Asian lands. This is supported by certain 

highly variable species which, in a single species, may 

contain much of the whole set of variations of the other 

species in their own genus, or even that of different other 

genera for example, Shorea roxburghii and Vatica 

pauciflora (respectively S. talura and V. wallichii: in 

15 

Maury 1978, Maury-Lechon 1979 a, b, Maury-Lechon 

and Ponge 1979). The new taxa should probably 

correspond to groups of species such as Hopea s. 

Dryobalanoides and all the Shorea of the ‘red-meranti’ 

group in the Malesian area, and perhaps also 

Balanocarpus Kosterm. in the Indo-Sri Lankan part. The 

limited taxa Vateriopsis, Vateria in the west and Upuna 

in the east, are residual genus with limited potential for 

differentiation. Anisoptera, with the fossil and present 

distribution area, perhaps partly shares this lack of 

evolutionary potential and could be a regressing group. 

With more limited areas, Parashorea and Pentacme


from one part, and Dryobalanops from the other, could 

also enter this evolutionary-limited-potential group. 

Sunaptea, having been several times merged into 

Vatica, requires fossil and living distributions. The 

Sunaptea morphological and anatomical characters in 

embryos, fruit-seeds and seedlings could have had a much 

larger distribution. The same remarks concern 

Cotylelobium. Kostermans (1992) changed 

Cotylelobium scabriusculum (Thw.) Brandis into 

Sunaptea scabriuscula (Thw.) Brandis. Perhaps 

information on living and fossil dipterocarps from China, 

Indo-China and Burma could modify the present 

perception of their extension. It is also unknown whether 

the Vaticoxylon were of the Vaticae, Pachynocarpus or 

Sunaptea types. The absence of Cotylelobium among 

fossil forms results from a lack of detailed criteria in 

wood anatomy that prevents assessment of its presence. 

Close cooperative works between paleobotanists, 

wood anatomists of living forms and systematicists are 

needed to consolidate present conclusions on the mixed 

taxa of Vatica and Cotylelobium. Future work should 

particularly consider separately the Sunaptea, Vaticae 

and Pachynocarpus. The same treatment is necessary 

for the Shorea and Hopea sections. These remarks are 

especially pertinent for the new molecular approaches 

being rapidly developed, and which have been applied in 

a few instances to dipterocarps (e.g. Chase et al. 1993, 

Wickneswari 1993). Some of the diverse opinions, in 

all disciplines, are due to the studies being limited to a 

restricted number of species. It is therefore necessary 

to examine the whole set of species instead, with 

particular attention to intermediate ones such as Vatica 

heteroptera and V. umbonata group. The V. pauciflora 

(ex V. wallichii) case has already been mentioned above, 

together with Shorea roxburghii (ex S. talura). Maury- 

Lechon’s previous conclusions (Maury 1978, Maury- 

Lechon 1979a, b: Fig. 16, p.100) based on cotyledonary 

shapes and structures have been vindicated by 

Wickneswari’s results (1993: Fig. 1). These conclusions 

concern affinities between the Sunaptea group and 

Cotylelobium and their joint affinities with Upuna, as 

well as the connection of these three taxa with the 

closely related group of Anisoptera first, and then 

Dipterocarpus and Dryobalanops. Shorea bracteolata, 

the only Anthoshorea in Wickneswari’s study, has 

cotyledonary characters that are distinct from species 

such as S. roxburghii and S. resinosa (Maury 1978, 

Maury-Lechon 1979 a, b, Maury-Lechon and Ponge 

16 

1979). Consequently, the position of S. bracteolata 

reflects perhaps only partially the position of the whole 

group of species presently included within the 

Anthoshorea. Shorea resinosa and S. roxburghii 

cotyledonary shapes locate the Anthoshorea close to 

Doona and to Dryobalanops, Dipterocarpus and 

Cotylelobium. The present heterogeneity of 

Anthoshorea suggests the need for a re-examination both 

by DNA analysis and other approaches. 

The Dipterocarpus genus should also be examined 

for eventual correspondence between chemotaxonomic 

groups (Ourisson 1979) and biological characters such 

as seed sensitivity to desiccation and cold temperatures, 

seed and seedling resistance to pathogens by defensive 

secretions, and chemical type of root exudates for 

mycorrhizal fungi association. The statement that there 

is no relation between chemical groups and 

morphological features should be re-examined in 

considering flower characters, particularly stamen 

shapes, pollen and pollination, sexual and non-sexual 

reproduction, the fruit-seed-embryo-seedling sequence 

and the habitat. 

Phytogeographical Regions of Extinct 

Dipterocarps or Related Taxa 

No living or extinct monotoid (Tables 5 and 6) has been 

reported in Asia while diverse supposed dipterocarpoid 

fossils are described from Europe: Woburnia porosa 

wood from Bedfordshire Lower Cretaceous, U.K. 

(Stopes 1912, Kraüsel 1922, Schweitzer 1958, all in 

Aubréville 1976), flowers of Monotes oeningensis 

(Heer) Weyland from Upper Eocene in Hungary 

(Boureau 1957, Boureau and Tardieu-Blot 1955), and 

Tertiary fruits of west Germany, Switzerland and Austria 

(Gothan and Weyland 1964, in Aubréville 1976). Doubts 

were cast on these identifications (Bancroft 1933, Harris 

1956 and Hughes 1961 in Aubréville 1976, Gottwald and 

Parameswaran 1966, 1968). Other doubtful fruits of 

Monotes type have even been reported from New York 

(USA) and from the Alaska Eocene putative but unlikely 

tropical forest (Wolffe 1969, 1977). 

Boureau (in Boureau and Tardieu-Blot 1955) doubted 

this sequence but he remained convinced of the real 

presence of Monotes in the European Cretaceous and 

Tertiary. Huge distances would then separate the living 

Monotoideae from the extinct ones, and the Asiatic- 

Malesian Dipterocarpaceae from the European fossils, 

without any fossils in between, notably in North Africa.


Tertiary dipterocarp fossils have been reported from 

the African Miocene of Ethiopia (Beauchamp et al. 

1973, Lemoigne 1978, Laveine et al. 1987) and from 

the putative (but probably earlier) Plio-Pleistocene of 

Somalia (Chiarugi 1933). That is in a continent where 

not a single living dipterocarpoid has been found, and 

where present monotoids are living. Lemoigne (1978, 

p.123) specifies ‘c’est avec des bois de la famille des 

Dipterocarpaceae, notamment ceux du genre Monotes 

que notre echantillon parait avoir le plus d’affinites... 

Certes les affinites avec la famille des Lauraceae sont 

aussi remarquables’. In spite of its name 

Dipterocarpoxylon, this fossil is thus of a Monotoid 

type (not Dipterocarpoid). In this case, is the African 

rain forest species Marquesia excelsa derived from a 

common ancestor with Monotes and adapted to a more 

humid climate, or is it the only surviving species of some 

Dipterocarpoid ancestor which could have fossilised in 

Ethiopia and Somalia (and Egypt?)? A study of the pollen 

exine structure in Marquesia is needed to clarify this 

genus situation, as well as a critical re-examination of 

all dipterocarpoid fossils (doubtful or not). 

Numerous accepted fossils from Asia (Awasthi 1971) 

testify that the present great species richness of the Asian 

flora (Table 6) probably existed since the Miocene and 

persists through the Pliocene and Pleistocene, up to the 

Quaternary (Anisopteroxylon, Dipterocarpoxylon, 

Dryobalanoxylon, Hopenium, Shoreoxylon, 

Vaticoxylon, Vaterioxylon). 

These fossils demonstrate a reduction of dipterocarp 

distribution area both in Africa (Fig. 1, Tables 5 and 6) 

and Asia (extinction of Anisoptera and Dryobalanops 

in India, and of the latter in Indo-China), and total 

extinction in Europe and North America (doubtful 

fossils?). Could thus the Tertiary distribution area of 

dipterocarps sensu stricto include Africa and Asia (and 

Europe?)? 

Hypotheses on the Geographical Origin of 


If Monotoideae and Pakaraimaea are to be connected 

to Asian dipterocarps (by a single family or into different 

families), a common ancestor and its migration path have 

to be found. During the transition from the later 

Cretaceous to the very early Eocene the paleogeographic 

changes, in combination with other effects, could have 

produced the present geography. Thus dipterocarp 

ancestors should have been present when land 

17 

connections still existed between South America, Africa 

and India and between them and southeast Asia (and 

probably with the European and north American Laurasia 

block with its intermittent ‘Grande coupure’). This 

situation occurred (Figs. 2 (A, B), 3) in the Permo- 

Triassic period. Later, parts of the northeastern Gondwana 

land detached from the Gondwana shelf, crossed the 

Tethys and joined southeast Laurasia (just as India would 

do later). These changes would have happened during the 

Permo-Triassic, Jurassic and Cretaceous times according 

to recent works on Cathaysian floras (Kovino 1963, 

1966, 1968, all in Vozenin-Serra and Salard-Cheboldaeff 

1994, Lemoigne 1978, Jaeger et al. 1983, Vozenin-Serra 

1984, Vozenin-Serra and Taugourdeau-Lantz 1985, 

Laveine et al. 1987, Taugourdeau and Vozenin-Serra 

1987, Renous 1989, Scotese and McKerrow 1990 in 

Vozenin-Serra and Salard-Cheboldaeff 1994), and on the 

Tethys Sea (Dercourt et al. 1992). 

Croizat (1964, 1952 in Aubréville 1976) and Ashton 

(1969) expressed the view of a dipterocarp Gondwanan 

origin of the present distribution area and to a further 

migration towards Indo-Malesia. Aubréville (1976) 

considered that Dipterocarpaceae probably occupied two 

main areas before the Cretaceous general drift of 

Gondwanan shelves: one in Asia and one in the joined 

Africa-India-Seychelles-Sri Lanka complex. He believed 

that the origin was in Europe from where ancestors of 

monotoids would have migrated towards Africa and 

further from there to India. He suggested two Tertiary 

centres of dipterocarps: one Indo-Malesian from 

Laurasian origin; and one Africano-Indian from a 

Gondwanan origin, on both sides of the Tethys Sea. More 

recent studies (Renous 1989, Dercourt et al. 1992, 

Vozenin-Serra and Salard-Cheboldaeff 1994) identify 

direct land connections between southeast Asia and 

Laurasia lands. These authors consider a possible series 

of small blocks detached from the northeastern part of 

Gondwana, moving through the Tethys, and forming an 

archipelago (Fig. 3: Tazrim-Sino-Korean block (1), north 

China block (2), north Tibet block (3), Khorat-Kontum 

block: east Thailand and most parts of Laos, Cambodia 

and Vietnam (4), south Tibet block (5), Kashmir block 

(6), Iran-Afghanistan block (7), Turkey block (8), Spain 

block (9)) together with the different southeast Asian 

plates. This putative archipelago would have served as a 

relay, perhaps owing to volcanism which would open 

routes for floral migration. Before the rise of the 

Himalayas, floristic exchanges would also have been 

possible around the Tethys (Dercourt et al. 1992).


Figure 2. Continental drifts concerning the area of Dipterocarps from Primary to Tertiary (adapted from 

Renous 1989). 

E 

D 

C 

B 

A 

 

 

 

 

 

 

 

 

LAURASIA 

 

GONDWANA 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Miocene 

Late 

Cretaceous 

Middle & late 

Jurassic 

Triassic 

Permian 

I: India 

M: Madagascar 

S.Am: South America 

N.Am: North America 

Af: Africa 

Au: Australia 

An: Antarctica 

SrL: Sri Lanka 

MY - Million of years 

18


Table 6. Dipterocarp fossil distribution in time and space. 

Fossil genera Number of species Geological periods 

# : early period; 

* : mid period; 

& : upper period; 

+ : early+mid-period; 

** : mid+upper period. 

The two successive hypotheses, of Aubréville’s 

double-cradle areas, and of the Vozenin-Serra and Salard- 

Cheboldaeff’s archipelago connection anterior to India’s 

contact with Laurasia, could support a possible very 

remote common ancestor of American, African and Asian 

total country II III IV 

Monotes oeningensis flower? 1 1 Hungary * 

Monotoid fruit remants? 1 Germany, Austria 

Switzerland 

* 

Cret. Eoc. Olig. Mio. Plio. Plei. 

Monotoids?? : 

3 

& 

Calicites alatus, C.obovatus 

2 2 New-York USA & 

Monotoid fossil? 

1 1 Alaska 

& 

Dipterocarpoxylon? Woburnia 1 1 Great Britain & 

Dipterocarpophyllum? 1 1 Egypt & 

Dipterocarpophyllum 1 1 Nepal, N. India * * 

Dipterocarpus-type pollen 1 1 Nepal, N. India * 

" 1 1 Vietnam III 

Dipterocarpoxylon? Monotoid 

(Lemoigne 1978) 

30 1 Ethiopia # 

Dipterocarpoxylon? Monotoid? 2 Somalia & # 

Dipterocarpoxylon 11 North India + # 

" 2 Burma III 

" 2 Vietnam III 

" 3 Sumatra III IV 

" 7 Java III 

Anisopteroxylon 7 5 N. and NW India ** # 

" 2 India ** # 

Vaticoxylon 2 1 Sumatra IV 

" 1 Java * 

Vaterioxylon 2 2 North India & # 

Shoreoxylon 23 13 3 India-Assam ** # 

" 1 Northwest India ** # 

" 1 North India III 

" 3 South India ** # 

" 1 Assam-Cambodia * 

" 1 Burma III 

" 1 Thailand # # 

" 7 Sumatra * * IV 

Pentacmoxylon?? 1 1 India III? 

Hopenium 4 2 North India * 

2 South India * 

Dryobalanoxylon 13 1 Cambodia IV 

" 1 South Vietnam # # 

" 

1 Borneo 

* 

Dryobalanops pollen 

1 Borneo * 

Dryobalanoxylon 6 Sumatra * * 

" 5 Java ** # 

19 

dipterocarps. They could explain the particularities of 

the Indian-Seychelles-Sri Lanka region, the existence of 

Upuna in Borneo and the post Cretaceous explosion of 

the dipterocarp family in the more humid and warm 

conditions found in southeast Asia.


Figure 3. Paleogeographical reconstitution of southeast Asian Permo-Triassic in conformity with paleobotanic 

data (from Vozenin-Serra and Salard-Cheboldaeff 1994). 

Be the origin in Europe or in Africa, both cases would 

have favoured dispersal and colonisation by the small, 

light, large-winged fruits over great distances and 

probably under drier conditions than those of the present 

rain forests. This could explain why certain present taxa 

suggest an ancestral form with these fruit characters and 

again brings forward the hypothesis (Maury et al. 1975a, 

Maury 1978, 1979, Maury-Lechon 1979b) of an origin 

in open (perhaps semi-dry) environment. Forms with 

wingless fruits would have had the only possibility to 

concentrate their evolutionary potential into the 

protective structures around the seed (e.g., Vateria, 

Vateriopsis, Stemonoporus, certain Vatica and other 

9 

1. Sinkiang block (Tarim) + Sino-Korean block (North China). 

2. Yangtse block (North China). 

3. North Tibet block (North Xizang + North-West Xunnan + Shan Plateau). 

4. Khorat-Kontum block (East Thailand, most of Laos, Cambodia and Vietnam, South of Song Ba suture). 

5. South Tibet block (Lhassa plate + lands located South of Yalu Tsangpo). 

6. Kashmir. 

7. Iran and Afghanistan (Helmand block). 

8. Turkey. 

9. Spain. 

8 

6 

7 

5 

1 

3 

2 

4 

20 

species with large wingless fruits), these structures could 

have favoured water dispersal. In the taxa possessing 

winged fruits the evolutionary potential might have 

remained available for new opportunities. Upuna and 

Monotes kerstingii are perhaps parallel in this respect, 

as they suggest the hypothesis that they represent similar 

situations developed in or near Laurasia (Upuna) and in 

Gondwana (Monotes kerstingii) during long periods of 

supposed separation. 

Past Continental Changes and Floral Evolution 

Past flora also underline a connection between large 

floristic-climatic changes and the main known collisions


or ruptures of continental drifts: a) by lack of marine 

influence the Permian continental block (Fig. 2 (A), 3) 

would have been drier than the previous split continents 

of the Carboniferous with its luxurious Cryptogamic 

flora; b) the split of Gondwanaland during the Secondary 

would have permitted marine humidity to enter the 

fragmented lands (Fig. 2 (B, C) and probably the first 

Angiosperm ancestral forms to originate; c) the supposed 

late Cretaceous - early Eocene (Renous 1989, Dercourt 

et al. 1992) connection of India with Eurasia (Fig. 2 (D)), 

between -65 and -40 million years, and later that of Africa 

(Fig. 2 (E)), would have created new dry and humid zones 

and corresponded to the differentiation of Angiosperms 

(and dipterocarp ancestors?). 

According to these reconstituted changes, the flora 

of past continents from Permian (Primary) to Miocene 

(Tertiary) times had a very ancient common history (land 

and climate). Later on the future southern part of the 

Eurasian southeast zone first separated from the rest of 

Eurasia (Fig. 2 (B): Triassic: hatched area, Fig. 3) and 

then (the upper Jurassic) connected with it (Fig. 2 (C, D, 

E)). Wallace’s line corresponds to the separation between 

lands of different origins: the two Gondwanan shelves, 

the Indian on the west and the Australian on the east (Fig. 

2 (C)). 

For long geological periods (lower to extreme upper 

Cretaceous period, Secondary) the Indian-Seychelles- 

Sri Lanka part of the Gondwana shelf remained under an 

insular situation. For similar long periods the present 

regions of mainland southeast Asia, China and Malesia 

pro-parte remained separated from the Indian island, but 

perhaps intermittently connected to Eurasia. The Indian 

collision with Eurasia produced huge changes (land, 

climate, flora) as well as possibilities (or difficulties) 

of colonisation and species evolution for both types of 

flora (the insular-Indian flora and the continental-Asian 

flora) in the new territories. 

Paths of Possible Flora Migrations 

Four main land connections are thus suggested for 

eventual migrations of the ancestors of the dipterocarps, 

at different periods after the Gondwana split: a) India- 

Sri Lanka-Madagascar-Africa-America-Eurasia (Fig. 2 

(C)); b) the putative eastern archipelago northeast of 

Gondwana to Eurasia (Fig. 3); c) later, India-Sri Lanka- 

Eurasia (Fig. 2 (C, D)); and d) finally northeast Africa - 

Southeast Eurasia (Fig. 2 (E)). Because of the distances, 

land dimensions and climate history, the first connection 

21 

could have favoured the success and survival of species 

with small winged fruits, the second could have aided 

species with water dispersal, while the third could have 

permitted the persistence and establishment of more 

diverse biological types. Perhaps excessively dry 

climates did not favour dipterocarp migrations in the 

fourth case. 

These geological events bring light to the present 

distribution over three continents and the paucity east 

of Wallace’s line. They explain certain endemic aspects 

such as the Monotes kerstingii disjuncted area in Africa 

(survival at the periphery of the rain forest newly 

established in the previously drier area of Monotes). 

They could justify Upuna in Borneo, and localisation of 

Vateriopsis, Vateria, Stemonoporus and Doona in the 

Indian island zone. These events underline the existence 

of a very long past of successive modifications, and help 

to explain the real difficulty in finding primitive features 

in present flora. If characters evolved independently 

from each other, a single present taxon might have 

retained some primitive aspects and modified others; 

these latter preventing consideration as an ancestral form. 

Endemicity of Dipterocarps Sensu Lato 

As expected, the higher endemicity is located at the 

extremes of the geographical area of distribution. It is 

due to monospecific genera westward in south America 

(100%: 2 sp.), Madagascar (100%: 1 sp.) and Seychelles 

(100%: 1 sp.). Endemicity is of different intensity (Table 

7) eastward in Sri Lanka (98%: 43/44 spp.), south India 

(85%: 11/13 spp.) and in New Guinea (73%: 11/15 spp.), 

and with a much lower proportion in Borneo (58 to 55%: 

158 to 155/267 spp. of which 1 is a monospecific 

endemic genus), north Peninsular Malaysia (49%: 23/ 

47 spp.) and the Philippines (47%: 21/45 spp.) and north 

India (40%: 4/13 spp.). A certain endemicity also exists 

in the other Malesian areas but the values rapidly 

decrease: Celebes (29%), Java (20%), Peninsular 

Malaysia (17-18%), Moluccas (16%). Peninsular 

Malaysia, Sumatra and Borneo only separated 10,000 

years B.P. and, if taken as one biogeographic region, its 

endemicity is 293/345 species or 85% when the 

boundary is determined by the Kangar/Pattani line, 303/ 

345 species or 87% when the boundary is the Isthmus 

of Kra. 

Endemicity is very reduced on a country to country 

basis (Vietnam 9%, Laos 5%), or absent in the mainland 

southeast Asian phytogeographical area (Burma, Thailand, 

Cambodia; however, for Indo-Burma as one 

biogeographic region it is high), and totally absent from


Table 7. Distribution and importance of endemicity in Dipterocarpaceae family. 

Geographical areas 

number of 

species 

* at least total of 16-24 sp. in China (Huang 1987, Yang Y.K. Personal communication), perhaps about 38%. 

Lesser Sundas and Lombok islands. In mainland China 

and Hainan Island data are too uncertain to draw any 

conclusion. More studies are needed in mainland 

southeast Asia and China. 

Asian Dipterocarp Vicariance 

Asian dipterocarps also present groups of twin vicariant 

species with similar function in different areas. 

Vicarious species (Ashton 1979a) have been noted in 

genera Dipterocarpus, Anthoshorea and Hopea (Hopea 

section) between Sri Lanka and either south India or 

Malesia, and between south India and Malesia or 

Indonesia. 

All these features correlate with the history of the 

continents and the combined action of island isolation 

and two other major forces: a) the drier climates in the 

Ashton 1982 Jacobs 1981 

% of 

endemicity 

total number 

of species 

number of 

endemics 

% of 

endemicity 

Seychelles 1 1 100 

Sri Lanka 44 43 98 

South India 13 11 85 

North India 13 4 40 

Andamans 8 1 12 

Burma 32 0 0 

China mainland 5* 3 60 ? 

Hainan 1 0 0 

Vietnam 35 3 9 

Laos 19 1 5 

Cambodia 27 0 0 

Thailand 63 0 0 

Peninsular Malaysia 155 18 156 26 17 

Sumatra 106 10 95 10 10 

Java 10 20 10 2 20 

Lesser Sundas 3 0 0 

Lombok 3 0 0 

Borneo 267 58 267 158 55 

Philippines 50 47 45 21 47 

Celebes 7 29 7 2 29 

Moluccas 6 16 6 1 16 

New Guinea 15 73 15 11 73 

North Peninsular Malaysia 

(Malayan-Burmese floristics) 

47 23 49 

22 

western lands which are accentuated in South America, 

Africa, Madagascar and lower for the Seychelles, south 

India, north India, and part of Sri Lanka; and b) the 

maintained humidity in the eastern extreme of the 

Eurasian lands. The Eurasian lands could have permitted 

the development and spread of winged, light-fruited 

dipterocarps for long periods of time. This would allow 

migrations and floristic exchanges first in the Eurasian 

and later into India on the west, as well as into recently 

emerged parts of Sunda and Malaysia regions, and the 

newly arrived parts of New Guinea. 

The Sunda region has been (and is still) submitted 

for long periods to an intense geological activity which 

could have interfered with the great diversification of 

dipterocarps within the wet Malesian region and more 

particularly in Borneo (Maury 1978).


Geographical Patterns in Biological 

Characters 

There is a relation between shapes and structures and 

the biological processes they permit. It is thus necessary 

to try to understand how the morphological or biological 

characters (which constitute the base of the taxonomic 

divisions and systematic affinities) are related to the 

survival of plants in a given habitat, particularly under 

eventual modifications. During the past geological time, 

climatic and/or geographic variations predominated, 

while presently the transformations by human beings 

predominate. This type of information is not yet available 

for the American and African putative dipterocarp taxa. 

Most importantly, some characters (which are 

essential in establishing phylogenies and classifications) 

are ancestral and do indeed constrain the ecological 

range of species; but others are plastic, derived and 

adaptive to ecological circumstances. This distinction 

still requires classification among dipterocarps. For 

Ashton, the greater the number of correlated/independent 

character states, the more ancient, conservative and 

phylogenetically important they are, thus this point 

should be a major basis for assessment of phylogenetic 

generic delimitation. However, Kostermans strongly 

disagreed with this approach when presenting his case 

for Sunaptea. 

Biological Groups 

All the African taxa, except one, fit into monospecific 

formations of savanna woodland or dry deciduous 

forests, under seasonal climates. Marquesia excelsa, a 

residual species of the Gabonese rain forest is close to 

the other savanna species of the genus, and the new South 

American genus Pseudomonotes which appears closely 

related to Marquesia, present the opposite situation. 

Pakaraimaea is abundant but not monospecific and 

could multiply both through coppicing and sexual 

reproduction. However in the laboratory the germinative 

potential of seeds was low and the survival of young 

seedlings in USA and France nearly impossible (Maguire 

and Steyermark 1981, Maguire and Maury unpublished). 

Most Asian dipterocarps remain in evergreen forests 

(some in seasonal regions, most in aseasonal areas). A 

few species of Dipterocarpus and Shorea live in fireclimax 

savanna woodlands, though closely allied to rain 

forest species. 

23 

There is a sharp discontinuity in Asian dipterocarps 

in ecological and geographical ranges of the family 

Dipterocarpaceae between the evergreen forest and the 

fire climax dry dipterocarp woodlands (Ashton 1979a). 

The species of the latter group present characters which 

are unusual within the family: thick, ruggedly fissured 

bark, some seed dormancy, cryptocotylar germination, 

easily coppicing, seedlings with prominent taproots as a 

result of frequent burning. 

The dipterocarp flowering (and consequent fruiting) 

phenology also changes: in seasonal areas species flower 

annually with varied intensities each year; in aseasonal 

regions sporadic flowerings occur each year in riparian 

species for example, but large gregarious flowerings 

happen at intervals of 3 or 4 years (Sri Lanka) or 5 or 10 

years (aseasonal Malesia) (Ashton 1988, 1989). The 

gregarious flowerings are synchronous within 

populations and occur over several months for the whole 

family (Ashton 1969, Chan 1977, 1980, 1981, Ng 1977). 

Certain understorey Stemonoporus and Shorea however, 

do not follow this timing. The climatic boundaries closely 

coincide with the boundaries of the regions of exceptional 

dipterocarp diversity. The abundance of species and 

gregarious flowerings both occur in aseasonal west 

Malesia. 

Detailed information on these aspects is needed for 

the American and African taxa. 

Morphological Trends Related to Biological 

Patterns 

Deciduousness is mainly connected to seasonal areas 

while evergreen trees are more frequent in the aseasonal 

zones (Ashton 1979a). Degree of hairiness decreases 

from seasonal to aseasonal; the extreme expression of 

glabrousness is in understorey taxa such as certain Vatica 

and many Hopea. The tomentum disappears first from 

leaves, then from twigs, followed by the young shoots 

and finally the inflorescence and floral parts. Similar 

trends exist in Africa with Marquesia species from the 

open savanna forests and the only species from the rain 

forest, Marquesia excelsa. 

Flower and fruit characters have strongly influenced 

dipterocarp classification. Flower size seems constant 

within genera and Shorea sensu lato subdivisions, except 

in Dipterocarpus. The larger flowered taxa have their 

crowns in or above the canopy. This is the case for 

Vateria, Vateriopsis, Dipterocarpus, Anisoptera, 

Parashorea, Shorea section Shorea (Ashton’s sub-


section Shorea), Pentacme, Doona and Anthoshorea 

which form a full complement of supraspecific taxa 

exclusively confined to canopy in the Asian seasonal 

tropics (Ashton 1979a). But many other emergent 

Shorea, especially the Richetioides group, have small 

flowers, whereas Vatica and Stemonoporus (many of 

which flower beneath the canopy) have large flowers (as 

big as Parashorea or Anthoshorea). 

The anther and stamen sizes broadly follow the same 

trend: a) in the seasonal area the anthers are large, 

elongate and bright yellow for Vateria, Vateriopsis, 

Dipterocarpus, Parashorea, and Pentacme; and b) the 

same type of anthers also characterise certain taxa 

confined to aseasonal regions: Dryobalanops, 

Cotylelobium, Neobalanocarpus, Stemonoporus, 

Ashton’s Shorea section Rubellae, and Doona. In the 

other taxa, such as Anthoshorea and Ovalis, anthers are 

smaller, white and subglobose to ellipsoid; Richetioides 

presents the smaller ones. The African taxa seem to 

possess numerous stamens of medium size (drawings 

of Verdcourt 1989). 

The number of stamens (Ashton 1979a) is often 15: 

Vateria, Vateriopsis, most Dipterocarpus species, 

Anisoptera section Anisoptera, Dryobalanops, Shorea 

sections Shorea and Rubellae, 6 species of Anthoshorea 

all in seasonal sites, Ovalis, 1 species of Brachypterae, 

1 species of Richetioides, and 3 species of Hopea (2 of 

which are in seasonal sites). Ten species have less than 

15 stamens: 10 stamens in 6 species of Hopea and 3 

species of Richetioides, 5 stamens in 2 species of 

Stemonoporus and 1 species of Vatica. 

Large flowers produce large pollen grains (Muller 

1979). Flower and pollen dimensions will interfere with 

potential pollinators. Clear relations have been 

demonstrated between ovary shapes and sizes within 

Shorea sensu lato subgroups and pollinator size or 

taxonomical group (Chan and Appanah 1980, Appanah 

and Chan 1981, Appanah 1990). Bees pollinate large 

yellow elongate anthers while thrips pollinate small, 

white anthers. Bees prevail in seasonal tropics and Sri 

Lanka. Pollination changes during geological to present 

times probably explain much of the present aspect of 

dipterocarps. This is an important point to consider when 

planting trees outside their original areas. Forest 

degradation may result in the absence of tree 

reproduction by extinction of pollinators. 

The biggest fruits are in taxa with large flowers, and 

more frequently in species producing wingless-fruits 

24 

than in species with winged fruits. There is also a relation 

between large dimensions and the development of a 

protective thickening of pericarp and/or calyx base to 

prevent dehydration of the embryo and sometimes 

permit floating of the fruit (Maury 1978, Maury-Lechon 

1979b, Maury-Lechon and Ponge 1979). Thickened sepal 

bases are a defining character of Shorea sensu Ashton, 

but do not occur in Anisoptera, Upuna, Cotylelobium 

or Sunaptea. Pericarp thickenings characterise 

particularly the Pachynocarpus and Vatica groups of 

genus Vatica and genera Stemonoporus and Vateria. The 

thickening is of different type in Monotoideae and the 

case of Dipterocarpus remains apart because of the 

variously thickened calyx ornamentations (tubercules, 

simple or folded wings). The protective thickenings 

mainly develop in the group of taxa forming 15 elongate, 

large, yellow anthers. Large fruits are produced in smaller 

numbers, and they represent an investment which lowers 

risks in weakly lit places. The increased size of seedembryo 

probably demonstrates a trial for better survival 

in unpredictable habitats with irregular supply of light 

and nutrients (and water) during the germination period 

of non-dormant seeds. However, fewer fruits are 

produced, so that investment is in fewer high-cost seeds 

bearing other risks of probably lower intensity. Animal 

predation is mainly by insects and seeds do germinate 

and develop normal seedlings in spite of insect larvae 

which continue their development within the fleshy 

cotyledonary limbs; human predation is more drastic and 

mainly corresponds to traditional and industrial oil 

extraction. 

The 5-winged fruits of Pakaraimaea, Monotes and 

Marquesia clearly disperse in open and windy habitats, 

as probably do that of Pseudomonotes (detailed 

information not yet available). In these taxa pollen and 

nuts show evident adaptations to the dry conditions of 

their seasonal climates: thick layers and protected 

apertures, while the thin coriaceous pericarp of the ripe 

fruit of Marquesia excelsa is an exception (Maury et 

al. 1975a, b, Maury 1978, Maury-Lechon 1979a, b, 

Maury-Lechon and Ponge 1979). In Asia the wingedlight 

fruits of Sunaptea and Cotylelobium, of certain 

species of Hopea, Shorea (Ashton sensu lato) and 

Upuna present thin pericarps, even in seasonal regions. 

Asian taxa have developed winged fruits in seasonal 

and aseasonal regions. In closed forests these fruit wings 

have limited possibilities for dispersal. However, over 

the canopy and at forest borders, storms and very strong 

winds at the beginning of the rainy season may transport


them for several hundred metres and sometimes about 

one kilometre. In the open dry deciduous forest of the 

seasonal Asian regions the dispersal of dipterocarp 

winged fruits becomes much more efficient. However, 

occasional dispersal over long distances has probably 

succeeded, judging from the present high diversity of 

Asian dipterocarp habitats in aseasonal regions. 

Classification 

The full set of taxonomic and systematic works available 

has been reported and discussed in the more recent flora 

books, monographs, theses and other publications of 

Ashton (1962, 1963, 1964, 1967, 1969, 1972, 1977, 

1979b, c, d, 1980, 1982), Bancroft (1933, 1934, 1935a, 

b, c, d, e, 1936a, b, 1939), Bisset et al. (1966, 1967, 

1971), Damstra (1986), Diaz and Ourisson (1966), Diaz 

et al. (1966), Gottwald and Parameswaran (1964, 1966, 

1968), Jacobs (1981), Jong and Lethbridge (1967), Jong 

and Kaur (1979), Jong (1976, 1978, 1979, 1980, 1982), 

Kochummen (1962), Kochummen and Whitmore 

(1973), Kostermans (1978, 1980, 1981a, b, c, 1982a, 

b, 1983, 1984, 1985, 1987, 1988, 1989, 1992), Maguire 

(1979), Maguire et al. (1977), Maguire and Ashton 

(1980), Maguire and Steyermark (1981), Maury et al. 

(1975a,b), Maury 1978, Maury-Lechon (1979a, b, 1982 

in Ashton: p. 265-266 ripe embryo, germinating 

seedlings and 268 palynology, 1985 in FAO: palynology, 

Maury-Lechon and Ponge (1979), Meijer (1963, 1968, 

1972, 1973, 1974a, b, 1979), Meijer and Wood (1964, 

1976), Ourisson (1979), Parameswaran (1979a, b), 

Parameswaran and Gottwald (1979), Parameswaran and 

Zamuco (1979, 1985 in FAO), Rojo (1976, 1977, 1979, 

1987), Rojo et al. (1992), Sidiyasa et al. (1986, 1989a, 

b), Smitinand (1958a, b, 1979, 1980), Smitinand and 

Santisuk (1981), Smitinand et al. (1980, 1990), Sukwong 

(1981), Sukwong et al. (1975), Tao and Tong (1982), 

Tao and Zhang (1983), Tao and Dunaiqiu (1984), Tewary 

(1984), Tewary and Sarkar (1987a, b), Verdcourt (1989), 

Vidal (1960, 62, 79), Whitmore (1962, 1963, 1976, 

1979, 1988), Wildeman (1927), and Wildeman and 

Staner (1933). 

Affinities of the Dipterocarpaceae Family 

Hutchinson (1959, 1969 in Maury 1978) put the family 

in the order Ochnales and later suggested a phylogenetic 

location of the order at the end of a series whose 

progressive steps were ordered as follows: Magnoliales, 

Dilleniales, Bixales, Theales and Ochnales. 

25 

Cronquist (1968) moved Dipterocarpaceae into the 

order Theales and Takhtajan (1969) regrouped Ochnales, 

Theales and Guttiferales in Theales and placed 

Dipterocarpaceae under it near Lophiraceae and 

Ancistrocladaceae. Dalgren (1975) placed the 

dipterocarp family in the order Malvales under 

superorder Dilleniflorae. 

After the description and detailed study of 

Pakaraimaea dipterocarpacea and its inclusion in 

Dipterocarpaceae, Ashton removed the family from 

Guttiferales and considered the imbricate calyx of 

Dipterocarpaceae and Sarcolaenaceae (Maguire et al. 

1977) as an isolated and derived character among 

otherwise Malvalian features, and hence a justification 

for inclusion in Theales. From the study of Pakaraimaea, 

a closer affinity was underlined between the 

Dipterocarpaceae and Sarcolaenaceae families. Indeed, 

the affinities between these two families within the 

Malvales could be regarded as greater than with the 

Tiliaceae (Maguire and Ashton in Maguire et al. 1977: 

p. 359-361). The Tiliaceae relation is stronger with the 

African taxa while for the American Pakaraimaea the 

strong affinities are to be found with the Sarcolaenaceae 

(De Zeeuw, in Maguire et al. 1977). The Monotoideae 

could be a link between Tiliaceae and Dipterocarpaceae. 

Previously Blume (1825), who first described the 

family Dipterocarpaceae, Pierre (1889-1891 in Maury 

1978), Heim (1892) and Hallier (1912 in Ashton 1982) 

mentioned the close affinity of Monotes and Tiliaceae. 

Heim and Hallier had concluded that Monotes did not 

belong to Dipterocarpaceae (Kostermans 1985). The 

tilioid structure of the pollen exine (Maury et al. 1975a, 

b) in Asian and African taxa again called attention to these 

possible affinities with Tiliaceae. 

Kostermans (1978) excluded Monotoideae and 

Pakaraimoideae from Asian Dipterocarpaceae and 

formally described the family Monotaceae (suggested 

by Maury 1978, Maury-Lechon 1979a, b, Maury-Lechon 

and Ponge 1979) and recognised close relations between 

Monotaceae and Tiliaceae. By the structure of the pollen 

exine the Monotoideae strongly differ from Asian 

Dipterocarpaceae in spite of the similarities of the tilioid 

aspect of the exine surface. 

Later Kostermans (1985) concluded that ‘it is very 

difficult to make a decision of alliance of the real Asiatic 

Dipterocarpaceae. They are not much allied to Guttiferae 

or Ternstroemiaceae’ (=Theaceae) ‘and apparently 

represent an ancient family, in which nowadays links with


other families have disappeared.’ Thus we can just ‘resign 

ourselves to leave the Dipterocarpaceae s.s. near the 

Guttiferae and Ternstroemiaceae (=Theaceae). With the 

Malvales or Tiliales they have very little or nothing in 

common.’ 

However, with recent help of molecular techniques 

on two species of Dipterocarpoideae (Shorea stipularis, 

Anthoshorea section, and S. zeylanica = Doona 

zeylanica) (Chase et al. 1993), Dipterocarpaceae are 

placed as an outlier in the order of Malvales. Its relations 

with Malvales are with Bombacaceae (Bombax), 

Tiliaceae (Tilia), Sterculiaceae (Theobroma) and 

Malvaceae (Thespesia, Gossypium). This work on 

chloroplast plastid gene rbcL1 confirms several major 

lineages which correspond well with Dalgren (1975) 

taxonomic schemes for Angiosperms. 

From the serology studies of John and Kolbe (1980) 

and Kolbe and John (1980) the further existence of the 

‘Theales’ is not justified if it contains Guttifereae, 

Dipterocarpaceae and Ochnaceae. 

In other works on DNA (Tsumura et al. 1993, 

Wickneswari 1993) parsimony analysis of molecular 

data revealed three major groups which resemble 

conclusions drawn from the anatomy of cotyledonary 

nodes (Maury 1978, Maury-Lechon 1979a,b). These are: 

an ancient group comprising Upuna, Cotylelobium, 

Vatica (V. odorata which is a Sunaptea), an intermediate 

group comprising Dryobalanops and Dipterocarpus, 

and an advanced group comprising Shorea, Hopea and 

Neobalanocarpus. 

Characters Specific to Dipterocarpaceae 

Among the numerous characters cited in the literature 

there is not a single character shared by all species of 

Dipterocarpaceae sensu lato. A detailed study is needed 

to verify the flower bud sepals in all species; a semiquincuncial 

sepal arrangement is reported in 

Pakaraimaea and Monotoideae (Maguire et al. 1977), 

while this arrangement is imbricate (=semi-quincuncial) 

or valvate in Dipterocarpoideae. 

On the contrary three biological characters exist in 

all species of Dipterocarpaceae sensu stricto: the stamen 

architecture (Kostermans 1985) and the pollen type 

(tricolpate grains, exine without endexine: Maury et al. 

1975a, b), and the absence of real post-germinative 

growth of cotyledons (Maury 1978, Maury-Lechon 

1979a, b, Maury-Lechon and Ponge 1979). 

26 

The detailed aspects of stamens were underlined by 

Kostermans (1985): Asian Dipterocarpaceae ‘without a 

single exception, have short, very much flattened, broadly 

(more rarely narrowly) triangular filaments, which 

terminate in a very short, thin, cylindrical part, which 

continues behind the pollen sacs as a cylindrical, often 

differently colored connectival part and often protrudes 

beyond the place of insertion of the pollen sacs, giving 

the impression of a sporophyllous ‘leaf’ to which the 

pollen sacs are attached at the interior surface. The 

anthers are actually dorsifixed, but they appear to be 

basifixed. It seems that the stamen architecture is one 

of the ‘old’ characteristics of Dipterocarpaceae s.s. which 

remained immutable (and hence defines the family, cf. 

Stebbins, 1974 and Melville 1983).’ (Stebbins, 1974 and 

Melville 1983 in Kostermans 1985). 

‘In Monotoideae the filaments are very long, 

cylindrical, thin, the 2-celled anthers are dorsally 

attached, versatile, there is no extension of the filament 

behind the pollen sacs and there is no protruding part 

(this is sometimes also lacking in Dipterocarpoideae, 

but often replaced by this setae). The pollen sacs of 

Dipterocarpoideae are separately and rather loosely (not 

completely) tied to cylindrical dorsal connectival part; 

the 2 or 4 sacs are not much connected, their tips are 

free and pointed or very pointed; the tips have no 

connective tissue. C. Woon and Hsuan Keng (1979) have 

depicted numerous stamens of the Asiatic 

Dipterocarpaceae and all have the same form. In the 

Monotoideae, on the contrary, the 2 pollen sacs are 

united at their apex (not in Marquesia) by a thick, 

triangular-ovoid connectival tissue’ (Kostermans 1985). 

Further investigation is thus needed in Asian, African and 

South American taxa (including the new Colombian taxon 

Pseudomonotes) to judge if these differences have been 

over-estimated. 

Anatomically the presence of wood resin canals and 

multiseriate wood rays, also characterise the Asian taxa. 

Chemically the presence in the resins of the Asian 

dipterocarps of dammaranic triterpenes and 

sesquiterpenes, combined with the absence of 

monoterpenes is also common to all Asian species 

examined by Ourisson’s team (Bisset et al. 1966, 1967, 

1971, Diaz and Ourisson 1966, Diaz et al. 1966). The 

triterpenes derived from the skeleton ‘epoxyde of 

squalene’ (precursor of sterols) constitute a familial 

feature for Dipterocarpaceae sensu stricto. The 

distribution of the other resin compounds (particularly


dipterocarpol) is considered of inferior rank (generic: 

for example, the hydroxydammarenon in genus 

Dipterocarpus; sub-generic: the sesquiterpenes derived 

from humulene at infrageneric rank). 

From the study of pollen, fruit, embryo and young 

seedlings (Maury 1978, Maury-Lechon 1979a, b) 

biological characters of seed germination have been used 

together with pollen types and exine structure, at familial 

and sub-familial levels. Other characters of the anatomy 

of very young seedlings and the morphology of ripe fruit, 

the epidermis of seedling cotyledons and the two first 

leaves, have been hierarchically ordered. Thus a new 

classification was proposed, without formal descriptions, 

to serve as a base for further studies on the delimitation 

of natural groups of taxa inside Dipterocarpaceae sensu 

lato (cf. below: present classifications). 

Supraspecific Taxa in Dipterocarpaceae 

Apart from the above familial and sub-familial rank, four 

principal taxonomic criteria have been expressed (Ashton 

1979c) for definition of supraspecific taxa in current 

revisions: 1) at least a pair of characters which are not 

functionally interrelated; 2) these characters should be 

common to all species in the taxon; 3) there should 

therefore be clear discontinuities in the variation 

between taxa; and 4) the prime goal of taxonomy should 

be to achieve nomenclatural stability. ‘Given these 

criteria, genera must be regarded as essentially artificial 

groupings in the sense that they are defined by breaks in 

the total range of variation’ (Ashton 1979b, p. 129). 

In Asian Dipterocarpaceae most characters 

correspond to two main trends expressed in the tribes 

Dipterocarpi and Shoreae sensu Ashton (1979b), which 

are nearly equivalent to ‘Valvate’ and ‘Imbricate’ groups 

sensu Maury-Lechon (1979a) except the Dryobalanops 

genus, which in the latter is intermediary (certain 

characters of Imbricate type and others of Valvate type). 

In these groups the characters are greatly or weakly 

predominant but their presence (and intensity) is not 

systematic in all species of the group. This situation 

explains the difficulty of establishing clear deliminations 

or affinities. Chromosome numbers (n=7 in most 

Imbricate species and n=11 in Valvate taxa) illustrate 

these facts and provide some explanation. 

The main differences between Ashton’s and Maury- 

Lechon’s two main groupings are: 

1. in Ashton: presence (Shoreae) or absence 

(Dipterocarpi) of the incrassate fruit sepal base (as 

27 

opposed to whole calyx tube), and in most cases the 

basic chromosome n-number is consistent for each 

of the two groups (7 in Shoreae, 11 in Dipterocarpi), 

as also are the scattered (Dipterocarpi) or tangential 

bands (Shoreae) of resin canals; 

2. in Maury-Lechon: some consistent characters for 

each of the two groups (a), and most frequent expression 

of some other characters in each group (b); 

a) Three consistent characters: 

• fruit-sepal base arrangement in ripe fruit: imbricate 

(Imbricate group), valvate (Valvate group) or 

intermediary (mainly Dryobalanops, but also 

Parashorea or Stemonoporus) according to their 

development from flower-bud to open flower. Sepals 

are clearly imbricate before the petals develop 

out of the sepal bud and remain so after the petals 

have grown out of the sepal bud in the Imbricate 

group. The sepals are imbricate at first and then only 

retain some traces of imbrication in Dryobalanops, 

Stemonoporus, Vateria, Marquesia, Monotes; imbricate 

at first and then valvate in all Vatica, and 

valvate all along their development in 

Dipterocarpus; 

• number of strata in pollen exine (3 in Imbricate 

group or 2 in Valvate group, and 4 in Monotoideae 

and Pakaraimoideae); basic chromosome n-number 

(mostly 7 in Imbricate, 11 in Valvate, intermediary 

cases), tilioid structure of exine absent (Imbricate) 

or present (Valvate), and columellae shape-type T 

and Y (Imbricate) or V and U (Valvate), 

• in secondary wood: arrangement of vessels grouped 

(Imbricate) or solitary (Valvate), resin canals in 

bands (Imbricate) or scattered (Valvate) with cellular 

divisions of canal formation radial (Imbricate) 

or oblique (Valvate); 

b) Three most frequent expressions: 

• fruit: number of incrassate bases of sepals (and 

number of accrescent sepals) 2 or 3 (Imbricate) 

or 0 or 5 (Valvate), bases of fruit sepals free (Imbricate) 

or fused (Valvate), type of pericarp tissue 

rigid (Imbricate) or rigid to soft (Valvate), fruit 

equatorial section circular (Imbricate) or circular 

to 3-symmetric (Valvate); 

• embryo: cotyledons ‘covering-piled’ (Maury 1978) 

(Imbricate), neither covering nor piled (Valvate), 

hypocotyl inferior or median-inferior (Imbricate), 

not inferior but apical or median (Valvate); 

• seedling: cotyledons bilobed (Imbricate), or entire 

(Valvate), number of root-xylem poles 4 (Im-


bricate) or 6, 8 or 10 (Valvate), cotyledonary vascular 

bundles uni to trilacunar (Imbricate) or tri to 

multilacunar (Valvate), stomatal types in first leaves 

paracytic, or para-cyclocytic, or anomo-cyclocytic 

(Imbricate) versus cyclocytic or anomocytic or 

anisocytic (Valvate), stomata elongate and sunken 

in the epiderm (Imbricate) or round and raised 

above the epiderm (Valvate). 

The overall pattern of infrageneric variation supports 

the establishment of the two Asian groups Dipterocarpi 

and Shoreae (tribes: Ashton 1979b) or Valvate and 

Imbricate (Maury 1979a). Wood anatomy 

(Parameswaran and Gottwald 1979), palynology and 

characters of fruit-embryo-seedling (Maury et al. 1975a, 

b, Maury 1978, Maury-Lechon 1979a, b, Maury-Lechon 

and Ponge 1979) locate the genus Dryobalanops at an 

intermediary position between Shoreae-Imbricate and 

Dipterocarpi-Valvate groups. Its calyx in ripe fruit is 

subvalvate, thus close to the Valvate group, but the 

chromosome number is 7 as in the Imbricate group. 

The basic distinctions of these two groups are: 

1. Valvate- Dipterocarpi group: base of sepals in calyx 

of ripe fruit valvate, vessels solitary, resin canals scattered; 

basic chromosome number n=11: Vateria, 

Vateriopsis, Stemonoporus, Vatica, Cotylelobium, 

Upuna, Anisoptera, Dipterocarpus. and 

2. Imbricate-Shoreae group: fruit sepals imbricate at 

the incrassate-cupped base of the ripe fruit, vessels 

always grouped, resin canals in tangential bands, basic 

number n=7; Shorea, Parashorea, Hopea, 

Neobalanocarpus. 

Loosely associated genera such as Vateria L. and 

Vateriopsis Heim are distinguishable on many characters 

such as floral parts, bark, fruit, embryo, germination, and 

wood anatomy, as are Stemonoporus Thw., Cotylelobium 

Pierre and Sunaptea Griff. by the same features. The 

other taxa in Vatica L. show the same nervation type and 

rather comparable adult wood or bark anatomy. However, 

a high diversity occurs in vascular structures in the 

seedling cotyledonary-node (Maury 1978; Fig. 677-679, 

p. 309-313, Maury-Lechon 1979a, b; Fig. 16 p. 100). A 

certain diversity also exists in flower-bud development 

(Maury 1978; vol. II tables VI, VII, p. 51-52) or stylestigma 

and stamen shapes (Woon and Keng 1979; Fig. 

30, p. 40). A high diversity is observed in fruit forms of 

sepal aestivation and wing-accrescence, and pericarp or 

sepal incrassatescence (Symington 1943, Ashton 1964; 

Fig. 10, 1968; Fig. 29, Maury 1978; Vol. II, Tables VI, 

28 

VII, p. 51-52). Stemonoporus present a unique terminal 

bud set in a depression at the twig apex which is prolonged 

beyond the last leaf insertion. In Anisoptera the two 

sections are based on floral and bark aspects. Parashorea 

stands out on account of its flower and its 5 or non-winged 

fruit. 

The infrageneric classification of the three genera 

Shorea, Hopea and Neobalanocarpus is more complex. 

Shorea and Hopea differ morphologically by the number 

of thickened bases of calyx (respectively 3 and 2) which 

eventually expand into wings; this is the sole consistent 

difference. Shoreas are mainly tall emergent or canopy 

trees while Hopeas mainly remain understorey or in the 

canopy. Flowers and leaf venation may distinguish the 

two genera but these characters also separate the sections 

within the two genera and are not constant for either genus 

as a whole (Ashton 1979b). Hopea sections differ by 

leaf characters only, while within each section the two 

subsections are principally classified by the shape of the 

floral ovary. 

Heim (1892) first and Symington (1943) afterwards 

produced sound groupings of Shorea and later noticed 

that floral characters closely correlate with field 

characters of bark morphology and wood anatomy in 

defining groups. However, Symington never gave a formal 

nomenclature to his groups (Ashton 1979b). Floral 

characters also correlate with pollination biology (Chan 

1977, 1981, Chan and Appanah 1980, Appanah and Chan 

1981, Appanah 1990). For these reasons and in light of 

the Woon and Keng’s (1979) results, additional emphasis 

has to be put on the importance of stamen shapes in the 

family Dipterocarpaceae. 

Some of the characters by which the groups in 

Shorea are recognised are also those which distinguish 

Cotylelobium, Vatica, Stemonoporus, Vateria and 

Vateriopsis as genera. This evidence was stressed by 

Parameswaran and Gottwald (1979) from wood anatomy 

studies. They stated that groups Anthoshorea, Shorea, 

Richetia, and Mutica merited generic status. However, 

Ashton maintains a different status for these groupings 

in Shorea. On the contrary he gives generic rank to Vatica 

and its allies. He justifies this difference of treatment 

owing to the presence of species with intermediate 

character states between most of the flowers within the 

Shorea group, as opposed to their absence between the 

taxa of the Vatica group. This situation could result from 

different degrees of diversification potential, and merits 

further investigation.


Consistent Criteria for Definition of Species and 

Sub-species 

Consistent criteria (Ashton 1979a) for definition of 

species and sub-species were expressed for 

Dipterocarpaceae as follows: 

1. Size differences are not by themselves sufficient, 

neither are differences of leaf size and shape combined. 

Differences in fruit size are likewise unreliable 

and rarely correlate with other characters; collections 

from one tree in different years often exhibit 

great variation. A consistent discontinuity in leaf 

size, when correlated with differences in androecium 

or gynoecium, in qualitative (not quantitative) characters 

of indumentum, with qualitative characters of 

the twig or stipule or with a discontinuity in the range 

in the number of leaf nerves does constitute an adequate 

criterion for separating species. 

2. Subspecies can be defined where discontinuities occur 

in the range of dimensions of parts, in tomentum 

distribution and in density. However, sometimes taxa 

which share a unique qualitative character, especially 

of fruit or flower, are recognised as subspecies even 

though they may differ qualitatively in vegetative 

parts. 

Experience has shown that this definition of 

subspecies is sometimes too conservative (for example, 

Shorea macroptera ssp. baillonii and ssp. 

macropterifolia occur together in some forests, Vatica 

oblongifolia ssp. multinervosa, ssp. crassilobata and 

ssp. oblongifolia do seem at times to intergrade. This 

definition of subspecies, albeit consistent, is essentially 

arbitrary but may be useful when evidence of 

hybridisation in nature is unavailable. 

Ontogenetic Aspects of Morphological and 

Anatomical Characters 

In Dipterocarpaceae decisions on the primitiveness or 

derived conditions of characters are drawn from personal 

hypotheses on the evolutionary trends within and between 

angiosperm families. Ontogenic trends may follow 

evolutionary trends. Even in the absence of this 

relationship, study of the embryonic trends helps to 

understand taxonomic relations. 

Chemotaxonomic studies have shown (Ourisson 

1979) the existence of certain chemical directions for 

molecular construction in the family: from the epoxyde 

of squalene to the triterpenes of the resins, in all species 

of dipterocarps sensu stricto. 

29 

The embryogenesis from seed germination to young 

seedling in dipterocarps analysed by Maury-Lechon has 

demonstrated a unique direction for the construction of 

the vascular structures in cotyledon node and petiole, 

from simple to very complex (Maury 1978, Maury- 

Lechon 1979a, b, Maury-Lechon and Ponge 1979). This 

trend exists in certain species at different developmental 

phases and morphological levels (node: base, mid-part, 

top of petiole) within a single plant (e.g. Vateria 

copallifera in Maury-Lechon 1979b; photographs Fig. 

49). In other species the trend may be visible by 

comparing plants of a given species (most genera of the 

family: Dipterocarpus, Dryobalanops, Parashorea, 

Vatica sections Vaticae and Pachynocarpus, Vateria, 

Hopea and Shorea) or by comparison of different 

species at a given stage of development and 

morphological level as, for example, from the simple 

trilacunar vascular structures of cotyledonary petiole in 

Shorea curtisii, to the increasing complexity of 

Stemonoporus affinis, S. reticulatus and finally the 

trilacunar appearance of the very complex structure of 

Vateria copallifera (Maury 1978, Maury-Lechon 

1979b). Simplest structures (unilacunar with a single 

resin canal in cotyledonary node) are remarkable in 

Sunaptea, Cotylelobium, Upuna and also exist in certain 

Hopea, Anthoshorea, Richetioides and Muticae. 

Monotes and Marquesia have different (no canals and 

different organisation of vascular bundles) and more 

complex structures than the Asian simplest forms. 

These simplest structures correspond to the taxa with 

small winged fruits, well dispersed by wind, thus again 

with the open areas and long distance migrations. These 

structures allow a better putative relation with the African 

and American taxa (simple but of different type and 

devoid of resin canals). They could evoke ancestral 

dipterocarp migrations in more open, windy and perhaps 

drier environments than those of the present rain forest. 

Polyploidy, Polyembryony, Apomixy and 

Variability of Dipterocarp Characters 

The two basic chromosome numbers tend to remain 

constant within a single genus and between groups of 

genera even in heterogeneous genera like Shorea and 

Hopea. It is premature to say which of the numbers is 

derived or ancestral (Jong and Kaur 1979). There is a 

low frequency of polyploidy series and intraspecific 

polyploids in the Asian genera Shorea and Hopea, 

especially in cases where polyploidy is associated with


agamospermy. Intraspecific polyploidy has been 

reported in Hopea odorata and Dipterocarpus 

tuberculatus (Jong 1976). 

It has been demonstrated that certain species produce 

polyembryonic seeds (Maury 1968, 1970a, b, Kaur et 

al. 1978). Shorea ovalis ssp. sericea is tetraploid with 

frequent polyembryony. These polyembryos originate 

from nucellar cells at the micropilar end of the ovule 

(Jong and Kaur 1979). However, fruit formation in S. 

ovalis requires stimulation of pollination (Chan 1980), 

which is pseudogamous. Pollen tubes have been observed 

in some embryological preparations, thus the possibility 

exists that a zygotic embryo is sometimes formed. Some 

participation by embryo-sac cells other than the egg in 

the formation of pro-embryos, in addition to those 

derived from the nucellus (Jong and Kaur 1979), could 

also be possible. 

On the basis of chromosome number (odd 

polyploidy) and other tentative evidence, it may be 

inferred that all triploids or near triploids (Kaur et al. 

1978) may also be apomicts with polyembryony. The 

triploid condition may have arisen in some cases from 

hybridisation between diploid and tetraploid congeners. 

Agamospermy may indeed provide a mechanism for 

overcoming chromosome sterility, and/or for the 

stabilisation of a heterozygous combination favoured by 

natural selection (Grant 1971 in Jong and Kaur 1979). 

A close association between agamospermy, 

polyploidy and hybridity has been demonstrated in a wide 

range of temperate angiosperms (Gustafsson 1947, 

Stebbins 1960). Even though much available evidence is 

indirect, such a pattern may also occur in Shorea and 

Hopea. 

Apomictic plants are troublesome for taxonomists 

because of the multitude of biotypes or microspecies 

that result from agamospermous reproduction; the 

periodic occurrence of hybridisation involving 

facultative apomicts and related sexual species generate 

additional variant forms which add to the complexity of 

the variation pattern. Some classificatory difficulties in 

Dipterocarpaceae at the supraspecific level presented by 

Shorea and Hopea may well be attributable to the 

presence in each genus of species groups or agamic 

complexes in which sexual and related agamospermous 

taxa exist side by side. Agamospermy whether facultative 

or obligate could well be an important contributory factor 

to the floristic diversity of the lowland mixed dipterocarp 

rain forests of southeast Asia (Kaur et al. 1978). 

30 

Present Classifications 

The four more recent classifications (Tables 1, 2, 8) of 

the family Dipterocarpaceae (Ashton 1964, 1968, 1982, 

Meijer 1963, 1979, Maury 1978, Maury-Lechon 1979a, 

b, Maury-Lechon and Ponge 1979, Kostermans 1978, 

1992) have retained large parts of the previous 

classifications from Heim (1892) and Symington 

(1943). 

Meijer has only taken into consideration the genera 

growing in Sabah and Kostermans has centered his works 

on Sri Lankan taxa and the three non-Asian genera. 

Ashton had a taxonomical approach, while Maury- 

Lechon concentrated on the definition of natural groups 

and their phylogenetic trends. They both utilised the 

results of anatomy studies produced by Whitmore (1963) 

on bark, Gottwald and Parameswaran (1964) and Brazier 

(1979) on wood. Likewise, they used works on cytology 

from Jong and Kaur (1979), embryology, 

chemotaxonomy (Ourisson et al. 1965, Diaz and 

Ourisson 1966, Diaz et al. 1966, Ourisson 1979) and 

Main aspects of Ashton’s classification (see also 

Tables 1, 2, 8) 

Taxonomical levels 

Family (1): Dipterocarpaceae (16 genera, 3 

sub-families, 2 tribes) 

Sub-families (3): Pakaraimoideae: 1 monospecific 

genus, 2 sub-species 

Monotoideae: 2 genera Monotes 

(more than 24 species), 

Marquesia (about 3 species) 

Dipterocarpoideae (2 tribes, 13 

genera, 17 sections, 12 subsections): 

Tribes (2): Dipterocarpi (8 genera, 4-5 

sections): Dipterocarpus, 

Anisoptera (2 sections), Upuna, 

Cotylelobium, Vatica (2-3 

sections), Stemonoporus, 

Vateria, Vateriopsis. 

Shoreae (5 genera, 13 sections, 

12 sub-sections): Hopea (2 

sections, 4 subsections), Shorea 

(11 sections, 8 subsections), 

Parashorea, Neobalanocarpus, 

Dryobalanops.


the herbarium collections of Asia (Forest Research 

Institute Malaysia, Kepong including Symington’s 

collection; Bogor in Indonesia, Peradeniya in Sri Lanka, 

Bangkok in Thailand) and Europe (Kew in U.K, Paris and 

Lyon in France, Leiden in Netherlands) where large 

collections including Ashton’s, Meijer’s, and Maury’s, 

are preserved. 

There is a certain complementarity in the above 

works, but a synthetic classification integrating all 

previous results is still not available. Traditional and 

modern approaches will have to be integrated, including 

DNA and mathematical analyses, as well as the use of 

computer systems for determination and treatment of 

the data. 

Main aspects of Maury-Lechon’s classification (see 

also Tables 1, 2, 8) 

The taxonomical levels have intentionally been left 

without formal definition to serve as a base for further 

research. The relative position of these levels is much 

more important than their names. However, to facilitate 

the explanations, the more proximal names usually 

adopted for these divisions were used (Maury-Lechon 

1979a). 

The separation of Monotoid taxa from the 

Dipterocarpaceae underlines the differences that 

introduce heterogeneity when these taxa are put together 

with the Asian group in the same family. The grouping of 

Monotaceae and Dipterocarpaceae in a supra-familial 

joint division (order or suborder), reminds the greater 

affinities of these two families among the other 

angiosperms. All the other groups aim to underline the 

closer resemblances on the basis of living biological, 

morphological and anatomical characters of the 

successive ontogenic phases (mainly fruit-seed, embryo, 

seedling) and the pollen types and structures. 

The characters of embryo-seedlings and pollens 

strongly emphasise the particular position of 

Anthoshorea close to Doona and partly to Pentacme 

within the Anthoshorinae group, and their position near 

Hopea and Neobalanocarpus in the Imbricate group. 

Still stronger relations exist with the intermediary 

genus Dryobalanops leading directly to the Valvate 

group. Tighter connections of the latter genus occur 

within the Dipterocarpae subgroup (Dipterocarpus and 

Anisoptera) as well as with the Vaticae subgroup through 

Sunaptinae taxa first (Cotylelobium mainly, Sunaptea 

too), and then to Stemonoporus. Through Dryobalanops 


Supra-family 

level (1) (order 

or sub-order): 

Dipterocarpales (2 families) 

Family level Monotaceae (3 genera: 

(2): 

Pakaraimaea, Marquesia, 

Monotes) 

Dipterocarpaceae (2 infra-family 

groups: sub-families, 4 sub-groups: 

tribes, 9 sub-subgroups: sub-tribes, 

19 genera) 

Sub-family Imbricate [2 tribes (a & b), 3 sub- 

level (2): 

tribes, 9 genera, 19 sections] 

Valvate [2 tribes (c & d), 7 subtribes, 

10 genera, 8 sections] 

Tribe level (4): a) Hopeae (2 genera, 4 sections): 

Hopea (4 sections), 

Neobalanocarpus 

b) Shoreae (3 sub-tribes, 7 genera, 15 

sections): 

Anthoshorinae (3 genera): Doona, 

Pentacme, Anthoshorea (2 

sections) 

Shorinae (3 genera): Shorea (2 

sections), Richetia (2 sections), 

Rubroshorea (7 sections) 

Parashorinae (1 genera): 

Parashorea (2 sections) 

c) Dipterocarpae (2 sub-tribes, 3 

genera, 2 sections): 

Dryobalanoinae (1 genus): 

Dryobalanops 

Dipterocarpinae (2 genera): 

Dipterocarpus, Anisoptera (2 

sections) 

d) Vaticae (5 sub-tribes, 7 genera, 6 

sections): 

Upuninae (1 genera): Upuna 

Sunaptinae (2 genera): 

Cotylelobium, Sunaptea (2 sections) 

Vaticinae (1 genus): Vatica pro 

parte (2 sections) 

Stemonoporinae (1 genera): 

Stemonoporus (2 sub-groups) 

Vaterinae (2 genera): Vateria, 

Vateriopsis 

31 

another line of similarities leads to Anisoptera, Vateria 

and Vateriopsis. 

Pollen, embryo and seedling characters, as well as 

parsimony analysis of molecular data (Tsumura et al. 

1993, Wickneswari 1993), show similar conclusions. By 

the pollen surface Anthoshorea resemble 

Dryobalanops, Doona, Neobalanocarpus heimii and 

Cotylelobium, and a basic similarity exists between 

Dryobalanops and Dipterocarpus, as with cotyledonary 

shapes.


Table 8. Comparative classifications of Asian Dipterocarpaceae. 

HEIM 1892 MAURY 1978 MEIJER 1964 ASHTON 1964-68-82 SYMINGTON 1943 

Hopea 

Euhopea, Hancea, 

Dryobalanoides, 

Petalandra 

Pierrea 

Duvallelia 

Balanocarpus 

Eubalanocarpus 

Pachynocarpoides 

Microcarpae 

Sphaerocarpae 

Parahopea 

Doona 

Pentacme 

Isoptera 

Richetia 

Shorea 

Eushorea 

Anthoshorea 

Hopeoides 

Richetioides 

Rugosae 

Brachypterae 

Brachypterae 

Smithianeae 

Pachycarpae 

Parashorea 

Dryobalanops 

Baillonodendron 

Dipterocarpus 

Sphaerales, 

Angulati, Plicati, 

Alati, Tuberculati 

Anisoptera 

Pilosae, Glabrae, 

Antherotriche 

Cotylelobium 

Sunaptea 

Dyerella 

Vatica 

Euvatica, Isauxis 

Retinodendron 

Pachynocarpus 

Stemonoporus 

Eustemonoporus 

Monoporandra 

Vesquella 

Vateriopsis 

Vateria 

Paenoe 

Hemiphractum 

IMBRICATE 

SUB-VALVATE 

VALVATE 

Hopea 

Hopeae 

Pierreae 

Dryobalanoides 

Sphaerocarpae 

Balanocarpus hemii 

Doona 

Pentacme 

Antoshorea 

Anthoshoreae 

Bracteolatae 

Shorea 

Shoreae 

Barbatae 

Richetia 

Richetioides 

Maximae 

Rubroshorea 

Muticae 

Muticae 

Auriculatae 

Ovalis 

Rubellae 

Brachypterae 

Brachypterae 

Smithianeae 

Pachycarpae 

Parashorea 

Tomentellae 

Smithianae 

Dryobalanops 

(Dryobalanoinae) 

Dipterocarpus 

Anisoptera 

Anisoptera, 

Glabrae 

Anthoshorinae Shorinae Parashorinae 

Upuna (Upuninae) 

------------------------ 

Cotylelobium 

Sunaptea 

Sunapteae 

Vaticoides 

------------------------ 

Vatica 

Vaticae 

Pachynocarpus 

-------------------------------------- 

Stemonoporus 

Sphaerae 

Ovoides 

------------------------ 

Vateriopsis 

Vateria 

Hopea 

Group I 

Group II 

Pentacme 

Shorea 

Anthoshorea 

Shorea 

Isoptera 

Barbata, Ciliata 

Richetia 

Rubroshorea 

Parvifolia subgr. 

Ovalis subgr. 

Pauciflora subgr. 

Smithiana subgr. 

Pinanga subgr. 

Parashorea 

Hopea 

Hopea 

Hopea, Pierrea 



Sphaerocarpae 


hemii 

Shorea 

Pentacme 

Doona 

Anthoshorea 

Shorea 

Shorea, Barbata 

Neohopea 

Richetioides 

Richetioides 

Polyandrae 

Mutica 

Mutica 

Auriculatae 

Ovalis 

Rubellae 

Brachypterae 

Brachypterae 

Smithiana 

Pachycarpae 

Parashorea 

*: Ashton 1964, 1968, spelling changes in and after 1982; subgr.: sub-group. 

Hopeae Shoreae 

Dipterocarpae Vaticae 

Dipterocarpinae 

Sunaptinae Vaticinae 

Stemonoporinae 

Vaterinae 

Doona 

Dryobalanops 

Dipterocarpus 

Anisoptera 

Pilosae 

Glabrae 

Upuna 

Cotylelobium 

Vatica 

Synaptea 

Isauxis 

Pachynocarpus 

SHOREA 

DIPTEROCARPI 

Dryobalanops 

Dipterocarpus 

Anisoptera 

Anisoptera 

Glabrae 

Upuna 

Cotylelobium 

Vatica 

Sunaptea 

Vatica 

(Pachynocarpus*) 

Stemonoporus 

Vateriopsis* 

Vateria 

Hopea 

Euhopea 

Pierrea 


Bracteata 

Balanocarpus 

hemii 

Pentacme 

Shorea 

Meranti Pa’ang 

Balau 

Damar hitam 

Parvifolia 

subgr. 

Ovalis subgr. 

Pauciflora 

subgr. 

Parashorea 

Dryobalanops 

Dipterocarpus 

Anisoptera 

Pilosae 

Glabrae 

Cotylelobium 

Vatica 

Synaptea 

Isauxis 

Pachynocarpus 

Red Meranti 

32


These features group together the living and fossil 

dipterocarps which have lived together in the same 

phytogeographical areas (Table 5). They also correspond 

to the subsequent hypothesis concerning their potential 

for differentiation (see above). They also suggest an 

eventual remote relation from Anthoshorea to 

Marquesia and then to Monotes. 

In the Valvate division the Dipterocarpinae group 

underlines the relations between the 2 subgroups: 1) 

Dryobalanops, and 2) Dipterocarpus with Anisoptera. 

Similarly the Vaticinae group emphasises the 

existence of 5 subgroups: 

1. Upuna alone but near subgroup 2; 

2. Cotylelobium close to Sunaptea; 

3. Vatica pro-parte intermediate between groups 1 and 

2, and 4 and 5; 

4. Stemonoporus position between subgroups 2 and 5; 

5. Vateria, and Vateriopsis with particular similarities 

(cotyledon position and shape, germination type) with 

Vatica and also Anisoptera, Stemonoporus and 

Cotylelobium. 

Vatica genus (excluding Sunaptea) stands somewhat 

isolated within the family Dipterocarpaceae by the pollen 

characters principally, and much less so by some aspects 

of embryo shape and structure (particularly seedling 

vascular structure). As mentioned, the tilioid surface of 

the pollen exine could suggest a proximity with 

monotoid taxa, however, its structure is definitely 

different (Maury et al. 1975a, b). New investigations are 

needed on a great number of species of Vatica genus 

sensu lato (including Sunaptea and Pachynocarpus) to 

permit a clearer view on infra-generic variation of these 

characters. 

Main lines of Kostermans’ classification (See also 

Tables 1, 2) 

Kostermans has mainly considered the Sri Lankan taxa 

so that, as in Meijer’s work, only the Asian genera 

represented in this geographical area were analysed in 

detail. Contrary to Maury-Lechon’s work he formally 

described the Monotaceae family, as well as genera 

Doona and Sunaptea (the latter including 

Cotylelobium). No publication remains on his views 

concerning the affinities of the genera inside his 

Dipterocarpaceae family, nor in eventual sections within 

the Shorea genus which includes Pentacme (Kostermans 

1992). In Stemonoporus he suggests 2 sub-divisions 

based on the pericarp aperture at germination (character 

used in Maury 1978 and Maury-Lechon 1979a, b). 


Family (2): 

Monotaceae (3 genera): Pakaraimaea, 

Marquesia, Monotes 

Dipterocarpaceae (15 genera): Hopea, 

Neobalanocarpus, Balanocarpus, Shorea, 

Doona, Parashorea, Dipterocarpus, 

Anisoptera, Dryobalanops, Upuna, 

Sunaptea (Cotylelobium included), Vatica, 

Stemonoporus, Vateria, Vateriopsis 

33 

Main Recent Taxonomic Changes: 

They successively concerned the: 

1. establishment of subgenera Shorea, Anthoshorea, 

Richetia and Rubroshorea (Meijer 1963, Meijer and 

Wood 1964); 

2. establishment of 11 sections in genus Shorea including 

the previous genera Doona and Pentacme (Ashton 

1964, 1968, 1980, 1982); 

3. proposition to re-install some ancient genera such 

as Doona Thw., Anthoshorea Heim and Richetia 

Heim outside genus Shorea Gaertn., Sunaptea Griff. 

outside genus Vatica and Vateriopsis Heim out of 

genus Vateria L. (Maury 1978, Maury-Lechon 

1979a, b); 

4. acceptance of the re-establishment of Vateriopsis 

genus by Ashton (1982); 

5. announcement of the discovery and description of 

Pakaraimaea (Maguire 1979, Maguire and Ashton 

1980, Maguire and Steyermark 1981); 

6. formal re-establishment of genus Doona Thw. outside 

Shorea (Kostermans 1984), genus 

Banalocarpus Beddome outside Hopea and independent 

from Neobalanocarpus (Kostermans 

1981a) genus Sunaptea outside genus Vatica but including 

genus Cotylelobium (Kostermans 1987); and 

7. discovery and description of Pseudomonotes 

tropenbosii (Londoño et al. 1995) which is included 

in Monotoideae sensu Maguire et al. (1977) close 

to the African Monotes and Marquesia. 

Discussion and Conclusions 

Morphology, as well as anatomy and ecophysiology, 

shows many characters tightly related to their biological 

functions, and these functions are connected to both the 

biotic associations and the climatic environmental 

features which influence flower pollination, seed


dispersal and seedling survival. It has been shown that 

the species diversity of ovary and style sizes and shapes 

in the Shorea sensu lato subdivisions correspond to 

their pollinator insect groups (Appanah 1990). The 

successive ontogenic phases of flower aestivation and 

seed germination, and seedling construction, also reveal 

the existence of particular directions such as the change 

of sepal position from imbricate in the young flower to 

subvalvate or valvate in ripe fruit (not the reverse), the 

multiplication of vascular bundles and resin canals (never 

the reverse) and the presence/absence of a postgerminative 

growth of cotyledonary limbs (Maury 1978, 

Maury-Lechon 1979a, b). 

The biological plasticity of ripe seeds and seedlings 

depends on their ability to maintain their biological 

functions (which are dependent on their structures). This 

plasticity determines the possibilities of survival in a 

changing environment or in new and different places. The 

exact knowledge of the present geographical distribution 

and the main ecological features of these places already 

allow useful speculations for the choice of species 

potentially able to adapt in more drastic conditions 

(Maury-Lechon 1993, 1996, Xu and Yu 1982). This is, 

for example, the case of species from the seasonal 

tropics, or from the aseasonal regions subjected to great 

changes (diurnal, seasonal or unpredictable climatic 

events) on dry sands of sea coastal areas, or temporary 

and alternatively flooded or dried areas. The evolutionary 

trends established by Hopea or Shorea allow a much 

wider range of possible future adaptations than do those 

established by Vateria or Stemonoporus (Maury-Lechon 

1979b). Thus a second level of prediction for adaptability 

is possible. 

Overall, the striking feature of the family is the high 

variability of characters within and between species, 

within and between individual trees in many cases, and 

even within a single seed in certain species. Furthermore, 

the present classification demonstrates a heterogeneity 

of levels between the two Asian subgroups Dipterocarpi 

and Shoreae sensu Ashton. A notable case is the 11 

sections of the Shorea genus in the Shoreae subgroup. 

They have unequal hierarchic levels when comparisons 

are established between the two Asian subgroups. 

Sections such as Anthoshorea, Shoreae or Richetioides 

of the Shoreae tribe have much higher rank than sections 

Rubellae or Ovalis for example, and a similar level to 

Doona, Pentacme, and Parashorea in the Imbricate 

34 

group (sensu Maury-Lechon). A similar situation appears 

for the Vatica genus in the Dipterocarpi subgroup (sensu 

Ashton). For this reason Sunaptea (Vatica pro-parte in 

certain cases) has again been raised to generic rank 

(Kostermans 1987). These difficulties underline the 

complexity of the family. However certain well defined 

genera exist, such as Dryobalanops, Dipterocarpus, 

Anisoptera and Upuna. It could thus be hoped that a more 

equal weighting of characters is still possible in building 

a more homogeneous classification in the complex parts 

of the family, and that criteria can be defined for Asian 

dipterocarps to determine generic rank. 

Present supraspecific taxa are mainly defined by 

groups of characters concerning the morphological 

aspects of leaves, the sequence fruit-seed-embryoseedling, 

flowers, bark and wood, and colour and 

consistency of resins. Anatomical structures (wood, bark, 

petioles, epidermis, germinating seeds and seedlings), 

stomatal types, and chemistry of resins, have historically 

clarified the definition of supraspecific taxa (but very 

few wood characters are specific) in Dipterocarpaceae. 

However anatomical or chemo-taxonomical groups are 

not yet totally integrated into the present taxonomic 

divisions. This is the case for genus Dipterocarpus for 

example, in which phytochemistry has recognised main 

groups without evident correspondence with the previous 

morphological divisions based on fruit characters 

(Meijer 1979). The main reason for this is that too few 

species have been studied in this way to permit a rigorous 

understanding of within group variability and thereby 

establish which characters provide useful criteria for 

defining groups and hierarchic levels. 

The morphological variability in the seasonal tropics 

may be the result of the frequent great changes with time 

in distribution of habitats and geographical boundaries. 

These changes favoured variability but rarely provided 

prolonged isolation mechanisms for fertility barriers to 

evolve. The frequency of hybrids suggests the same 

conclusion. It is not really known to what extent 

competition eliminates these hybrids. In the aseasonal 

tropics dipterocarps appear to be outbreeding species. 

Many other species in aseasonal tropics present 

allopatric differentiation and clear discontinuities in 

variation. Facultative apomixis may produce successful 

genotypes and accelerate ecotypic differentiation and 

short term evolution. Apomixis could serve to maintain 

fecundity where sexual reproduction is inadequate, and


perhaps allow rapid spread of favourable ecotypes in 

heterogeneous terrain. However apomixis should 

probably lead to lowered genetic variability within 

populations, and thus could still result in increased 

probability of extinction when unadapted to the 

environmental changes. Constant differences in habit, 

morphology and reproductive biology exist between 

emergent and understorey trees (Richards 1952, Hallé 

et al. 1978, Yap 1982 in Ashton 1984). Ontogenesis thus 

follows predictable patterns in the larger trees and 

selection acts on these characteristics, which include tree 

habit and leaf shape, irrespective of their systematic 

relationships. These species present an ecological 

complementarity. It is however, not certain that species 

sharing a common habitat and geography will identically 

respond from flower-bud initiation to sensitivity to light, 

mycorrhizal invasion, water stress, seed predation, or 

share the same pollinators and seed vectors (M.S. Ashton 

1992, 1995). Ecological criteria such as seed water 

content and seed resistance to desiccation are 

expressions of the species’ biological plasticity and the 

possible complementarity between species (Maury- 

Lechon 1993). Certain species are abundant and others 

rare. Detailed systematic and biosystematic comparisons 

between rare and abundant congeners should bring some 

answers (Ashton 1992). We need to confirm whether 

the means exist for gene flow, and then to directly 

measure the level and pattern of genetic variability. 

Amenable source of evidence could also be tested for 

the presence or absence of associations between 

population distributions in space, and demographic and 

population genetic studies are also required on the basis 

of repeated observations. These observations and the use 

of species’ complementarity would reinforce the 

understanding of systematics and help the forest 

managers to make decisions in rehabilitation and 

conservation programmes (Maury-Lechon 1991, 1993, 

1996). 

Bearing in mind the economic value and the present 

status of Dipterocarpaceae, it is also urgent to relate 

phylogeny to comparative ecology within genera and 

sections by a combination of: 

1. molecular phylogenic studies, concentrating on genera 

and below; 

2. comparative demographic studies of groups of related 

species, especially those which co-occur: 

a) during the reproductive phase (bud to recruit, and 

including fecundity), 

b) during stand development and trough to mortality, 

35 

c) population genetics of selected species under selected 

conditions; 

3. comparative ecophysiological experiments on seeds 

and seedlings; and 

4. competition experiments. 

The teams exist within the frame of the International 

Working Group on Dipterocarps (IWGD-IUFRO S.07- 

17 Working Party) and contacts have already been taken 

between the authors of this paper and their direct 

commentators for this purpose. Field and laboratory 

works will be organised on the base of complementarity. 

Overall, a cooperative, integrated and detailed reassessment 

is needed for the whole family 

Dipterocarpaceae sensu lato, with the establishment of 

an evolutionary classification based on a general 

consensus. Several groups around the world are currently 

working on projects that can lead to such a solution. They 

include: Forest Research Institute Malaysia, Kepong and 

Unité Mixte de Recherche 5558 du Centre National de 

la Recherche Scientifique with Lyon University (France), 

in association with Harvard University (USA), who are 

working on the genetic analysis of DNA sequences; 

Massachusetts University (USA) on genetic and breeding 

system studies (Murawski and Bawa 1994, Murawski et 

al. 1994); and Cambridge-Edinburgh (U.K.) on computer 

identification keys (Newman et al. 1995). Several more 

complementary works are needed in palynology (pollen 

and exine), stamen architecture and shape (the flower 

being observed from the pollination point of view), 

ontogenesis (structure and morphology of fruit-embryogermination, 

anatomy of cotyledonary node and petiole, 

epidermis of primordial leaves), wood anatomy, 

chemotaxonomy and architecture of juvenile stage. These 

complementary works will require broad cooperation 

with colleagues and institutions from the Asian, African 

and South American zones. More effort is needed for 

the Asian dipterocarps from China, Burma and Indo- 

China. Likewise, there is a need for contact with African 

colleagues 


We express our sincere thanks to the Center for 

International Forestry Research and most particularly to 

C. Cossalter for having initiated and supported the 

realisation of a book which provides the first broad 

synthesis of the present status of knowledge on 

dipterocarps that complements the proceedings of the


five Round Table Conference Proceedings on 

Dipterocarps (1977 to 1994). Special thanks are also 

addressed to T.C. Whitmore (Cambridge University) and 

P.S. Ashton (Harvard Institute for International 

Development) for their advice and corrections, to R. 

Grantham (Lyon University) for his contribution to the 

final presentation, to S. Appanah (Forest Research 

Institute Malaysia) who reviewed this text and shared the 

editing tasks with C. Cossalter, and to C. Elouard (French 

Institute of Pondicherry). Thanks also to colleagues from 

Lyon and Paris who contributed to the bibliographic 

effort. 

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Conservation of Genetic Resources 

in the Dipterocarpaceae 

K.S. Bawa 

Introduction 

The biological and economic importance of 

Dipterocarpaceae lies in the extraordinary dominance of 

its members over vast areas in forests of southeast Asia. 

With approximately 510 species and 16 genera, the family 

may not be particularly large among tropical woody 

groups. Other families such as Euphorbiaceae, 

Myrtaceae, Rubiaceae, Annonaceae, and Lauraceae have 

more taxa than the Dipterocarpaceae, however, they are 

pantropical in distribution. Although members of 

Dipterocarpaceae are also found in the African and 

American tropics, 13 out of 16 genera and 470 out of 510 

species are largely restricted to Asia, and there, restricted 

primarily to south and southeast Asia. In Malaysia, it is 

certainly among the six largest families that are 

predominantly woody, the others being Euphorbiaceae, 

Myrtaceae, Rubiaceae, Annonaceae, and Lauraceae. 

Moreover, the members of the family are exceedingly 

abundant in lowland forests of southeast Asia, for 

example, in many areas, 80% of the emergent individuals 

and 40% of understorey trees are dipterocarps (Ashton 

1982). Thus, when one considers the relatively restricted 

distribution of the family, both diversity and abundance 

are its main attributes. 

The diversity of the family is under assault from 

deforestation and habitat alteration. Effective in-situ and 

ex-situ conservation strategies are required to conserve 

the existing genetic resources. To conserve genetic 

resources, it is essential not only to maintain existing 

diversity, but also to understand the ecological and 

evolutionary processes that have been responsible for the 

origin, evolution, and maintenance of diversity at 

intraspecific and higher taxonomic levels. This chapter 

has two broad objectives. One is to review genetic 

mechanisms responsible for the origin and maintenance 

of diversity. The second is to identify areas of research 

that may elucidate patterns and processes of diversity and 

Chapter 2 

a more complete understanding of factors regulating 

diversity. It is assumed that a better understanding of 

diversity and the mechanisms maintaining diversity may 

be helpful in developing effective strategies for 

conservation of genetic resources. The chapter ends with 

a brief commentary on the institutions involved in 

research related to the conservation genetics of the family. 

Diversity 

Genetic mechanisms responsible for diversification at 

intraspecific and specific levels are considered and then 

patterns of genetic variation within and among 

populations are described. 

Chromosomal Differentiation 

Information about chromosome numbers is available for 

9 out of 15 genera and 68 out of 510 species of the family 

(Jong and Kaur 1979, Ashton 1982). Species and genera 

are remarkably uniform with respect to chromosome 

number. Perhaps all species in the genera Dryobalanops, 

Hopea, Neobalanocarpus, Parashorea, and Shorea have 

x=7 as the basic number. Anisoptera, Dipterocarpus, 

Upuna, and Vatica seem to have x=11 as the basic 

number. Several species in the genera with x=7 as the 

basic number have a somatic chromosome number of 20, 

21 and 22. Thus, x=11 may have been derived from x=7 

through alloploidy. 

Polyploid species are known in only two genera: 

Hopea and Shorea. In Hopea, polyploidy has been 

reported in 5 out of 9 species and in Shorea in 3 out of 36 

species. Five of these polyploid species are triploids 

(2n=21; also 2n=20 and 22) and one (2n=20) seems to 

be an aneuploid derivative of a triploid. Many of the 

triploids are apomictic (see below).

Conservation of Genetic Resources in the dipterocarpaceae 46 

Table 1. Infraspecific variation in chromosome number 

(from Ashton 1982). 

Species Chromosome 

Number 

Dipterocarpus alatus 20, 22 

D. tuberculatus 20 

D. tuberculatus var. turbinatus 30 

Hopea beccariana 20, 21, 22 

H. odorata 14, 20, 21, 22 

H. subalata 20, 21, 22 

Aneuploid series are common in Anisoptera and 

Dipterocarpus. Both genera have species with 2n=20 or 

2n=22. In some taxa, both variants occur within the same 

species (Table 1). 

Thus, both polyploidy and aneuploidy indicate the 

importance of chromosomal variation in diversification 

at the species level. However, the largest genus, Shorea, 

shows remarkable uniformity in chromosome number; 

31 out of the 34 species, for which chromosome numbers 

are known, have the same diploid number, viz. 2n=14. 

Intraspecific variation in chromosome number has 

been reported in several species, particularly in 

Dipterocarpus and Hopea (Table 1). Of the 68 species 

for which chromosome numbers are available, 6 species 

have been recorded to show intraspecific variation. Such 

variation has not been reported for species of Shorea, the 

largest genus of the family even though data are available 

for 34 species. 

Inter and intraspecific variation in chromosome 

numbers is difficult to interpret for two reasons. First, 

more than one chromosome numbers for the same taxon 

have been reported by different rather than the same 

author. Second, much of the reported variation due to 

reports of a single author, Tixier (1960) and most of 

Tixier’s counts have not been confirmed by others. 

It should, also be kept in mind that information on 

chromosome number for large tropical trees is usually 

obtained from very small sample sizes. Often only one 

or two individuals in a population are examined and rarely 

is there data from more than one population. Thus, it is 

impossible from available data to determine the magnitude 

of intraspecific variation in chromosome number. 

Furthermore, even in these cases, where such variation 

has been reported, one cannot estimate the extent of 

variation and therefore its significance. For example, for 

species of Dipterocarpus as well as Hopea listed in Table 

1, variation is in the form of either aneuploid or 

polyploid chromosomal series, but whether this variation 

is in the form of occasional aneuploid or polyploid 

populations is not known (Ashton 1982). 

Breeding Systems 

Breeding systems are one of the primary determinants of 

the pattern of genetic diversity in natural populations of 

plants (Hamrick 1982, Hamrick and Godt 1989). 

Outcrossing combined with extensive movement of pollen 

and seed can lead to a high degree of genetic variation 

within populations but reduce differentiation among 

populations. Selfing and limited mobility of pollen and 

seed can have the opposite effect of reducing variation 

within, but promoting differentiation among populations. 

Dipterocarpaceae have bisexual flowers which are 

pollinated by a variety of animal vectors (see below). 

Controlled pollinations have revealed the presence of selfincompatibility 

systems in a large number of species. At 

least 14 out of 17 species appear to be self-incompatible 

(Table 2.) The self-incompatibility system in several 

species is apparently weak, as is the case in many other 

tropical species. In most of the species subjected to 

controlled pollination so far, a certain proportion of selfpollinated 

flowers set fruits. Dayanandan et al. (1990) 

and Momose et al. (1994) suggest that fruit set in self 

and cross-pollinated flowers is initially high but during 

development, fruits from self-pollinated flowers suffer 

from higher abortion rates than fruits from crosspollinated 

flowers. T. Inoue (personal communication) 

has implicated the existence of a post-zygotic 

incompatibility system in Dryobalanops lanceolata. Such 

systems have also been reported for other tropical forest 

trees (Bawa 1979, Seavey and Bawa 1983). 

On the basis of controlled pollinations, most 

dipterocarps appear to be strongly cross-pollinated. 

Outcrossing is the usual mode of reproduction in tropical 

forest trees (Ashton 1969, Bawa 1974, 1979, 1990, and 

references therein.) However, in dipterocarps, studies of 

breeding systems conducted so far are based on very small 

sample sizes in very few species. The data of Dayanandan 

et al. (1990) are from 2-3 trees, mostly two of each 

species; of Chan (1981) from 1-2 trees, and of Momose 

et al. (1994) from only one tree. Considering the 

variability among trees and that the distinction between 

self-compatibility and self-incompatibility in the family 

appears to be quantitative, large sample sizes will be 

required to precisely define the self-incompatibility 

systems.


Table 2. Breeding systems of Dipterocarps. 

Species/Section 

Percent Fruit Set Inferred Breeding References 

Selfed Crossed 

System 

Shorea cordifolia 

Section Doona 

0.5 21.0 Self-incompatible 1 

S. disticha 

Section Doona 


S. trapezifolia 

Section Doona 


S. trapezifolia 

Section Doona 


S. hemsleyana 

Section Muticae 


S. macroptera 



S. lepidota 



S. acuminata 



S. leprosula 



S. splendida 

Section Pachycarpae 


S. stenoptera 

Section Pachycarpae 

n/a n/a Self-incompatible 2 

S. ovalis 

Section Ovales 

16.2 17.6 Self-compatible 2 

Dipterocarpus oblongifolius 69.3 64.0 Self-compatible 2 

Dryobalanops lanceolata n/a n/a Self-incompatible 3 

Hopea glabra 

n/a n/a Self-compatible or 

2 

Section Richetioides 

apomictic 

S. maxima 

Section Richetioides 

n/a n/a Self-incompatible 2 

S. multiflora n/a n/a Self-incompatible 2 

1: Dayanandan et al. (1990); 2: Chan (1981); 3: Mamose et al. (1994). 

Outcrossing Rates 

More recently genetic markers in the form of allozymes 

have been used to quantify mating systems in species of 

Dryobalanops, Hopea, Shorea, and Stemonoporus. 

Analysis of mating systems on the basis of markers allows 

examination of the progeny arrays of many trees in the 

population. Moreover, outcrossing rate (tm) can be 

quantified between zero and one; zero representing 

complete selfing and one indicating 100% outcrossing. 

Mating systems of species examined so far are shown in 

Table 3. The outcrossing rates range from 0.617 in Shorea 

trapezifolia to 0.898 in Stemonoporus oblongifolius. 

The average rates expressed in Table 3 mask considerable 

variation among trees and years. The rate varied from 

0.49 to 1.00 among trees in Shorea congestiflora 

(Murawski et al. 1994a, b). S. megistophylla trees in the 

logged forests had a lower outcrossing rate than trees in 

undisturbed forests (Murawski et al. 1994b). The 

difference seems to be dependent on the density of 

reproductive trees. Such density-dependent differences 

in outcrossing rates have also been shown in several other


Table 3. Outcrossing rates of Dipterocarps. 

Species Outcrossing Rate tm 

(± standard error) 

tropical tree species (Murawski and Hamrick 1990, 

1991). Apparently, the low density of trees in logged 

stands reduces the inter-tree movement of pollinators 

and promotes self-pollination. Selfing, in part, may be 

aided by a weak self-incompatibility system. 

Kitamura et al. (1994) compared outcrossing rates 

of Dryobalanops aromatica in primary and secondary 

forests but found no significant differences. 

Self-incompatibility as well as mating system studies 

suggest that dipterocarps are predominantly outcrossed. 

Outcrossing in large populations can allow populations 

to harbour considerable genetic variation. In dipterocarps, 

mass flowering is also likely to enhance outcrossing by 

allowing exchange of gametes among a very large number 

of individuals. Not surprisingly, therefore, populations 

of dipterocarps show considerable genetic variation (see 

below). 

Although the analysis of mating systems shows that 

the rates of outcrossing are high, it is also clear that there 

is a considerable potential for selfing in almost all species 

examined so far. Moreover, apomixis has been reported 

in several species (see below). While outcrossing 

continuously generates new genetic variation, potential 

for self-pollination and apomixis allows occasional new 

variants to spread in the population or colonise new sites, 

and thereby promote differentiation of taxa. 

Pollen and Seed Dispersal 

Pollen dispersal influences the mating system, and both 

pollen and seed dispersal affect population genetic 

structure. Limited dispersal results in inbreeding, small 

effective population sizes, and a high level of 

differentiation among populations. Extensive dispersal 

has the opposite effect. 

Outcrossing in dipterocarps is achieved through a 

wide variety of pollinators that differ in their foraging 

References 

Dryobalanops 

0.794 (±0.059) – Kitamura et al. (1994) 

aromatica 

0.856 (±0.063) 

Shorea congestiflora 0.874 (±0.021) Murawski et al. (1994) 

S. megistophylla 0.860 (±0.058) Murawski et al. (1994) 

S. trapezifolia 0.617 (±0.033) Murawski et al. (1994) 

Stemonoporus 

0.898 (±0.022) Murawski & Bawa 

oblongifolius 

(1993) 

ranges and therefore disperse pollen 

over varying distances. Appanah and 

Chan (1981) implicated thrips as 

pollen vectors for several species of 

Malaysian species of Shorea, section 

Muticae. The thrips breed in flower 

buds of the species they pollinate and, 

as flowering progresses, they multiply 

in number. The adult thrips feed on 

stamens and petals. As the petals of the 

flowers are shed from the tree the 

thrips fall on the ground and then move 

to a new cohort of subsequently opened flowers. The 

distances over which thrips move are not known but, 

because of their relatively small body size, they 

apparently do not fly over long distances. It is presumed 

that their restricted movement is not a drawback in their 

effectiveness as pollinators because the species they 

pollinate are relatively abundant. 

Dayanandan et al. (1990) present evidence for 

pollination of Shorea megistophylla (section Doona) and 

Vateria copallifera by bees (Apis spp.). They also 

observed a wide variety of other insect floral visitors 

including thrips. However, the thrips acted as flower 

predators rather than pollinators, particularly in V. 

copallifera. 

More recently, Momose et al. (1994) have presented 

evidence for pollination of Dryobalanops lanceolata, a 

large canopy tree species in Sarawak, by medium sized, 

stingless bees (Trigona spp.). They also noted the presence 

of many other types of flower visitors (Coleoptera and 

Diptera). Momose et al. suggest that medium sized, 

stingless bees constitute an important group of pollen 

vectors for canopy and subcanopy trees in Sarawak (see 

also Chan and Appanah 1980). 

Clearly, the dipterocarps are pollinated by a wide 

variety of insects. The three detailed studies, respectively 

by Appanah and Chan (1981), Dayanandan et al. (1990), 

and Momose et al. (1994) have revealed three different 

classes of pollinators. Ashton (1982) in his extensive 

review also lists beetles and moths as flower visitors, 

although their role in pollination has not yet been 

demonstrated. Among the pollinators implicated so far, 

all, except thrips, are capable of moving pollen over long 

distances. The extent of gene flow via pollen (and seeds) 

is further discussed in subsequent sections. 

Seed dispersal in most dipterocarps is by wind 

(Ashton 1982). In most species, sepals are modified into


wing like structures in the fruits that allow the single 

seeded fruits to gyrate toward the ground. Many species 

growing in swamps or river banks have fruits with short 

sepals and may be dispersed by water (Ashton 1982). In 

some dipterocarps, such as species of Stemonoporus, 

fruits are without wing-like sepals. When mature, they 

simply fall on the ground and are apparently not removed 

by any disperser (Murawski and Bawa 1994) although 

rodents are known to hoard the seeds and, perhaps, aid in 

dispersal (P. Ashton, personal communication). Seeds 

disseminated by wind and water can potentially disperse 

over long distances. In Shorea albida, dissemination by 

wind up to 2 km has been documented (Ashton 1982) 

and although dispersal by water has not been observed in 

any species seeds may move over long distances in water 

channels. 

Apomixis 

Apomixis has been reported in several taxa of the family. 

There is embryological evidence for the existence of 

multiple embryos originating from a single ovule in 

Shorea ovalis ssp. sericea and S. agamii ssp. agamii 

(Kaur et al. 1978). Multiple seedlings from a single fruit, 

indicative of polyembryony, have been reported in 

Anisoptera curtisii, Dipterocarpus baudii, D. cornutus, 

D. costulatus, Dryobalanops aromatica, Hopea 

odorata, H. subalata, Parashorea densiflora, Shorea 

argentifolia, S. gratissima, S. macrophylla, S. 

parvifolia, S. pauciflora, S. smithiana, Vatica pallida 

and V. pauciflora (Kaur et al. 1978 and references 

therein) and in Shorea trapezifolia (S. Dayanandan, 

personal commnication). The percentage of multiple 

seedlings is low in all these species except for S. 

macroptera, S. resinosa, H. odorata, and H. subalata 

in which 30-70%, 98%, 90% and 21% seeds respectively 

have multiple seedlings. 

Interestingly, a recent study by Wickneswari and 

Norwati (1994) indicates that multiple seedlings from 

the same seed in Hopea odorata have different 

genotypes raising the possibility that multiple seedlings 

may not necessarily involve apomixis. Furthermore, 

using genetic markers, a high outcrossing rate has been 

estimated for the species (Table 3). Isozyme surveys also 

reveal high amounts of genetic diversity within 

populations (Wickneswari et al. 1994). 

Apomixis is associated with triploidy in Shorea 

resinosa and Hopea subalata (also possibly in H. 

latifolia) but other species displaying polyembryony are 

mostly diploid. The ovary in Dipterocarpaceae is usually 

three locular with two ovules in each loculus. Normally, 

only one ovule develops into a seed thus, multiple 

seedlings can result from occasional development of 

seeds from more than one ovule and the presence of such 

seedlings need not always imply apomixis. 

Apparently, in some species apomixis is widespread 

while in others it occurs occasionally. Obligate apomixis 

for either individual trees or populations (and species) 

remains to be demonstrated but is a possibility in taxa 

with a triploid chromosome number. Certainly among 

tropical woody families apomixis at a scale comparable 

to Dipterocarpaceae has not been reported. Moreover, 

considering that most species in the family have a low 

diploid chromosome number, the common occurrence 

of apomixis is puzzling because apomixis is usually 

associated with polyploidy and hybridisation. 

Apomixis could have played an important role in 

evolution of the family. New genetic combinations 

arising through mutations or hybridisation that may be 

partially or completely sterile can be perpetuated by 

apomixis. Vegetative multiplication can also maintain 

heterozygosity for a long time. In addition, apomixis and 

self-pollination may allow new genetic variants to spread 

at new sites. Subsequent restoration of sexual 

reproduction and outcrossing, combined with mutation, 

can introduce genetic variation in the new isolates. 

Hybridisation 

Hybridisation has played an important role in the 

evolution and diversification of angiosperms (Stebbins 

1950). Hybrids in tropical trees are assumed to be rare 

(Ashton 1969). In dipterocarps, however, hybrids have 

been frequently reported. Ashton (1982) suggests that 

many triploid taxa in the family could be of infraspecific 

hybrid origin. His list includes the following: Hopea 

subalata, H. odorata, Shorea ovalis ssp. sericea, 

Neobalanocarpus heimii, Shorea leprosula, S. curtisii, 

and hybrids between Vatica rassak and V. umbonata, and 

Anisoptera costata and A. curtisii. Many examples of 

putative hybrids between species of Dipterocarpus have 

also been reported (Symington 1943). Apomixis, already 

noted in several taxa of the family, could certainly allow 

the hybrids to persist until sexual fertility is restored. 

Although several of the interspecific hybrids are 

polyploids, polyploidy in the family has so far been 

recorded in relatively few taxa. On the other hand, the 

base number x=11 observed in several genera of the 

family itself could be of ancient alloploid derivation.


Genetic Diversity Within and Among Populations 

The existence of self-incompatibility and high 

outcrossing rates suggest that populations of 

dipterocarps should harbour high levels of genetic 

variation. Indeed, recent studies, based on analysis of 

variation at isozyme loci, have revealed considerable 

genetic variation in natural populations. A high level of 

enzymatic polymorphism in natural populations of 

Shorea leprosula was first detected by Gan et al. (1977). 

More recently, genetic diversity within and among 

populations of several species of Hopea (Wickneswari 

et al. 1994), Shorea (Harada et al. 1994), 

Stemonoporus (Murawski and Bawa 1994, Dayanandan 

and Bawa, unpublished data) has been quantified. Genetic 

diversity in many Malaysian species of Hopea and 

Shorea were studied using Random Amplified 

Polymorphic DNAs (RAPD). Considerable variation was 

found both within and among populations. The level of 

diversity in species of Hopea (Wickneswari et al. 1996) 

was less than in species of Shorea (Harada et al. 1994). 

In these studies, methods to characterise genetic 

diversity depended upon several assumptions about the 

segregation and homology of bands. Moreover, results 

from most RAPD surveys cannot be compared with those 

obtained from isozyme surveys because dominance at 

RAPD ‘loci’ makes it impossible to distinguish 

heterozygotes from homozygotes. Thus, genetic diversity 

cannot be characterised in conventional terms. Bawa and 

his associates have used isozymes to estimate genetic 

diversity in species of Stemonoporus and Shorea. In 

Stemonoporus oblongifolius, the percent of 

polymorphic loci range from 89% to 100%, the average 

number of alleles per polymorphic locus is 3.1 and mean 

genetic diversity for the species is 0.297. The number 

of loci sampled was 9 and was the same sampled for other 

dipterocarps and tropical trees. The estimates of genetic 

diversity are among the highest reported for plant species 

(Murawski and Bawa 1994). The values for the above 

parameters are lower for Shorea trapezifolia, but remain 

toward the higher end of the value reported for tropical 

trees. Similarly, a high level of genetic variation has been 

observed in Shorea megistophylla (Murawski et al. 

1994b) and several other species of Stemonoporus 

(Murawski and Bawa, unpublished). Wickneswari et al. 

(1994) also report high levels of variation in Hopea 

odorata on the basis of isozyme studies. 

Inter-population differentiation on the basis of 

isozyme surveys has been studied in only three species: 

Stemonoporus oblongifolius (Murawski and Bawa 

1994), Shorea trapezifolia (Dayanandan and Bawa, in 

preparation) and Hopea odorata (Wickneswari et al. 

1994). In all cases, there is a high level of variation among 

populations. In Stemonoporus oblongifolius, the mean Gst 

value, which is a measure of population differentiation, 

is 0.16. In other words, 16% of total genetic diversity is 

due to differences among populations. Interestingly, the 

distance among sampled populations ranged from 1.3 to 

9.7 km. Thus, populations seem to differ over a relatively 

small spatial scale. In Shorea trapezifolia too the Gst value 

was high (0.11); in this case the most distant were 

separated by 43.5 km. The mean genetic distance between 

populations in Hopea odorata was 0.10 (Wickneswari et 

al. 1994). 

The high level of genetic differentiation could be due 

to either restricted gene flow or local selection. Direct 

observations of gene flow in dipterocarps are lacking. 

Seed dispersal in Stemonoporus oblongifolius seems to 

be passive; the one seeded, heavy, resinous fruit drop 

under the maternal tree and the seed germinates without 

being removed by any disperser (Murawski and Bawa 

1994). In Shorea trapezifolia, the seeds are dispersed by 

gyration, assisted by wind with most seeds falling within 

the vicinity of the parent. Gyration of fruits, referred to 

earlier, may have evolved as an adaptation to restrict 

dispersal to the sites in which the parents are found. Thus, 

gene dispersal via seeds in both species does not generally 

occur over large distances. 

The degree of gene dispersal via pollen would depend 

upon the pollinators. Medium sized to large bees should 

be able to bring about long-distance dispersal more 

frequently than small bees or thrips. Both Stemonoporus 

oblongifolius and Shorea trapezifolia are pollinated by 

medium-sized bees (Apis spp.). 

Gene dispersal has been indirectly measured in 

Shorea trapezifolia (more than one migrant per 

generation). Nm estimates the degree of migration 

between populations, and a value of Nm>1 is enough to 

prevent population differentiation due to drift for neutral 

loci (Wright 1931, Maruyama 1970, Slatkin and 

Maruyama 1975). In S. trapezifolia, the value of Nm is 

1.62. This high value indicates that differentiation in S. 

trapezifolia is not due to restricted gene flow. 

Ashton (1982, 1988) has shown that congeneric 

species in the family often occupy different edaphic zones. 

Moreover, within the same habitat related species may 

be differentiated along environmental gradients that


define the regeneration ‘niche’. Therefore, genetic 

selection within taxa of the family can readily be moulded 

by fine and coarse grain variation in the environment. 

Thus, the inter-population differentiation observed in 

Stemonoporus oblongifolius and Shorea trapezifolia is 

consistent with the hypothesis that slight variation in the 

habitat can allow genetic variants to differentiate along 

environmental gradients despite low or moderate levels 

of gene flow. 

Summary of Diversification Processes 

Most dipterocarps are outcrossed and diploid. Speciation 

seems to have involved allopatric differentiation of widely 

outcrossing populations; differentiation seems to have 

occurred in response to differences in soils and habitats 

(Ashton 1969). Aneuploidy, polyploidy, and hybridisation 

may have also assumed a role in the spread of some 

variants arising as a result of hybridisation and changes 

in chromosome number. At the intraspecific level, 

outcrossing maintains high levels of genetic variation in 

populations. Mass flowering combined with abundance 

of adults probably ensures large effective population 

sizes. Nevertheless, despite extensive gene flow, 

selection results in differentiation of populations over 

relatively small scales. 

Research Needs 

Future research needs may be best examined in the context 

of threats to diversity. Genetic resources are imperilled 

by deforestation and forest fragmentation. Moreover, 

selective logging often can lead to reduction in genetic 

variation (Kemp 1992) and alter population structure with 

concomitant changes in demography and genetics of 

subsequent generations (Bawa 1993). Global climatic 

change is also expected to influence plant populations, 

but the potential effects, deleterious or beneficial, are not 

well defined, particularly for the areas where dipterocarps 

are dominant. 

Deforestation and forest fragmentation may influence 

diversity in several ways. Species or populations may 

become extinct or severely endangered. At the population 

level, once seemingly large, contiguous populations may 

be broken into relatively small, remnant patches, 

physically isolated from each other. Over time, gene 

exchange among the remnant patches may be completely 

eliminated and the small populations may be subject to 

inbreeding. Habitat fragmentation can also increase 

overall levels of variation if isolated populations diverge 

from each other. The consequences of fragmentation 

depend upon the degree and duration of isolation and the 

size of the isolated population. 

Fragmentation of habitats may have deleterious 

effects on both the ecosystem dominants as well as rare 

species. The ecosystem dominants may have very large 

populations, and fragmentation may result in loss of 

genetic diversity (Holsinger 1993). Rare species may face 

severe reduction in population size following 

fragmentation. Many species of dipterocarps have adult 

population densities as low as 0.07 to 0.30 individuals 

per hectare (Ashton 1988). Some of these species occur 

in low population densities at more than one site and may 

be particularly prone to inbreeding. In addition, there may 

be selection for apomixis in such situations (P. Ashton, 

personal communication). 

Selective logging can also increase the potential for 

inbreeding. Logging temporarily reduces adult population 

densities. In many tropical tree species, inbreeding has 

been shown to be a function of stand density (Murawski 

and Hamrick 1990, 1991). In Shorea megistophylla, as 

noted above, the rates of inbreeding are higher for trees 

from logged stands than for trees in unlogged stands. 

However, it should be noted that stands in properly 

managed forests regenerate from seedlings established 

prior to logging. In dipterocarps, the potential for 

inbreeding is also increased by the fact that selfincompatibility 

barriers are not strong; trees in many 

species are capable of setting seeds after self-pollination, 

but here again selfed seeds may be selected against in the 

presence of outcrossed seeds in the same inflorescence. 

The longevity of trees may not allow many of the 

assumed deleterious consequences of forest fragmentation 

and selective logging to be manifested for a long time. 

Even in small patches, trees may set fruits and seeds and 

regenerate without apparent ill-effects. Comparative 

studies of reproductive output, mating patterns, and 

regeneration processes involving trees in large contiguous 

forests and small fragments may reveal the consequences 

of habitat alteration. 

Thus, in order to fully understand the effects of 

deforestation, forest fragmentation, and forest 

management practices on forest genetic resources of 

dipterocarps, we need a better understanding of patterns 

of diversity and processes that maintain diversity. Areas 

of research that require immediate attention are outlined 

below.


Species Level Research 

First, we have to identify species that are endangered or 

threatened with extinction. In some instances, populations 

of species themselves might be large, but the types of 

forests in which such species occur may be disappearing 

at a very rapid rate. Examples are the moist seasonal 

evergreen forests on the western slopes of western Ghats 

in India and throughout Indochina, the mixed dipterocarp 

forests in the southwest region of Sri Lanka, and the 

dipterocarp forests in Philippines. Fortunately, due to the 

work of Ashton (1982, 1988) and others, as compared to 

other tropical families, there is far more information 

available on the geographical ranges of various species 

and the type of habitats and soil types occupied by these 

species. More recently, P. Ashton has reviewed the 

conservation status of all Asian dipterocarps for the World 

Conservation Monitoring Centre at Cambridge, UK. All 

this information along with other data on land use patterns, 

fragmentation, and deforestation should be combined in 

a geographical information system to provide easily 

comprehensible graphical information on the current 

status of distribution of species and the conservation status 

of the forests in which they occur. Such a database would 

be particularly useful because, in many cases, information 

on the conservation status of family members is equivalent 

to information on conservation status of dipterocarp 

forests, the most important and dominant vegetation in 

very large areas of south Asia and southeast Asia. 

Second, we need to identify centres of taxonomic 

diversity and active speciation in the family. Centres of 

taxonomic diversity, of course, are known on the basis 

of morphological criteria (Ashton 1982, 1988). Molecular 

techniques, however, provide means to rapidly assess 

species relationships and to elucidate patterns of 

speciation. For example, within section Doona of Shorea, 

molecular data indicates that the ‘Beraliya’ group is 

evolving at a higher rate than the remaining species (S. 

Dayanandan, personal communication). 

Third, comparative studies of genetic diversity in 

species that occupy centres of diversity and those that 

occur away from zones of diversification may provide 

further insights into patterns of genetic diversity. As 

mentioned earlier, Murawski and Bawa (1994) observed 

an unusually high level of genetic variation in natural 

populations of Stemonoporus oblongifolius. The genus 

is endemic to Sri Lanka and has undergone active 

speciation in a small region in the southwest region of 

the island. The high diversity observed by Murawski and 

Bawa may be due to the fact that this species is found in 

a region which is the centre of active speciation. 

Similarly, comparative studies of related common and 

rare species, or species in different ecological zones 

may provide additional insights into patterns of genetic 

diversity among species. 

Fourth, there is an urgent need to study the effects of 

logging on genetic diversity and other population genetic 

parameters such as inbreeding and gene flow. Gene 

Resources Areas that are being established in Malaysia 

(Tsai and Yuan 1995) may provide excellent opportunities 

for such comparative research. 

Fifth, we need a better understanding of the 

importance of chromosomal variation, apomixis, and 

hybridisation in diversification at the species level and 

infraspecific levels. We know the species and genera in 

which these processes occur. However, our knowledge 

with respect to the incidence and ecological and 

evolutionary importance, particularly, of apomixis and 

hybridisation is very limited. Again, molecular 

techniques now offer new opportunities to assess the 

significance of these processes. 

Finally, information on breeding systems and 

pollination mechanisms is required for many taxa to 

characterise genetic factors maintaining genetic variation. 

Such information is available for only a few species. Many 

large genera such as Dipterocarpus and Hopea remain 

unexplored. 

Intraspecific Level Research 

First, the most urgent need is the characterisation of the 

patterns of genetic variation in important species. 

However, in addition to ecosystem dominants and species 

of commercial importance, we also need to analyse the 

genetic structure of rare species. A better understanding 

of the spatial organisation of genetic variation is critical 

to the assessment of the effects of deforestation and forest 

fragmentation on genetic diversity. 

Second, comparative studies of gene flow in 

contiguous and fragmented forests can provide 

information about the effective size of populations, 

microevolutionary forces responsible for genetic 

differentiation among populations, and the potential 

effects of deforestation and fragmentation on genetic 

isolation of populations that were once contiguous. 

Third, comparative studies of central and peripheral 

populations may be useful in revealing pockets of high 

genetic diversity. Populations in the centre of a species


range may often show more genetic variation than 

peripheral populations due to a higher rate of gene 

exchange in central populations. 

Fourth, the effect of genetic diversity and inbreeding 

on population viability should be an area of utmost 

concern. Are reduced levels of genetic diversity and 

outcrossing associated with a decline in fitness? Decrease 

in fitness may be manifested as reduction in fruit and 

seed set, seedling vigour and overall recruitment and 

regeneration. It is thus critical to link genetic studies and 

demographic studies. Comparative studies of gene flow 

in fragmented and contiguous forests, described earlier, 

should incorporate comparative studies of the effects of 

genetic variation and inbreeding on reproductive output 

and regeneration. Mass flowering in dipterocarps also 

offers opportunities to gain insights into the relationship 

between genetic diversity and population recruitment. 

Sporadic flowering in off years may reduce the effective 

population size, increase inbreeding and mortality of seeds 

due to predation and lead to a disproportionately low level 

of recruitment. Comparative genetic and demographic 

studies during mass and sporadic off year flowering can 

provide useful information about possible effects of 

reduction in population size. 

Fifth, many species of dipterocarps display 

intraspecific variation in chromosome number and 

apomixis. However, the frequency of chromosomal 

variants or apomixis within or among populations is not 

documented. There are now molecular tools to rapidly 

assay populations for the incidence of chromosomal 

variation, apomixis and hybridisation. 

Site-specific Research 

The rates of deforestation vary widely among the regions. 

Species diversity of dipterocarps is also not uniform 

throughout South and Southeast Asia. Thus, from a 

geographical perspective, high priority should be 

accorded to regions that are undergoing rapid 

deforestation and those that have very high species 

richness. 

The Philippines, Sri Lanka and the Western Ghats of 

south India have been converted into other forms of land 

uses at a high rate during the last fifty years. These areas 

have certainly lost unique populations and perhaps species 

of dipterocarps. In such areas, there is an immediate need 

to assess the conservation status of various taxa building 

on P. Ashton’s earlier review. Sri Lanka particularly 

deserves serious consideration because of the high 

degree of endemism: 6 out of 7 genera and 45 out of 46 

species of dipterocarps are endemic to the country. 

The greatest species diversity in the family is found 

in northwest Borneo. However, much of the cytology 

and genetic research cited in this paper has been 

conducted on species from Peninsular Malaysia and Sri 

Lanka. Data from genetics and population biology of the 

taxa that occur in northwest Borneo should provide useful 

insights into mechanisms regulating differentiation 

within and among species. 

Institutional Capability and Constraints 

P.S. Ashton , J. Liu, P. Hall and their associates (Harvard 

University), S. Appanah, H. Chan, and others (Forest 

Research Institute Malaysia (FRIM)) have played a key 

role in advancing our knowledge of the systematics, 

biogeography, and ecology of the family. Research in 

systematics and ecology is being continued at Harvard 

University. At FRIM the scope of research in genetic 

resources has been recently enlarged to include such 

areas as molecular evolution and population genetics. 

In Sri Lanka, N. Gunatilleke and S. Gunatilleke at 

the University of Peradeniya have a major research 

programme on conservation biology of dipterocarps. 

This programme includes research on population biology 

and population genetics. N. Gunatilleke and S. 

Gunatilleke have collaborated with P. Ashton (Harvard 

University), K. Bawa and D. Murawski (University of 

Massachusetts, Boston). 

Another major centre of research on population 

biology and genetics of dipterocarps is the Kyoto 

University. T. Inoue, K. Momose and R. Terauchi are 

involved in detailed studies of phenology, pollination 

biology and genetics of dipterocarp species in Sarawak. 

The work is a part of a major programme on canopy 

research in dipterocarp forests. 

S. Dayanandan and R. Primack (Boston University) 

are working in collaboration with P. Ashton on a diverse 

range of issues in dipterocarp biology, from molecular 

biology to population dynamics. 

Recently, the Center for International Forestry 

Research (CIFOR) and International Plant Genetic 

Resources Institute (IPGRI) have initiated a project on 

population genetics, specifically on the effects of forest 

fragmentation, logging and non-logging disturbance on 

genetic diversity of some dipterocarps. This programme


involves a number of institutions in India, Indonesia, 

Malaysia and Thailand. 

Apart from insufficient funding, the major factor 

constraining progress has been the lack of a coordinated 

programme with clear objectives and predetermined 

priorities. With the establishment of institutions such 

as CIFOR and IPGRI, it should be possible to undertake 

a cohesive research programme with well defined goals. 


I thank Peter Ashton (Harvard Institute for International 

Development), Tim Boyle (Center for International Forestry 

Research), S. Dayanandan (University of Alberta), 

and S. Appanah (Forest Research Institute Malaysia) for 

their comments. This work is supported in part by Center 

for International Forestry Research and in part by 

grants from the U.S. National Science Foundation, Pew 

Charitable Trusts, and the MacArthur Foundation. 

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Guttiférales récoltées au Laos. Revue Cytolologique 

Biologique vég. 29: 65-70. 

Tsai, L. M. and Yuan, Ct. T. 1995. A practical approach to 

conservation of genetic diversity in Malyasia: genetic 

resource area. In: Boyle, T. J. B. and Boontawee, B. 

(eds.) Measuring and monitoring biodiversity in 

tropical and temperate forests, 207-217. CIFOR, 

Jakarta. 

Wickneswari, R. and Norwati, M. 1994. Genetic 

variation in polyembryos of Hopea odorata Roxb. 

In: Koh, C.L. (ed.) Proceedings of the 1st National 

Congress on Genetics, 76-78. Genetics Society of 

Malaysia, Kuala Lumpur. 

Wickneswari, R., Zawawi, I., Lee, S.L. and Norwati, M. 

1994. Genetic diversity of remnant and planted 

populations of Hopea odorata Roxb. in Peninsular 

Malaysia. Proceedings International Workshop, BIO 

REFOR, Kangar, Malaysia. 

Wickneswari, R., Norwati, M., Tsumura, Y., Kawahara, 

T., Yoshimaru, H. andYoshimura, K. 1996. Genetic 

diversity of tropical tree species: genetic variation 

of Hopea species (Dipterocarpaceae) using RAPD 

markers. Meeting on Global Environmental 

Research Project on Tropical Forest Ecosystem 

Project, National Institute of Environmental 

Sciences, Tsukaba, Japan. 

Wright, S. 1931. Evolution in Mendelian populations. 

Genetics 16: 97-159.

Seed Physiology 

P.B. Tompsett 

Seed is the natural vehicle for gene movement and 

storage. It is the usual form in which germplasm is 

collected. When procedures can be devised to transport 

and retain material in this form, many of the technical 

problems associated with other methods can be avoided. 

This advantage renders seed especially appropriate for 

users in tropical and subtropical countries. In general, 

seed is the most common form of propagation for 

afforestation and is the form in which breeding stock is 

usually retained. There are, however, considerable 

problems remaining in the use of seed. Some of these 

are discussed below for dipterocarp species in relation 

to the underlying seed physiology processes. 

Much pioneering work on agricultural crop seed 

physiology was conducted over the last 20 years (see 

below for some references) and the principles 

discovered often apply to seed of woody species. These 

earlier results have been translated into technological 

principles. Thus, manuals have been published on the 

design of seed storage facilities (Cromarty et al. 1982), 

seed management techniques (Ellis et al. 1984) and a 

handbook on seed technology for genebanks (Ellis et al. 

1985). Knowledge of seed physiology has thus improved 

practical handling and management of crop seeds. 

Compared to crop species, relatively little research 

has been published on tropical and subtropical tree seed 

technology and physiology. Publications have been 

produced following IUFRO Seed Problems Group 

meetings, the most recent of which was held in Tanzania 

(Olsen 1996). Another source of information is a 

summary of some relevant seed physiology projects 

which has recently been published in database form 

(Tompsett and Kemp 1996a, b). Also, a seed compendium 

has been published which supplies succinct entries on 

many tropical trees (Hong et al. 1996). 

A considerable amount of empirical work on the 

storage of forest tree seed has been carried out; a 

sampling is given in Chapter 4. A more physiological 

research approach is relatively new. Many tree species 

have seed that is desiccation-sensitive (‘recalcitrant’), 

Chapter 3 

so that moisture physiology is especially important for 

this group. However, in a recent review article on water 

in relation to seed storage, the section on desiccationsensitive 

seeds comprised only 4% of the article 

(Roberts and Ellis 1989). More attention is, however, 

now being given to recalcitrant seeds (see, for example, 

Berjak and Pammenter 1996). 

Framework of the Review 

Germination is basic to all aspects of seed studies; work 

on germination physiology, especially in relation to 

temperature, is thus considered first. Another important 

experimental consideration is the physiological 

condition of the seed at the time of harvest, moisture 

content being the single most important factor; this is 

considered next in the review. Thirdly, the effect of 

desiccation is discussed; knowledge of this factor enables 

seed to be classed as orthodox (tolerant) or recalcitrant 

(intolerant). Finally, the effects of seed storage are 

considered. 

The Review 

Germination 

In general, dipterocarp seeds germinate quickly under 

moist, warm conditions. 

Early germination studies took the form of nursery 

assessments leading to ecological conclusions. A major 

study of this type, in which 56 dipterocarp species were 

assessed for germination rate and final germination, is 

that of Ng (1980); no conclusions can be made 

concerning temperature effects. Conditions were more 

closely controlled in experiments on Shorea roxburghii, 

S. robusta and S. almon by Tompsett (1985); results 

showed optimum germination in the range 26-31°C for 

these three species. Corbineau and Come (1986) found 

that final germination reached nearly 100% for both 

Hopea odorata and S. roxburghii over a broad range of 

temperatures, but 30-35°C was deemed optimal because 

germination rates were faster. More recently, a standard

Seed Physiology 58 

Table 1.Optimum germination temperatures, germination rates and base temperature values (Tompsett and Kemp 

1996a, b). 

Species Base 

temperature 

(°C) 

approach was adopted in a series of germination studies 

so that physiological parameters could be assessed 

(Table 1). Optimum germination for 30 species was 

confirmed as lying between 26°C and 31°C and the value 

for the time to 50% final germination, at the optimum 

temperature, generally ranged between 2 and 13 days. 

There were two Dipterocarpus species (D. alatus and 

D. costatus) that were much slower to germinate, 

however. 

A physiological parameter relating to the theoretical 

value at which zero growth occurs was also assessed; 

this is referred to as the ‘base temperature’. The base 

Time to 50% final 

germination at the 

optimum 

temperature (d) 

Optimum 

temperature 

(°C) 

Radicle length 

defining 

germination 

(mm) 

Shorea almon n/a n/a 26 5 

Shorea siamensis n/a n/a 26 5 

Shorea smithiana n/a n/a 26 3 

Shorea pinanga n/a n/a 31 10 

Hopea parviflora n/a 11 31 5 

Shorea roxburghii n/a 11 31 5 

Dipterocarpus alatus n/a 20 26 5 

Shorea robusta n/a 4 31 5 

Anisoptera marginata n/a 6 26 3 

Hopea foxworthyi 6.4 4 31 3 

Hopea odorata 7.1 2 31 5 

Shorea leprosula 7.5 4 31 3 

Shorea parvifolia 8.3 4 26 3 

Shorea contorta 8.8 7 26 3 

Shorea affinis 9.4 5 26 3 

Parashorea smythiesii 9.7 3 31 3 

Dipterocarpus costatus 10.2 30 31 3 

Dryobalanops aromatica 10.5 3 26 3 

Parashorea tomentella 10.5 9 31 10 

Shorea guiso 11.0 4 26 3 

Anisoptera costata 11.4 5 26 3 

Shorea ferruginea 12.2 6 26 3 

Dipterocarpus obtusifolius 12.4 3 26 5 

Cotylelobium burckii 13.0 9 26 3 

Vatica mangachapoi 13.0 9 31 3 

Dipterocarpus turbinatus 15.2 5 26 10 

Cotylelobium melanoxylon 15.4 6 31 3 

Dipterocarpus tuberculatus 15.8 13 31 5 

Shorea amplexicaulis 15.8 7 26 3 

Dipterocarpus zeylanicus 16.2 11 26 10 

Shorea argentifolia 16.4 4 26 3 

temperature is the intersect on the temperature axis for 

the plot of germination rate against temperature. More 

details of technique are given in Tompsett and Kemp 

(1996a, b); the parameter is further discussed below. 

Chilling damage 

Studies show germination is reduced or does not occur 

at temperatures below 16°C for several dipterocarp 

species (Tompsett 1985, Corbineau and Come 1986), 

due to chilling damage. This type of damage is also 

observed in the results from storage research; data show 

a reduced ability to survive low temperature conditions


(Sasaki 1980, Yap 1981, Tompsett 1985, Corbineau and 

Come 1986). 

Sasaki (1980) considered seeds of (i) Shorea species 

in the ‘yellow and white meranti’ groups, (ii) Hopea, (iii) 

Dipterocarpus, (iv) Vatica, (v) Dryobalanops, (vi) 

Balanocarpus and (vii) Parashorea to be tolerant down 

to 4°C. By contrast, he believed that seeds of Shorea 

species in the ‘red meranti and balau’ groups were 

intolerant of temperatures below 15°C. He classified 

Anisoptera as a tolerant genus in a separate publication 

(Sasaki 1979). Yap (1981) later proposed a three-group 

classification: firstly, seed of species in the 

Dipterocarpus, Dryobalanops, Neobalanocarpus and 

Vatica genera were said to be intolerant of temperatures 

below 14°C; secondly, seed of Shorea species in the 

sections Mutica, Pachycarpae and Brachypterae were 

considered intolerant of temperatures below 22°-28°C; 

and, finally, seed of Shorea species in the Anthoshorea 

section and seed of Hopea and Parashorea could be 

cooled to 4°C (but were recommended to be stored at 

14°C). Further details relating taxonomic classification 

to chilling damage are given elsewhere (Tompsett 1992). 

A possible explanation for the above inconsistencies is 

that different authors have studied the effects of chilling 

for different periods of time, leading to different 

conclusions; exposure of seed to longer periods of 

chilling can show up chilling damage which might 

otherwise have ben missed in the case of relatively chillresistant 

species. 

The processes behind the chilling physiology 

phenomenon have not been adequately studied. However, 

differences among species in susceptibility to chilling 

damage are confirmed by the base temperature data in 

Table 1. In particular, Hopea species appear the most 

resistant to chilling damage, since they have the lowest 

base temperatures. A low value for the base temperature 

is expected if germination ability decreases relatively 

slowly as germination temperature is reduced. It should 

be emphasised, however, that these results apply 

exclusively to moist seeds. Storage of dry orthodox 

dipterocarp seeds at low temperatures is described in 

the storage section below. 

The differences in chilling tolerance of seeds among 

dipterocarp species are quantitative rather than qualitative. 

Seed of the ‘tolerant’ species S. roxburghii eventually 

suffers damage at 2°C -5°C relative to seed kept at 

warmer temperatures (Purohit et al. 1982, Tompsett 

1985). Another example of chilling damage which 

occurred over a lengthy period of time is that to H. 

hainanensis. For this species, seed at 5°C almost all 

died after 6 months; by contrast, at 15°C -20 °C no loss 

of viability occurred (Song et al. 1984). 

Harvest and Maturity 

The condition of seed at harvest is of primary concern 

in the planning of all physiological experiments. 

Moisture contents at or near harvest are given for 25 

species in Table 2, including examples from both seasonal 

and aseasonal dipterocarp forests. Seeds of the three 

species with the lowest moisture contents, which are 

found in seasonal forests, were collected from the 

ground after natural desiccation. For these seeds, drying 

occurs very swiftly after abscission because the open 

canopy exposes them to direct sunlight. Of the remaining 

species listed, some are derived from the dry forest and 

others from moist areas; they possessed a relatively high 

range of post-processing moisture contents between 29 

and 56% (usually, seeds were just de-winged). 

It has been realised for some time that there can be a 

considerable difference between whole-seed moisture 

content and moisture content of the embryo or embryo 

axis (Grout et al.,1983). Since axis or embryo moisture 

content is more closely related to basic physiological 

processes than whole seed moisture content, it is a 

preferable measure to use herein. Axis values have been 

determined for dipterocarp species (Table 2) and range 

from 51 to 74%, except in the case of the much lower 

value for the dry-zone species Dipterocarpus 

tuberculatus, which was collected after natural drying. 

Seed maturation 

A few developmental studies have been carried out on 

dipterocarp species; whole-seed moisture content has 

been employed in most of these as the main physiological 

criterion. Sasaki (1980) reported that the moisture 

content (wet basis) of Shorea roxburghii declined from 

60 to 50% in the final 3 weeks of maturation on the tree. 

Panochit et al. (1986) reported a comparable decline 

from 40 to 30% for the same species, whilst a reduction 

from 59 to 49% was reported for S. siamensis (Panochit 

et al. 1984). 

Nautiyal and Purohit (1985a) assessed changes 

during maturation of S. robusta seed; they described 

these changes as biphasic. Over the 60 days from anthesis 

to maturity, concentrations of soluble carbohydrates, 

starch, soluble protein and acid phosphatase were


Table 2. Percentage moisture content and oil values for processed, dewinged whole seed and excised 

seed parts (Tompsett and Kemp 1996a, b). 

Species Whole-seed moisture 

content* (percentage) 

*: calculated on wet weight basis; 

**: calculated on dry weight basis; 

determined; in addition, declining moisture content, 

increasing germination and increasing weight of the seed 

were recorded. One theory proposed was that early 

desiccation of the seed coat may be connected with poor 

viability; this explanation appears unlikely since S. 

roxburghii has similar seed coat structures and is much 

longer-lived. 

An interaction of maturity with chilling damage has 

been noted. Increased resistance to such damage was 

observed as maturity approached for S. siamensis 

(Panochit et al. 1984); germination declined to zero and 

25% for seed collected 4 and 2 weeks respectively 

before maturity after storage for 28 days at 2°C, but 

mature seed still gave about 60% germination after 56 

days of similar storage. The same effect was noted for 

S. roxburghii (Panochit et al. 1986). 

Axis moisture 

content* 

(percentage) 

Embryo oil content 

** (percentage) 

Dipterocarpus intricatus *** 8 n/a 16 

Dipterocarpus alatus *** 11 n/a 7 

Dipterocarpus tuberculatus *** 11 13 19 

Shorea ferruginea 29 n/a 61 

Shorea argentifolia 29 51 n/a 

Hopea ferrea 32 n/a 9 

Shorea parvifolia 32 62 n/a 

Hopea foxworthyi 34 52 n/a 

Hopea odorata 36 54 20 

Shorea gibbosa 37 64 n/a 

Dipterocarpus costatus 38 n/a 10 

Shorea macrophylla 38 66 n/a 

Parashorea tomentella 40 63 n/a 

Shorea amplexicaulis 40 69 57 

Dipterocarpus grandiflorus 40 70 n/a 

Anisoptera costata 42 n/a 33 

Shorea fallax 42 70 n/a 

Shorea affinis 44 63 n/a 

Dipterocarpus chartaceus 47 n/a 8 

Shorea leptoderma 47 61 n/a 

Parashorea malaanonan 48 66 n/a 

Dryobalanops keithii 50 56 n/a 

Stemonoporus canaliculatus 53 64 n/a 

Shorea macroptera 55 n/a n/a 

Dipterocarpus obtusifolius 56 74 n/a 

***: seeds of OLDA (orthodox with limited desiccation 

ability) dried naturally in the field. 

Tang and Tamari (1973) were the first to report the 

post-harvest-maturation phenomenon for dipterocarp 

seeds. They found that Hopea helferi and H. odorata 

seeds blown down prematurely by a high wind increased 

in germination during storage. The effect was observed 

over a period of about one week for seeds held at 15 °C. 

Desiccation Studies 

Knowledge of its storage physiology category, which can 

be derived from desiccation studies, is the single most 

useful piece of physiological information about a seed. 

It is the key to correct seed handling procedures. 

Seed storage physiology categories 

Three storage category designations are recognised. Of 

these, the main two are:


§ orthodox; and 

§ recalcitrant. 

The orthodox type is capable of desiccation to a low 

moisture content (approximately 5%) and storage for 

several years at -20 o C with little loss of viability (Roberts 

1973). By contrast, the recalcitrant type is not capable 

of desiccation to a low moisture content without loss of 

germination capacity and cannot be stored for long 

periods of time (Roberts 1973). 

A third category of seed storage physiology has been 

described. It was first defined in 1984 in relation to 

Araucaria columnaris seed (Tompsett 1984) and was 

termed ‘orthodox with limited desiccation ability’ 

(OLDA). A similar category was later defined for coffee 

seed and termed ‘intermediate’; the name denotes its 

partial tolerance of desiccation (Ellis et al. 1990, 1991). 

Some recent evidence (Tompsett, unpublished), however, 

confirms there may be little physiological difference 

between this third category of seed and the orthodox 

type. There are, however, important practical handling 

difficulties associated with this third category. These 

problems justify its retention as a distinct storage type. 

Some tropical seed is additionally subject to lowtemperature 

damage when stored in the moist condition 

(chilling damage, see pages 58-59). As a result, further 

categories could have been included. However, it was 

considered preferable, for the sake of simplicity, to 

employ only the three desiccation-damage-based 

categories described above. To date, all dipterocarp 

species examined have been found to be subject to 

chilling damage when moist. 

Desiccation physiology 

Curves of germination percentage against moisture 

content percentage can be plotted for the results from 

controlled desiccation studies. These curves illustrate 

whether the seed is recalcitrant or not and give 

parameters for the way the seed responds when it is dried. 

One parameter is the lowest-safe moisture content 

(LSMC), defined as the value below which viability is 

immediately lost on drying. The value for this parameter 

provides a guide to the moisture content below which 

seed should not be held during handling procedures. 

LSMC values were assessed under standard drying 

conditions and were found to vary between 26% and 50% 

(Table 3). In Table 4 further LSMC data are given; 

although these were assessed using various desiccation 

methods, the results are in broad agreement with those 

in Table 3. 

Slope and intercept parameters are presented for 

some species (Table 3); these define the relationship 

between germination and moisture content during 

desiccation. 

Desiccation rates 

It is possible that desiccation rate may influence viability; 

for example, seeds dried quickly might give lower 

germination than seeds dried more slowly and gently to 

the same moisture content. However, in the case of the 

‘recalcitrant’ seed of Araucaria hunsteinii 

(Araucariaceae) no such differences were observed 

(Tompsett 1982). No intensive study of this sort has been 

carried out on dipterocarp seed. However, 

Amata-Archachai and Hellum (unpublished) found that 

immature fruits of Dipterocarpus alatus clearly dried 

quicker than mature fruits; they suggested that the 

difference could be because of the death on drying of 

the immature seeds. However, the faster loss of moisture 

by immature seeds could also be explained by their 

smaller size. Small seeds have a higher ratio of surface 

area to volume than large seeds, enabling quicker 

moisture loss. In this connection, Tamari (1976) found 

small seeds of S. parvifolia (0.3 g) gave low viability, 

whilst large seeds (0.5 g) gave higher viability. One 

explanation for the latter finding is that the smaller seeds 

had dried quicker and thus lost more viability than larger 

seeds prior to testing. 

Clear-cut differences in desiccation rates among 

seeds of species in the same genus have been reported 

by Tompsett (1986, 1987); rates for Dipterocarpus 

seeds varied greatly and depended on their size and 

structure. At the one extreme D. intricatus seed required 

only one week to dry to 7% moisture content; at the other 

extreme, seed of D. obtusifolius under identical 

conditions retained c. 30% moisture content even after 

5 weeks. Likewise, Yap (1986) found S. parvifolia seeds 

dried quicker than those of two larger-seeded species 

of Shorea; he believed the difference in rates to be 

related to pericarp thickness. 

Differences in desiccation rates of the type discussed 

above may possibly affect both the initial 

post-desiccation viability and the subsequent storage life 

of the seed. Further studies are needed to assess these 

effects.


Table 3. Relationship between germination and moisture content during desiccation (Tompsett and Kemp 1996a, b)*. 

Species Lowest-safe moisture 

content values 

(percentage)** 

*: Results can be summarised by regression as a straight line if 

germination percentage is first transformed into probits; 

The basis of desiccation damage 

If the basic causes of desiccation damage could be 

determined, a way might be found to reduce the effect, 

thus enabling better survival of the seed. In this 

connection, Nautiyal and Purohit (1985b, c) assessed 

various factors for S. robusta seed. The quantity of 

nutrients leaking from the seed increased as moisture 

content (and germination ability) declined; it was 

Slope of probit line (probits 

per unit of moisture content 

percentage) 

Intercept of probit line 

(probit percentage 

germination) 

Shorea leprosula 26 n/a n/a 

Shorea argentifolia 28 0.1814 -3.5780 

Shorea ferruginea 29 0.1912 -4.8300 

Hopea ferrea 30 0.1050 -3.5660 

Hopea mengerawan 30 0.1400 -3.6700 

Hopea foxworthyi 31 0.0994 -2.5555 

Hopea odorata 32 0.1303 -3.6450 

Shorea parvifolia 32 n/a n/a 

Shorea roxburghii 32 0.1000 -2.7700 

Shorea obtusa 33 0.0660 -2.4420 

Shorea ovalis 33 0.1816 -5.0550 

Cotylelobium melanoxylon 34 0.1303 -3.1790 

Vatica mangachapoi 34 0.0919 -3.4460 

Cotylelobium burckii 35 0.1400 -3.2200 

Parashorea smythiesii 35 0.1743 -4.9920 

Shorea macrophylla 35 0.0978 -2.6190 

Shorea trapezifolia 37 n/a n/a 

Dipterocarpus costatus 38 0.0789 -1.9360 

Dipterocarpus obtusifolius 38 0.0584 -2.2470 

Dipterocarpus zeylanicus 38 0.1427 -3.8090 

Shorea fallax 38 0.0869 -2.2470 

Shorea macroptera 38 0.0867 -4.2300 

Parashorea tomentella 40 0.2000 -7.1400 

Shorea amplexicaulis 40 0.1307 -5.5600 

Shorea congestiflora 40 0.0904 -3.6510 

Shorea robusta 40 0.1300 -4.2200 

Vatica odorata ssp. odorata 41 0.0961 -3.6290 

Shorea affinis 42 0.0475 -1.6120 

Shorea almon 42 0.1400 -5.4700 

Shorea leptoderma 42 0.0460 -2.5280 

Dipterocarpus turbinatus 43 0.1300 -5.5300 

Dryobalanops lanceolata 43 0.1200 -6.0400 

Dryobalanops keithii 50 0.1233 -5.0770 

Parashorea malaanonan 50 0.0965 -3.9830 

**: LSMC for seeds dried at 10-15% relative humidity and 15- 

20°C. 

concluded that cellular membranes in the seed had lost 

their semi-permeability. However, whether the apparent 

loss of semi-permeability was a primary result of 

desiccation, or whether it was one aspect of a general 

loss of metabolic capability could not be distinguished 

from the data obtained. A small decline in the absolute 

concentration of nutrients in the seed was observed, but 

the significance of this decline was not clear. Protein


Table 4. Lowest-safe moisture content values (wet weight basis) for mature seeds*. 

Species Source LSMC (%) 

Dipterocarpus alatus** Tompsett (unpub.) 25 

Shorea siamensis Tompsett (unpub.) 51*** 

Shorea singkawang Yap (1986) 55 

Shorea xanthophylla Tompsett (unpub.) >41*** 

Stemonoporus canaliculatus Tompsett (unpub.) 43*** 

Vatica umbonata Mahdi (1987) 74**** 

*: no slopes and intercepts available for these species; 

**: seed is OLDA (orthodox with limited desiccation ability); 

changes accompanying loss of viability of S. robusta 

have also been reported (Nautiyal et al. 1985). 

Some authors have confused the effects of 

desiccation itself with the effects of ageing; in order to 

determine the effects of ageing, moisture contents 

should be kept constant. However, in studies by Song et 

al. (1983) on Hopea hainanensis it is clear that 

desiccation effects per se were being examined. At 36% 

moisture content the ultrastructure was intact, but on 

desiccation to 26% moisture content, which severely 

reduces germination percentage, various changes were 

observed. Vesicles appeared in the cytoplasm, vacuolar 

membranes ruptured and cell contents became less 

distinct. Cell walls and cytoplasm became separated and 

nuclear membranes could not be distinguished from the 

nucleolus. These changes illustrate a general 

deterioration of cellular structure rather than an effect 

confined to the cell membrane. In a further study (Song 

et al. 1986), desiccation to 31% was shown to disturb 

the ribosomes and endoplasmic reticulum, but these 

changes were reversed on re-hydration. 

More recently, Krishan Chaitanya and Naithani 

(1994) measured changes in superoxide, lipid 

***: based on at least 25 seeds per germination; 

****: unusually high value. 

peroxidation and superoxide dismutase for seeds of S. 

robusta during desiccation. They concluded that the loss 

of viability observed may be caused by the cumulative 

effect of peroxidation products of polyunsaturated fatty 

acids and peroxidation of the membrane lipids. 

Storage Physiology 

Some aspects of seed storage are considered elsewhere: 

practical aspects, including the effects of gases, are 

discussed in Chapter 4; chilling physiology is considered 

above under germination effects. Topics discussed below 

include the following: best recorded storage periods; use 

of viability constants; the significance of oil contents; 

some aspects of tissue culture; and various associations 

with storage physiology. 

Best storage records 

An up-to-date summary of best storage records is given 

in Table 5. These records should not be confused with 

practical recommendations; if the recommended storage 

conditions were employed, longer storage would be 

expected in many cases. The best record for an OLDA 

species is 2829 days for Dipterocarpus alatus and the


Table 5. Temperatures, moisture contents and germination of mature seeds for the optimum reported storage conditions. 

Species 

Germination 

(%) 

Optimum storage achieved 

Days Temp. 

(°C) 

MC 

(%) 

Other conditions Source 

Anisoptera costata 44 30 18 44 ventilated incubator, 99% 

rh, rib-channel PB, 

ventilated weekly 

Anisoptera marginata 45 84 21 48 ventilated incubator, 99% 

rh, PB, ventilated weekly 

Cotylelobium burckii 52 28 21 29 gas box, over water, 


Cotylelobium 

46 67 21 36 gas box, over water, 

melanoxylon 


Dipterocarpus alatus** 44 2829 -13 11 hermetic, laminated 

aluminium foil bag 

Tompsett 

(unpub.)* 

Dipterocarpus baudii 25 30 14 n/a n/a 

Tompsett 

(unpub.)* 

Tompsett 

(unpub.)* 

Tompsett 

(unpub.)* 

Tompsett 

(unpub.)* 

Yap (1981) 

Dipterocarpus 

87 4 n/a 40 No information. Tompsett 

grandiflorus 

(unpub.)* 

Dipterocarpus 

30 28 15 26 PB, sealed, inflated with Maury-Lechon 

humeratus 

nitrogen 

et al. (1981) 

Dipterocarpus 

30 2373 -20 10 hermetic, laminated Tompsett 

intricatus** 

aluminium foil bag (unpub.)* 

Dipterocarpus 

20 60 18 59 ventilated incubator 99% Tompsett 

obtusifolius 

rh, rib-channel PB, 


(unpub.)* 

Dipterocarpus 

77 1369 -20 12 hermetic, laminated Tompsett 

tuberculatus** 

aluminium foil bag (unpub.)* 

Dipterocarpus 

20 177 16 42 closed box, over water, Tompsett 

turbinatus 

ventilated weekly (unpub.)* 

Dipterocarpus 

53 100 21 39 ventilated incubator 99% Tompsett 

zeylanicus 

rh, loose 

(unpub.)* 

Dryobalanops 

aromatica 

50 16 14 38-40 n/a Yap (1981) 

Dryobalanops keithii 54 23 16 45 PB tied, sawd. (18% Tompsett 

MC), ventilated weekly (unpub.)* 

Dryobalanops 

92 62 21 56 PB sealed and inflated, Tompsett 

lanceolata 


Hopea ferrea 40 300 16 30-50 PB then perl. Tompsett 

(unpub.)* 

Hopea foxworthyi 68 365 18 35 ventilated incubator at Tompsett 

99% rh, rib-channel PB, 


(unpub.)* 

Hopea hainanensis 80 365 18 35-38 n/a Song et al. 

(1984, 1986) 

Hopea helferi 85 40 15 48 n/a Tang and 

Tamari (1973) 

Hopea mengerawan 40 67 21 44 ventilated incubator at Tompsett 

99% r.h., loose 

(unpub.)* 

Hopea nervosa 19 330 25 n/a n/a Sasaki (1980) 

Hopea odorata 48 93 16 38 polythene rib-channel Tompsett 

bag within polythene box, 


(unpub.)*


Table 5. (continued) Temperatures, moisture contents and germination of mature seeds for the optimum reported storage 

conditions. 

Species 

Germination 

(%) 


Days Temp. 

(°C) 

MC 

(%) 

Other conditions 

Source 

Hopea parviflora 84 104 18 41 PB sealed and inflated, Tompsett 


Hopea subalata 40 51 4 32-43 n/a Sasaki (1980) 

Hopea wightiana 5 60 4 n/a n/a Sasaki (1980) 

Monotes kerstingii 16 90 2 7 PB, ventilated weekly Tompsett 

(unpub.)* 


heimii 

80 50 14 28-47 n/a Yap (1981) 

Parashorea densiflora 90 60 25 54 n/a Yap (1981) 

Parashorea 

67 141 18 45 rib-channel PB, 

Tompsett 

malaanonan 


Parashorea smythiesii 50 317 18 44 ventilated incubator 99% 

rh, PB, perl. (0% MC), 


Parashorea tomentella 40 91 16 40 PB, 4% moisture content 

perl., ventilated weekly 

(unpub.)* 

Tompsett 

(unpub.)* 

Tompsett 

(unpub.)* 

Shorea acuminata 70 30 21 38-43 n/a Sasaki (1980) 

Shorea affinis 56 253 21 35 ventilated incubator 99% Tompsett 

rh, loose 

(unpub.)* 

Shorea almon 18 32 16 45 stored on agar, some Tompsett 

seed germinated and are 

included 

(1985) 

Shorea amplexicaulis 30 168 21 45 PB, sealed, perl. 0-8% Tompsett 

MC, ventilated weekly (unpub.)* 

Shorea argentifolia 60 45 21 43 No information. Sasaki (1980) 

Shorea assamica 50 98 4 n/a n/a Sasaki (1980) 

Shorea bracteolata 4 60 4 n/a n/a Sasaki (1980) 

Shorea congestiflora 52 49 21 39 PB, top folded over, Tompsett 

within gas box, ventilated 

weekly 

(unpub.)* 

Shorea contorta 35 32 21 67 ventilated incubator 98% Tompsett 

rh, PB, ventilated weekly (unpub.)* 

Shorea curtisii 20 30 25 n/a n/a Yap (1981) 

Shorea dasyphylla 24 14 21 40-46 n/a Sasaki (1980) 

Shorea fallax 50 50 21 40 PB tied, with sawd., Tompsett 


Shorea ferruginea 36 77 21 34 PB, sealed and inflated, Tompsett 


Shorea hypochra 10 60 4 n/a n/a Sasaki (1980) 

Shorea javanica 87 30 20 13-15 moisture low Umboh 

(1987) 

Shorea leprosula 45 30 21 32 No information. Sasaki (1980) 

Shorea macrophylla 40 22 21 31 PB tied, with sawd., Tompsett 


Shorea obtusa 44 11 16 36 PB, sealed and inflated, Tompsett 


Shorea ovalis 87 92 21 37 No information. Sasaki (1980) 

Shorea pachyphylla 28 16 18 66 ventilated incubator 99% Tompsett 

rh, over water 

(unpub.)*


Table 5. (continued) Temperatures, moisture contents and germination of mature seeds for the optimum reported 

storage conditions. 

Species 


Source 

Germination Days Temp. MC Other conditions 

(%) 

(°C) (%) 

Shorea parvifolia 40 57 18 35 ventilated incubator Tompsett 

99% rh, PB, with perl., 


(unpub.)* 

Shorea pauciflora 67 45 25 38-51 n/a Sasaki (1980) 

Shorea pinanga 50 112 21 46 ventilated incubator Tompsett 

99% rh, loose 

(unpub.)* 

Shorea platyclados 80 58 25 n/a n/a Yap (1981) 

Shorea robusta 54 49 15 0 partial vacuum, MC Khare et al. 

unknown 

(1987) 

Shorea roxburghii 52 307 16 36 PB, tied and inflated, Tompsett 

ventilated weekly (1985) 

Shorea siamensis 83 56 15 40-48 No information. Panochit et 

al. (1984) 

Shorea smithiana 28 46 26 44 PB sealed and inflated, Tompsett 


Shorea sumatrana 60 15 25 n/a n/a Yap (1986) 

Shorea trapezifolia 100 63 21 n/a ventilated incubator Tompsett 

97% rh, PB, ventilated 

weekly, seed was pregerminated. 

(unpub.)* 

Stemonoporus 

20 77 18-21 47 ventilated incubator Tompsett 

canaliculatus 

99% rh, loose 

(unpub.)* 

Vatica mangachapoi 24 85 21 40-45 PB, perl. 0% MC, Tompsett 


Vatica odorata ssp. 

48 148 18 40 ventilated incubator Tompsett 

odorata 

99% rh, loose 

(unpub.)* 

Vatica umbonata 10-50 60 10-15 n/a n/a Mori (1979) 

*: data based on at least 25 seeds per germination; 

**: seed has OLDA storage physiology (orthodox with limited 

desiccation ability); 

MC: moisture content based on wet weight; 

corresponding value for a recalcitrant species is 365 

days for Hopea hainanensis. Five recalcitrant species 

were stored for over 300 days and a further eight were 

stored for over 100 days (Table 5). The three OLDA 

species all stored for over 1300 days. 

Viability constants 

A brief background to viability constants (meaning and 

derivation) is given. 

The rate at which orthodox and OLDA seeds age in 

storage increases with temperature and with moisture 

contents between certain limits. Successive samples 

from a seedlot of high initial viability in storage will 

show progressively lower germination percentages, 

perl.: stored in perlite; 

sawd.: stored in sawdust; 

PB: stored in ventilated polythene bag. 

Note: not all reports specified the seed maturity. 

producing a curve of germination against time which is 

sigmoid in shape. This curve can be transformed by 

probit analysis to produce a straight line relationship. 

Results from many storage treatments (various moisture 

content and temperature combinations) can be analysed 

to give constants in a predictive equation. The equation 

was developed by Ellis and Roberts (1980a, b) for 

herbaceous species and is: 

V = Ki - P/10K E - C W log 10 m - C H t - C Q t2 ............. (Eqn 1). 

In this equation, V is the predicted viability, K i is the 

initial viability, P is the number of days in storage, m is 

the moisture content (percentage fresh weight basis) and 

t is the temperature ( o C). Viability is expressed as probit 

germination. The constants K E , C W , C H and C Q are


Table 6. Viability constants and standard errors for two OLDA species of dipterocarps (Tompsett and Kemp 

1996a, b). 

Species KE (se) CW (se) CH (se) CQ (se) 

Dipterocarpus 

alatus 

Dipterocarpus 

intricatus 

6.44 (0.72) 3.09 (0.61) 0.0329 (0.0017) 0.000478 (0.000000) 

6.34 (0.81) 2.70 (0.68) 0.0329 (0.0017) 0.000478 (0.000000) 

common to all seedlots of a species. The equation has 

been shown to apply to tropical and temperate tree 

species (Tompsett 1986). 

Equation 1 was derived using the following equation: 

log10 s = K - C log m - C t - C t2 

E W 10 H Q ............. (Eqn 2). 

In this equation, s represents the rate of loss of viability 

in days per probit. 

The viability constants K E , C W , C H and C Q of Eqn 2 

are reported for two dipterocarp species in Table 6. 

Ageing was observed under at least 4 temperature 

conditions and with several moisture content treatments 

at each temperature in order to obtain the parameters 

presented for these species. Further details of method 

are in Tompsett and Kemp (1996a, b). 

The constants for the two Dipterocarpus species were 

similar and can be used to predict viability at the end of 

any storage period when moisture content and 

temperature are known. Thus, for D. alatus, a 64-year 

period is predicted before seed ages to 85% germination, 

provided the initial viability of the seed is 99.4% and 

storage is at -13°C with 7% moisture content. 

Calculations should be based on sound seed only. This 

approach enables decisions to be made about the 

cheapest conditions commensurate with attaining the 

objectives of storage for different purposes. The 7% 

moisture content value for D. alatus was chosen, in part, 

because it has proved difficult to dry the seed further. 

Oil content of the seed 

Details of embryo oil contents for dipterocarps are given 

in Table 2 and show much lower values for Hopea and 

Dipterocarpus than for Shorea. 

In the predictive viability equation given above, the 

water status of seed was assessed using moisture content. 

However, a more accurate measure of seed water status 

in relation to physiological activity is seed water 

potential. Water potential is in turn related to the relative 

humidity which produces, at equilibrium, the moisture 

content under consideration. These relationships have 

been considered in connection with storage life by 

Roberts and Ellis (1989). The reason why relative 

humidity is of importance may be illustrated by 

considering the influence on longevity of the reserves 

in an oily seed. For a species with an oil content of 50%, 

ageing-associated physiological responses would be 

predicted at a moisture content which is about half the 

moisture content for the same responses in a non-oily 

seed, provided all other factors are identical. This is 

because of the hydrophobic nature of the oily reserve. 

The relative humidity value at equilibrium for the same 

physiological responses, however, would be expected to 

be similar for both species. Since seeds of 

Dipterocarpus alatus are not oily, it is not surprising 

that optimum longevity is at a relatively high moisture 

content near 7%. By contrast, the oily seed of Swietenia 

humilis (Meliaceae) is best stored at near 3% moisture 

content. 

Tissue Culture 

Tissue culture has been suggested as a means of storage 

of gene resources under slow growth conditions. 

Additionally, this technique can be employed for 

micropropagation. It is also likely that tissue culture 

would be needed to grow the resulting tissue after 

cryopreservation if the latter method proves practical. 

However, tissue culture of dipterocarps is not easy, high 

rates of cell necrosis having been observed for some 

species. High resin content within the tissues may be at 

least partly responsible for this effect for some species. 

However, some success has been achieved by Smits and 

Struycken (1983), Scott et al. (1988) and Linington 

(1991) in culturing the tissues of some Shorea and 

Dipterocarpus species. 

Association of storage physiology with seed 

characters and tree habitat 

Various associations have been noted for dipterocarp 

seeds. The LSMC, defined as the moisture content below


which some germination loss occurs on desiccation, is 

associated with various properties of the seed and its 

parent tree. Seed size, seed desiccation rate, seed 

longevity and the habitat of the parent species have all 

been found to be related to storage physiology. 

Storage physiology and seed size 

For three Shorea species a relationship has been noted 

between seed size and desiccation tolerance; lowest-safe 

moisture content values increase as size increases from 

the small, desiccation-tolerant seed of S. roxburghii to 

the larger, desiccation-intolerant seed of S. almon 

(Tompsett 1985). A similar relationship was found in the 

Dipterocarpus genus (Tompsett 1987), but in this case 

it is the size of the embryo that appears more important. 

Thus, two relatively small-embryoed species (D. 

intricatus and D. tuberculatus) were shown to be OLDA 

in their storage physiology (and can therefore be dried 

with relatively little damage); on the other hand, two 

species with large embryos (D. obtusifolius and D. 

turbinatus) were shown to have high LSMC values and 

recalcitrant physiology. There are other species that fit 

this pattern (Tompsett 1986). 

A further association which has been observed is that 

recalcitrant seeds tend to be smooth surfaced (globular), 

whilst OLDA seeds have tubercles or other projections 

from the calyx. These projections may enhance 

desiccation rate, leading to better storage on the forest 

floor (Tompsett 1987), as explained below. 

Storage physiology in relation to habitat and longevity 

Seeds of three recalcitrant Shorea species from 

different habitats have been found to have different 

desiccation tolerances. The low-rainfall area species S. 

roxburghii has seed which can be dried safely down to 

35%, whereas the two monsoon or rain forest species S. 

almon and S. robusta cannot be safely dried below 40% 

moisture content (Tompsett 1985). Interestingly, the seed 

with the greatest desiccation tolerance (S. roxburghii) 

is also the seed with the greatest longevity. 

A more extreme example is found in the genus 

Dipterocarpus. Two dry-zone, deciduous species (D. 

intricatus and D. tuberculatus) have OLDA-physiology 

seeds, whilst two other species with distributions 

extending into the relatively wet, evergreen areas (D. 

turbinatus and D. obtusifolius) have recalcitrant seeds 

(Tompsett 1987). The longevity of dry OLDA seeds is 

relatively great (Table 5), whilst recalcitrant seeds cannot 

be stored in the long term at present. As with other 

factors, these patterns have been found to extend to seeds 

of other species; trees from low-rainfall and sandy-soiled 

areas tend to have greater longevity and lower LSMC 

values (Tompsett 1986). 

Storage physiology in relation to seed desiccation 

rate 

The OLDA seeds of Dipterocarpus intricatus and D. 

tuberculatus can dry to below 10% in 2 weeks, whereas 

the recalcitrant seed of D. obtusifolius remains above 

28% moisture content even after 5 weeks in the same 

drying conditions (Tompsett 1987). This situation may 

have evolved because OLDA species benefit from 

desiccation in terms of enhanced storage life as follows. 

If the wet season arrives late, so that the seeds lie on the 

ground for several weeks, viability is nonetheless 

preserved by their low moisture content under natural 

conditions. Conversely, the slow desiccation rate 

characteristic of recalcitrant seeds is protective against 

desiccation damage. The differences in desiccation rates 

observed are generally associated with seed size (small 

seeds dry faster) and probably also to seed anatomy. 

Induction of Flowering and Seeding 

Little work has been done on the artificial induction of 

flowering and seeding. However, Tompsett, 

Tangmitcharoen, Ngamkhajornwiwat and 

Sornsathapornkul (unpublished) have found a positive 

effect of the growth inhibitor paclobutrazol in promoting 

the flowering of Dipterocarpus intricatus in north-east 

Thailand. The best effect was found by applying the 

substance at 20 g/l to buds between late September and 

early November. The ability to control flowering would 

aid breeding programmes and may enhance seed 

production in years when it is otherwise poor. 

Future Research 

More work is needed to assess the seed storage 

physiology categories of dipterocarp species, exploring 

desiccation tolerance to assess whether the currently 

known species with OLDA seed are the only ones in 

existence. There are currently three such species known. 

A broad range of species should be included to enable a 

steady flow of material, despite the infrequent fruiting 

and the logistical problems of locating, collecting and 

transporting materials.


Dipterocarp seed of the OLDA type has a shorter 

storage life than seed of crop species if compared at the 

same moisture content. Thus, it has been estimated that 

the relevant K E and, C W viability constants (which 

indicate seed longevity) are, respectively, only 6.4 and 

2.9 on the average for dipterocarp seed (Table 6), 

compared with 8.4 and 4.7 (Tompsett 1994) for 

herbaceous crops. Further research is needed to extend 

these findings to other OLDA dipterocarp species. 

The stage of fruit development at harvest is important 

to ensure optimum desiccation tolerance, and 

consequently to ensure maximum storage potential. 

Further research is needed for dipterocarps in order to 

closely assess the relationship between harvest 

condition, postharvest handling, and desiccation 

tolerance. 

Studies are needed to increase knowledge of the 

optimum moisture and temperature conditions for 

storage of recalcitrant seeds, employing controlled 

conditions. Especially, research is needed in relation to 

the chilling injury. Studies to quantify chilling damage 

in relation to moisture content are needed. Also, research 

is required to determine its relationship to underlying 

biochemical processes. 

Although the database DABATTS (Tompsett and 

Kemp 1996a, b) includes a large amount of previously 

unpublished information on dipterocarp seed, a high 

proportion of its contents are the results produced by 

the authors. Unpublished information from other sources 

needs to be databased, building on DABATTS and 

increasing the total sum of research results readily 

available. 

Research on the induction of flowering is necessary 

to improve knowledge of the causes underlying the 

irregular flowering of dipterocarps. Such research may 

provide artificial means for the induction of flowering 

in relation to breeding and to seed production in nonmast 

years. 

These approaches might with benefit be extended to 

other tropical tree families such as the Palmae and 

Sapotaceae. 

Relevant Institutions 

As described in Chapter 4 and above (in the work of 

Sasaki, Mori, Tang, Tamari and Yap), Forest Research 

Institute Malaysia (FRIM) has played a leading role in 

early dipterocarp seed research, particularly in the areas 

of germination ecology and storage research. Current 

work at FRIM on cryopreservation and seedling storage 

is referred to elsewhere. Over the last decade, the seed 

physiological studies at the Royal Botanic Gardens Kew 

have contributed basic knowledge, creating a firm 

foundation for practical recommendations. The Forest 

Research Centre, Sandakan, Malaysia, has a seed research 

laboratory constructed under an FAO aid programme and 

has undertaken significant dipterocarp research. 

In Thailand the Royal Forest Department’s ASEAN 

Tree Seed Centre, Muak Lek, has been involved in 

dipterocarp studies for a number of years and has good 

facilities; additionally, the central laboratory in Bangkok 

has an active research team on the topic. 

The Ecosystems Research and Development Bureau, 

Republic of the Philippines, is engaged in dipterocarp 

seed research, as are the Forest Research and 

Development Centre, the Biotechnology Centre and 

BIOTROP in Bogor, Indonesia. In India, research on 

biochemical aspects has been recently conducted at the 

High Altitude Plant Physiology Research Centre of 

Garhwal University, Srinagar and the Forest Research 

Institute, Dehra Dun has been involved in dipterocarp 

research in the recent past. 

In China, biochemical, ultrastuctural and 

physiological research on dipterocarp species has been 

performed by staff of the Tropical Forest Research 

Institute, Chinese Academy of Forestry, Guangdong. 

Although not involved in dipterocarp research, the 

Agriculture and Horticulture Department at Reading 

University, UK, is developing experience in the area of 

tropical tree seed physiology. Other institutes have 

contributed information in this field, but space available 

limits the numbers that can be included. 


I thank the Royal Botanic Gardens Kew and the Center 

for International Forestry Research for facilities and 

financial support 

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Seed Handling 

B. Krishnapillay and P.B. Tompsett 

In considering seed handling, it is important to be aware 

of the sources of seed quality. Many benefits flow from 

the use of better quality seeds, selected and handled 

optimally; advantages include the improved survival of 

seedlings and greater overall commercial returns. 

However, methods to ensure high quality of seed 

supply are not as advanced for dipterocarps as for other 

forest species such as pines and eucalypts. The primary 

problem is seed supply and this factor is a major 

constraint in dipterocarp forest management. Thus, the 

lack of seeds in sufficient quantity and quality has 

discouraged the raising of seedlings in the nursery and 

direct sowing of seeds in the field. 

The majority of dipterocarps do not flower regularly. 

In the aseasonal zones flowering occurs at intervals of 

two to five years and its accurate prediction is impossible. 

Consequently, it is difficult to plan major planting 

activities. Even in flowering years, interference by 

drought can cause premature fruit drop. On the other 

hand, flowering is generally on an annual basis in the 

seasonal climatic zones so that planning seed collection 

in these areas is easier. Although seed production can 

vary between years in any particular place, foresters can 

make more secure plans by widening the area monitored 

for seed supply. 

A second problem in practice is the life span of 

dipterocarp fruits; most species have short-lived 

‘recalcitrant’ seed. If seed collectors do not harvest 

mature seed and sow it immediately, a proportion will 

soon become inviable. A few species, however, have longlived 

seed. Early descriptions of the short-lived nature 

of dipterocarp seeds include those of Troup (1921), Sen 

Gupta (1939) and Dent (1948). The period between 

collection and sowing should thus generally be as short 

as possible. In practice, reports of fruiting are often 

received at short notice; thus, in order to produce 

dipterocarp seedlings, a collection team has to be hastily 

prepared for collection, transport and sowing in the 

nursery. Few agencies can liaise these activities 

efficiently. Schaffalitzky de Muckadell and Malim 

(1983) considered some relevant factors. 

Chapter 4 

In seasonal forests, on the other hand, the scope for 

forestry operations with dipterocarp species is wider, 

flowering being more regular and seed being longer-lived. 

Even in this climatic zone, however, most species are 

recalcitrant. Much work has been carried out on the factors 

controlling the longevity of dipterocarp seeds (see 

Chapter 3). Researchers have achieved success for 

species from both seasonal and aseasonal zones but have 

made relatively more progress with species that do not 

possess recalcitrant seed. Alternative means of raising 

planting material have been investigated as a 

complementary approach. 

Several handbooks have been produced on the 

handling of tropical tree seed, a notable example being 

that of Willan (1985). In this chapter, wider aspects of 

seed handling, including biology and ontogeny, are 

described. In addition, seed collection, seed storage, 

seedling storage and cryopreservation are covered and 

future research priorities and prospects for successful 

forest seed programmes are considered. 

Factors Affecting Seed Viability 

When seeds (more correctly fruits) reach maturity on 

the mother plant, they begin to deteriorate; the rate of 

deterioration depends on the environmental conditions 

they experience. Progressively, germination rate is 

reduced, the number of abnormal seedlings is increased 

and field emergence is lowered. Cumulative damage 

occurs until the seed is incapable of germinating. 

Preferably, foresters should use seed before its viability 

has dropped significantly. Various factors operating 

before seeds arrive at the seed centre can influence initial 

germination percentage. These factors in relation to seed 

handling considerations are summarised below and more 

detail is given in Chapter 3. 

The effect of climate and pest infestation 

Climatic conditions prior to seed harvest and the 

physiological state of the mother tree may influence

Seed Handling 74 

viability of the seed but experimental proof is lacking. 

In some years there are heavy infestations of the 

developing seed by pests and insects. It is possible that 

heavy infestations occur relatively more frequently in 

years when there are light crops on the tree but 

confirmation of this relationship is needed. 

Maturity 

Seed germination continues to improve up to near the 

time of peak maturity, emphasising the need for optimal 

harvest timing. 

Physiological and other associated damage 

During the period between collection and arrival at the 

seed centre, material is at risk. This applies particularly 

if seed is held under conditions that are either too humid 

or too dry, and if temperatures are too high or too low. 

Necrosis is liable to occur under such conditions, 

associated with fungal growth and viability loss. 

Seed Storage Categories 

Researchers have divided seeds broadly into 3 major 

groups on the basis of their storage behaviour. The 

following descriptions give the general basis for each 

type; more accurate definitions are presented in Chapter 

3 (pages 60-61). 

Orthodox seeds 

This category includes seeds that can be dried to low 

moisture contents (about 5%) without serious 

deleterious effects. Under optimal conditions, the life 

span of this group of seeds can be extended for decades 

or longer. 

Recalcitrant seeds 

This group of seeds differs from orthodox seeds in two 

ways; their seeds die if they are dried below relatively 

high moisture contents (values are given for lowest-safe 

moisture contents in Chapter 3, page 62) and if they are 

subject to damage at low temperatures (< 16 o C). Even 

under optimal conditions survival of seeds in this group 

is limited. The difficulties in storing the seed led to their 

being described as ‘recalcitrant’. 

Intermediate (OLDA) seeds 

A third category of seed storage physiology has been 

recently defined. In practice, the seeds in this group have 

desiccation characteristics that are intermediate between 

those of the orthodox and recalcitrant seeds and they 

have thus been termed ‘intermediate’. When harvested 

in the usual way, seeds of this type can be dried to 

moisture levels of about 8-12% whilst retaining a 

substantial amount of (but not all) their original viability. 

There is also a greater susceptibility to chilling and 

freezing damage than is the case with orthodox seed, even 

when the seeds are relatively dry. 

When this type of seed was first studied in detail, its 

physiological similarity to orthodox seeds led to the 

description ‘orthodox with limited desiccation ability’ 

(OLDA). However, employing the term ‘intermediate’ 

to indicate a practical difference from orthodox seeds 

is useful. This matter is further discussed in the Seed 

Physiology chapter. 

Tropical Forest Tree Seeds 

Tompsett (1994) has estimated that 72% of tree species 

found in the tropics may bear ‘recalcitrant’ seeds. 

Recalcitrant seeds are shed from the mother plant with 

very high moisture contents (about 40-60% on a wet 

weight basis) and germinate soon after shedding. Whilst 

recalcitrant dipterocarp species provide real problems, 

those of the OLDA type are more amenable, as described 

above. Tompsett (1994) found that, in the case of 

dipterocarp species, 94% of those examined possessed 

recalcitrant seed. 

Seed Ontogeny 

Ontogeny covers development from floral initiation 

through growth and differentiation to maturity of the 

seed. To date, very little work has been published on the 

ontogeny of dipterocarp species; Owens et al. (1991) 

presented a generalised, basic development diagram 

which may relate to certain species of the dry forest in 

Thailand. 

Phenology 

Phenology, in a broad sense, refers to the relationship 

between changes in seasons and climate and to the 

phenomena of leaf and bud formation, leaf fall, floral 

anthesis, fruit set and ripening. In the aseasonal 

dipterocarp forests from south Asia to Malesia 

phenological observations are an essential part of the 

strategy for seed procurement of dipterocarps, owing to 

the irregularity of their flowering and fruiting patterns.


Table 1. Likely periods for flowering and seed production of important Dryobalanops, Dipterocarpus, Shorea, and Anisoptera 

species (Krishnapillay, unpublished). 

Species 

Over the last 25 years various authors have reported 

detailed phenological records. Studies include those of 

Burgess (1972), Cockburn (1975) and Ng (1981, 1984) 

for the Malaysian aseasonal forest, and Sukwong et al. 

(1975) for the dry forest of Thailand. In Table 1, there is 

a general summary for the important timber species of 

Peninsular Malaysia. 

The infrequency and irregularity of dipterocarp 

flowering and fruiting in the aseasonal areas have already 

been referred to above. A further feature is that flowering 

tends to be gregarious and may be limited or may extend 

throughout an entire region. 

Flower and seed surveys indicate: 

1. whether flowering is scattered and confined to 

particular species or whether it is a mast flowering; 

2. whether the amount of seeds available is sufficient to 

meet seed collection requirements; 

3. whether the crop is sound or has been attacked by 

pests or insects; and 

4. the time when the seeds will mature. 

The natural trigger for mast flowering and fruiting 

among dipterocarps has been sought by looking for 

associations with several factors. Foxworthy (1932) and 

Months 

J F M A M J J A S O N D 

many others suggested an association between flowering 

and strong droughts but Wood (1956) disputed the 

conclusion. Ng (1981) suggested that a dry spell 

preceding leaf flush accompanied by a rising gradient of 

daily sunshine induces flowering. Again, Ashton et al. 

(1988) proposed that the environmental trigger is a 

protracted low night temperature over a period of about 

3-4 days. However, experimental evidence is required 

to establish cause and effect. The matter is further 

discussed in the Seed Physiology chapter. 

Seed Procurement 

Frequency 

Dryobalanops aromatica x x x x x x x x x x x biennial 

Dryobalanops oblongifolia x x x x x x x x x x biennial 

Shorea leprosula x x x x x x x 3-4 years 

Shorea parvifolia x x x x x x x 3-4 years 

Dipterocarpus baudii x x x x x x x annual 

Dipterocarpus costulatus x x x x x x x 4-5 years 

Anisoptera scaphula x x x x 4-5 years 

Anisoptera laevis x x x x 4-5 years 

Dipterocarpus kerrii x x x x 4-5 years 

Shorea macrophylla x x x x x x x x x x x x annual 

Shorea macroptera x x x x x 3-4 years 

Shorea ovalis x x x x x x x 3-4 years 

Shorea platyclados x x x x x 3-4 years 

Shorea acuminata x x x x x x x 2-3 years 

Shorea bracteolata x x x x x x x 2-3 years 

Shorea curtisii x x x x x 3-4 years 

Current research on artificial regeneration has been 

reviewed by Mok (1994), whilst Barnard (1950) and 

Appanah and Weinland (1993) outline some procedures 

that have been used to procure dipterocarp seeds for 

planting programmes. A more detailed procurement 

procedure is needed. At present, most methods involve 

collection of seeds on an ad hoc basis or the collection 

of wildings. Seed procurement should involve planning, 

collection, transporting, processing, testing, temporary 

storage and nursery facilities. A general description of


the basic activities involved in seed procurement is given 

below. If a large collection region is monitored, some 

seeding may be found every year; in practice, however, 

logistical and other problems make annual collection from 

aseasonal regions difficult. 

Planning 

When trees start fruiting, procurement planning has to 

be initiated immediately so that good-quality planting 

material can be obtained. The period between collection 

and storage or sowing should be as short as possible to 

reduce the chance of seed deterioration. Transport and 

processing should be carefully planned and, when 

necessary, the nursery advised so that germination space 

is available. 

Collection 

The choice of collection technique for forest tree seed is 

dependent on many factors, including the way the tree 

disperses its seeds or fruits. For recalcitrant-seeded 

dipterocarp species collecting seeds directly from the tree 

crown by climbing has several advantages. These are: 

a) mature seeds can be selectively collected; 

b) seed from each mother tree can be kept separate when 

the need arises; 

c) potential losses to insect and animal interference can 

be minimised; and 

d) damage incurred after falling onto the ground, such 

as that resulting from desiccation and ageing, can be 

limited. 

Generally, collections of seeds should be made from 

healthy trees that have good shape and form, avoiding 

trees that are obviously diseased. Inclusion of immature 

seeds and seeds that have been lying on the ground for 

some time should be minimised. Various methods of 

collection used by the seed collection team at the Forest 

Research Institute Malaysia (FRIM) are described below 

along with their advantages and limitations. The methods 

can be divided into two main types. Firstly, those that do 

not involve climbing, the overall operation being confined 

to the ground (Methods 1-3). Secondly, those involving 

an element of tree climbing (Methods 4-5). 

Factors to be considered for harvesting in the 

aseasonal zones are given in the summary at the end of 

the chapter. 

1. Ground collection 

Ground collection does not require employment of staff 

possessing both tree climbing skills and the ability to 

collect seed efficiently; the cost is thus reduced. 

Nevertheless, this method necessitates good preparation: 

trees must be selected and marked; and all vegetation, 

debris and old or premature seeds below the trees must 

be cleared. Proper supervision of collection is also 

necessary. The limitations of this method are: 

a) seed collection is protracted; 

b) collections have to be made daily until most of the 

seeds have fallen; 

c) there is competition with mammals, birds and insects; 

d) fungal problems, seed deterioration and premature 

germination are encountered; and 

e) ground cover surrounding the tree is destroyed. 

2. Collection using nets or canvas 

With this method, nets or canvas are laid under the tree. 

This procedure is desirable in that undergrowth is not 

destroyed. The limitations of this method are: 

a) it is not suitable under dense undergrowth; and 

b) daily collections of fallen seeds need to be made. 

3. Shaking of seed-bearing branches 

This method is referred to as the ‘fishing line’ method. A 

local home-made catapult is used to shoot a singlefilament 

fishing line, attached to a lead weight, over 

smaller branches of the tree from which seed is to be 

collected. A polythene rope is then pulled over the branch 

and back down to the ground using the fishing line; the 

rope is then pulled vigorously to shake down the seeds. 

The method is suitable for small trees and for those 

standing in the open. The limitations of this method are: 

a) it cannot be used with very tall trees, which may be 

the ones possessing the best genotypes; 

b) a clear view of the terminal branches is required for 

the lead weight to be aimed accurately; 

c) it usually requires several attempts before the line is 

satisfactorily positioned on the right branch; and 

d) the lead weight and line are not fully controllable and 

minor injuries may sometimes be experienced by the 

operator. 

4. Free climbing 

This method is employed by professional tree climbers. 

It involves the use of a neighbouring smaller tree for the 

initial ascent, after which the climber crosses to the main 

seed tree at a height where the bole is small enough to 

hold safely and ascend the tree. The climber cuts off and


drops down small branches bearing the seeds. The 

limitations of this method are: 

a) a suitable smaller proximal tree (or group of trees) is 

required; 

b) it is very strenuous and time consuming which limits 

the number of trees that can be worked on per day; 

and 

c) it is dangerous. 

5. Methods of climbing using equipment 

With the following three methods climbing gear is used 

to gain access to the canopy making the overall procedures 

much safer. 

a) Tree bicycle 

Trees can be climbed without causing serious damage to 

the tree trunk. The equipment consists of two unequally 

long bearing pieces with rests for the feet. Flexible steel 

bands are positioned around the tree trunk at the far end 

of the bearing pieces,. By a bicycling motion the tree 

climber ascends the tree moving the steel bands upwards 

parallel to the tree axis. During this procedure the climber 

wears a security belt with ropes fastened around the tree. 

The equipment offers a comfortable and safe basis for 

standing during working in the crown. This method is 

not suitable, however, for trees that have branches on the 

bole. Also, use is limited to those trees having a girth that 

can be easily encircled by the fastening ropes. 

b) Climbing using spurs 

With this method the climber uses a pair of spurs fastened 

under his shoes in addition to the security belt and 

fastening ropes which were mentioned above for the 

bicycle method. The climber uses the spurs by pricking 

its spikes into the tree bark to secure a foothold for every 

upward movement. The holes made by the climbing spurs 

are vulnerable to fungal, viral and bacterial attack, a 

problem which is aggravated if trees are often climbed in 

this way. It is thus advisable that, if this method is 

employed, an interval of at least a year should be allowed 

before a further collection is made; healing of the 

damaged parts on the trunk can then occur. As is the case 

for the tree bicycle method, the circumference must not 

be too large. 

c) ‘Roping up’ method 

In this method a line is shot up into the crown over two 

or more strong branches. The climbing rope is then drawn 

up over the branches and, on return to ground level, the 

free end is fastened at the base of the trunk. The climber 

then uses the rope to pull himself up using a shoemore. 

This method can be used whatever the girth of the trunk 

and does not damage the tree. 

A combination of elements from different methods 

may be necessary; for example, it may be desirable to 

combine the laying-nets as in Method 2 with the shaking 

element of Method 3. 

Seed Transportation 

The length of time between collection of moist 

dipterocarp seed and its arrival at the seed centre is 

crucial in determining viability. Transport should be 

carefully planned to minimise delay; staff in the nursery 

or seed store should be advised of the schedule so that 

seed can be handled immediately on receipt. 

Methods for transport of OLDA seeds collected in 

the dry condition are given in the summary at the end of 

the chapter. The following points are relevant in relation 

to the transport of moist dipterocarp seeds. 

Ventilation and Moisture Content 

Moist dipterocarp seeds respire intensively and so require 

good ventilation. If large quantities are closely packed, 

the seeds become anaerobic, physiological breakdown 

takes place, fungal growth takes hold and overheating 

occurs; these changes accelerate deterioration of the seed. 

Recalcitrant-seeded species will deteriorate rapidly if their 

moisture content is reduced significantly; ventilation must 

be provided, but without drying the seed. 

If plastic bags are used to contain the seeds, their tops 

should either be left open and folded over or they should 

be tied and small holes made in their sides. Hessian or 

jute bags with a loose weave are also suitable for transport. 

Desiccation is more likely to occur if transport is in open 

vehicles; air movement may accelerate the process. 

Temperature 

Temperatures below 16 o C or above 32 o C should be 

strictly avoided for moist, recalcitrant seeds. Good 

ventilation reduces heat build-up from respiration. Seeds 

should be shaded from direct sunlight at all times during 

transport. 

Long Journeys 

Efforts must be made to dispatch the seeds to their 

destination within two days of collection. If seeds begin


Table 2. Seed (fruit) weight and size indicators at harvest (Tompsett and Kemp 1996a, b). 

Species Mean seeds per kilo Mean length (mm) Mean width (mm) 

Shorea pinanga 30 59 32 

Shorea macrophylla 33 n/a n/a 

Dipterocarpus grandiflorus 50 58 38 

Shorea amplexicaulis 64 46 26 

Dipterocarpus kunstleri 80 55 43 

Dipterocarpus humeratus 90 35 29 

Dipterocarpus obtusifolius 90 20 19 

Dryobalanops keithii 100 n/a n/a 

Dipterocarpus cornutus 110 29 28 

Dipterocarpus caudatus ssp. penangianus 120 26 23 

Dipterocarpus zeylanicus 120 36 23 

Dryobalanops lanceolata 120 26 23 

Shorea palembanica 140 n/a n/a 

Shorea beccariana 160 36 24 

Shorea fallax 160 n/a n/a 

Stemonoporus canaliculatus 160 n/a n/a 

Dipterocarpus turbinatus 170 30 20 

Parashorea tomentella 180 30 20 

Dipterocarpus chartaceus 200 28 22 

Shorea smithiana 200 29 17 

Anisoptera megistocarpa 220 27 20 

Dipterocarpus tuberculatus 230 27 23 

Shorea almon 270 n/a n/a 

Dipterocarpus alatus 360 38 30 

Shorea ferruginea 440 26 13 

Parashorea malaanonan 540 15 14 

Shorea robusta 588 n/a n/a 

Shorea trapezifolia 670 16 9 

Shorea siamensis 680 26 16 

Dipterocarpus tuberculatus var. grandifolius 690 n/a n/a 

Dipterocarpus costatus 760 n/a n/a 

Dipterocarpus gracilis 790 15 13 

Shorea ovalis 790 17 11 

Shorea gibbosa 930 n/a n/a 

Parashorea smythiesii 940 17 11 

Shorea argentifolia 1100 n/a n/a 

Shorea macroptera 1100 19 10 

Shorea roxburghii 1195 16 8 

Anisoptera costata 1200 11 11 

Dipterocarpus intricatus x tuberculatus 1200 24 17 

Shorea congestiflora 1300 19 8 

Shorea parvifolia 1300 17 10 

Shorea selanica 1300 n/a n/a


Table 2. (continued) Seed (fruit) weight and size indicators at harvest. 

Species Mean seeds per kilo Mean length (mm) Mean width (mm) 

Dryobalanops rappa 1400 17 9 

Shorea faguetiana 1400 n/a n/a 

Shorea laevis 1600 14 9 

Anisoptera marginata 1800 10 10 

Shorea leprosula 1800 16 10 

Shorea affinis 1900 n/a n/a 

Shorea leptoderma 1900 n/a n/a 

Shorea ovata 1900 n/a n/a 

Dipterocarpus intricatus 2800 20 17 

Cotylelobium burckii 2900 10 10 

Cotylelobium melanoxylon 2900 9 8 

Shorea obtusa 2900 n/a n/a 

Hopea dryobalanoides 4000 10 7 

Vatica odorata ssp. odorata 4000 8 7 

Hopea parviflora 4100 7 6 

Shorea guiso 4200 11 6 

Hopea odorata 5300 8 6 

Hopea foxworthyi 5500 8 5 

Hopea ferrea 5800 n/a n/a 

Hopea mengerawan 6300 10 4 

Hopea nigra 9000 8 5 

Vatica mangachapoi 17000 5 5 

Monotes kerstingii* 45000 n/a n/a 

*: Assessment refers to seeds inside the fruit. 

to germinate they can be saved by storing in rigid 

containers lined with moist newspaper or other absorbent 

materials to keep them moist during transit. 

Size Considerations 

There is a large range in sizes of dipterocarp seeds (Table 

2) which implies that different handling procedures may 

be needed for moist seed of particular size ranges. For 

example, smaller seeds (< 2 g) would benefit from the 

inclusion of packing material to increase the size of air 

spaces between the seeds. Crumpled newspapers and 

polystyrene chips have been used for this purpose. 

Seed Processing 

The fruit of dipterocarp species, which is the unit 

employed for handling, is generally referred to as the 

‘seed’. Removal of calyx lobes (‘wings’) by manual 

abscission is carried out for all physiology types. This 

enables easier sowing and better contact of the seeds with 

the soil. 

Factors which should be considered in the drying of 

OLDA seeds for storage are described in the summary 

at the end of the chapter. 

Insect infestation can be a major problem in the 

handling of seed, especially in the genus Dipterocarpus 

(Table 3, Prasad and Jalil 1987, Eungwijarnpanya and 

Hedlin 1984); sometimes 100% of individual seedlots are 

rendered useless. Methods to reduce this problem are 

required and would be best supported by studies on insect 

behaviour. Some studies have already been carried out 

on recalcitrant material of the rain forest (Toy et al. 

1992, Toy and Toy 1992); extension of such studies to 

include seasonal-climate species would be advantageous. 

Further discussion of infestation problems can be found 

in Chapter 7. 

Methods for Storage of Dipterocarp 

Seeds 

In the past half century, various methods of storage have 

been proposed for recalcitrant dipterocarp seeds and,


Table 3. Mean insect infestation statistics for species received 

at Kew (Tompsett and Kemp 1996a, b). 

Genus Mean 

percent 

infestation 

more recently, species with OLDA seeds have been 

considered. Successful long-term storage has been 

achieved in the case of some OLDA species. 

In the case of recalcitrant seeds, some methods 

currently available are useful to ensure the survival of 

seed material during extended field collection trips, for 

planting and for storage in the short to medium term. 

However, the methods cannot ensure a continuous supply 

of planting materials throughout the long periods when 

mother trees are not fruiting. 

Work on dipterocarp seed storage is reviewed in the 

Seed Physiology chapter but some practical storage 

methods are briefly discussed. 

Imbibed Storage in Media such as Sawdust, 

Perlite and Vermiculite 

Storage of recalcitrant dipterocarp seeds in sawdust, 

ground charcoal, perlite and vermiculite has been 

employed to maintain high moisture content. This is the 

most commonly used method for prolonging recalcitrantseed 

viability. With care, seeds can be kept viable in this 

way for several months. Table 4 shows some of the work 

carried out on imbibed storage but the limitations of the 

method are: 

a) a proportion of the seeds may germinate due to the 

high moisture content under these conditions; and 

b) in many cases, because of the difficulties in 

controlling aeration and moisture content, necrosis 

may occur and microbial infection may set in; seed 

viability is then severely affected. 

Storage in Airtight Containers 

Dry seeds of the OLDA type have been successfully 

stored in airtight containers. For example, Dipterocarpus 

intricatus has been retained for 2829 days with no loss 

of viability observed (Tompsett and Kemp 1996a, b). 

Storage under a partial vacuum has been attempted 

for seeds of the recalcitrant species Shorea robusta at 

15°C (Khare et al. 1987); 54% viability after a period 

of 49 days in storage was reported, beyond which further 

storage resulted in the death of most of the seeds. 

Unfortunately, moisture content was not measured during 

storage so the extent to which this factor contributed to 

viability loss is unknown. Seed storage in airtight 

containers is not appropriate for recalcitrant-seeded 

species as it leads to an increasing depletion of oxygen 

in the containers, associated with progressive loss of 

viability. 

Storage in Inflated Bags with Different Gaseous 

Environments 

Sasaki (1980), working on recalcitrant-seeded 

dipterocarps of Malaysia, reported that ventilation with 

ambient air was essential for dipterocarp seeds to 

preserve viability. For example, he found that the viability 

of S. roxburghii (syn. S. talura) seed could be prolonged 

to seven months with adequate ventilation. 

Table 4. Examples of optimum recorded storage in various media for imbibed seed of recalcitrant-seeded Shorea, 

Hopea and Parashorea species. 

MC: moisture content. 

Number of 

species 

examined 

Number of 

species 

infested 

Dipterocarpus 35 10 10 

Shorea 16 18 12 

Hopea 8 5 4 

Parashorea 4 2 2 

Dryobalanops 1 3 1 

Species Source 

Optimum storage recorded 

Days Temp. 

( o C) 

Germination 

(%) 

MC 

(%) 

Medium 

Shorea platyclados Tang (1971) 20 16 64 27 Vermiculite 

Hopea ferrea Tompsett (1992) 300 16 40 30-50 Mainly perlite 

Parashorea smythiesii Tompsett (1992) 317 18 46 45 Perlite 

Shorea fallax Tompsett (1992) 50 21 50 40 Sawdust


Most, but not all, studies concerning the effects on 

viability of gases other than ambient air have been carried 

out under poorly controlled conditions. In most cases 

inflated polythene bags were used so that the gas under 

test was liable to mixing over time with ambient air and, 

in addition, respiration of seeds inside the bag altered 

the gas environment. 

Various gaseous environments have been assessed. 

Song et al. (1984) was able to maintain 80% viability of 

Hopea hainanensis seeds for up to 365 days by 

maintaining oxygen levels above 10%. On the other hand, 

Yap (1981) was able to store seeds of D. oblongifolius 

in bags filled with nitrogen and reported a 60% 

germination after a period of 60 days. However, 

Tompsett (1983) reported a stepwise decrease in 

longevity of the seed as oxygen was lowered 

progressively from 10% to zero per cent for the 

recalcitrant seed of the tropical tree species Araucaria 

hunsteinii (Araucariaceae). Carefully controlled 

conditions were employed; a continuous flow of the gas 

under test was supplied to the seed at the correct relative 

humidity. This study also highlighted two further points. 

Firstly, increased concentrations of carbon dioxide and 

ethylene had no beneficial effects (Araucaria 

hunsteinii; Tompsett 1983) and, secondly, oxygen levels 

above 21% did not enhance storage life (D. turbinatus; 

Tompsett, unpublished). There appear to be no reports 

that altering the gaseous environment from that of 

ambient air can increase longevity for recalcitrant seeds. 

Storage Using Germination Inhibitors 

An alternative method to prevent germination during 

storage is by incorporating germination inhibitors into 

the storage system. Substances that have been used are 

polyethylene glycol (PEG), sucrose, sodium chloride 

and abscisic acid (ABA). Goldbach (1979) reported that 

by treating seeds of Meliococcus (Sapindaceae) and 

Eugenia (Myrtaceae) with 10 -4 molar ABA solution at 

15°C it was possible to store seeds for four to six months 

with at least 89% final viability. This general approach 

for recalcitrant seed storage has subsequently not been 

confirmed as successful; a problem encountered with 

the ABA method is the speedy germination of seed during 

storage. 

Fungicide Treatment Followed by Partial Desiccation 

and Storage at Controlled Temperatures 

Partial desiccation was proposed as a favourable 

approach by King and Roberts (1979). Furthermore, 

several researchers have mixed fungicide with stored 

seeds to protect against fungal growth. However, few have 

conducted controlled experiments to test the effects of 

applying combinations of fungicide treatments with 

partial desiccation treatments. Nevertheless, Hor (1984) 

treated cacao seeds with a 0.2% benlate/thiram mixture, 

partially desiccated the seeds by air drying and then stored 

them loosely packed in polythene bags at 21-24°C. The 

viability of the seeds in his study was prolonged from 

one week to about 24 weeks with a final 50% germination. 

This approach needs to be further assessed with the factors 

separately examined. 

Partial Desiccation without Fungicide 

Maury-Lechon et al. (1981) reported partial drying of 

dipterocarp seeds but did not use fungicides. From their 

results they recommended drying seeds to half the original 

moisture content. This latter procedure prevents 

pre-germination in storage. However, as their experiments 

did not include undried controls, the overall benefit was 

not established. 

Storage at Harvest Moisture Content without 

Fungicide Application or Partial Desiccation 

The examples cited from Tompsett (1992) in Table 4 

were not subjected to partial desiccation and were not 

combined with fungicide. Further examples are given in 

the Seed Physiology chapter and show a total of 13 

species capable of storage for longer than 100 days. 

The pre-germination problem associated with the 

storage of moist seed is illustrated by results for S. 

roxburghii; seeds of this species stored at 16°C with 

approximately 40% moisture content had about 50% pregermination 

after 44 weeks of storage (Tompsett 1985). 

However, provided desiccation and mechanical damage 

to the radicle are avoided, viable seedlings can still be 

produced by a high proportion of the pre-germinated 

seeds. 

Research on Seedling Storage and 

Cryopreservation 

Despite the improvements in short to medium-term 

storage, it is not feasible to store recalcitrant dipterocarp 

seeds in the longer term. Complementary methods are 

being sought to ensure a continuous supply of planting 

material. Two approaches have been attempted at FRIM 

in unpublished work of Sasaki and of Krishnapillay; these 

comprise seedling storage and cryopreservation of seed 

materials.


Seedling storage under low light conditions 

It is well established that dipterocarp seedlings usually 

have high survival and slow growth rates over periods of 

several months when grown under low intensity light. 

Many studies, including those of Brown and Whitmore 

(1992) and Press et al. (1996), report this phenomenon. 

The idea of using this phenomenon for the storage of 

recalcitrant-seeded species was first clearly proposed by 

Hawkes (1980). 

The two methods outlined below, have been tested at 

FRIM: (i) storage of seedlings in a seedling chamber; 

and (ii) storage of seedlings on the forest floor under 

subdued-light conditions. 

Seedling chamber storage 

With this method, freshly collected seeds were surface 

treated with a fungicide (0.1% benlate/thiram mixture) 

and allowed to germinate under ambient conditions in 

containers kept at high humidity with moistened tissue 

paper. After radicle emergence, germinated seeds were 

packed loosely in polythene bags, trays or boxes lined 

with moist tissue paper and stored in a specially 

constructed seedling chamber in which temperature, 

humidity and light were controlled. The temperature was 

16°C, the relative humidity was 80% and the photoperiod 

was 4 hours. Light was supplied from a fluorescent source, 

giving 80-1000 lux. Development of the germinated seeds 

into seedlings occurred slowly in the chamber. Seventeen 

dipterocarp species have been tested (Krishnapillay, 

unpublished); these species, with the periods they have 

been stored, are listed in Table 5. 

Seedlings developed slowly in the chamber, barely 

attaining the heights of 20-25 cm over the storage periods 

tested. Seedlings which were transferred to the nursery 

and grown in polythene bags needed to be weaned in at 

least 70% shade for a period of 2-3 weeks before they 

could be placed under direct sunlight. Survival percentage 

was between 60 and 80%, dependent on the species. 

Forest Floor 

The second approach for storage of seedlings is on the 

forest floor under subdued light. Areas were cleared of 

undergrowth and freshly collected seeds were sown. 

Seedlings developed very slowly and so can remain within 

manageable heights for long periods of time. 

Seedlings of Hopea odorata did not grow to a height 

greater than 10 cm under these conditions over a period 

of three years. Seedlings transferred to the nursery and 

Table 5. Storage periods for Hopea, Dipterocarpus, 

Shorea and Dryobalanops species in a subdued-light 

chamber (Krishnapillay, unpublished). 

Species Period of storage 

(months) 

Hopea odorata 9-12 

Hopea helferi 9 

Dipterocarpus cornutus 6 

Shorea macrophylla 4 

Shorea leprosula 6-9 

Shorea acuminata 8 

Shorea longisperma 6 

Shorea parvifolia 8-9 

Shorea ovalis 8-9 

Shorea curtisii 8-9 

Shorea platyclados 8-9 

Shorea bracteolata 6 

Shorea macroptera 6 

Shorea maxwelliana 4 

Shorea pauciflora 6 

Dryobalanops aromatica 5 

Dryobalanops oblongifolia 4 

grown in polythene bags began to increase in size rapidly. 

Weaning in 70% shade for 2 weeks before transfer to 

direct sunlight was, however, necessary. Survival was 

approximately 80-90%, depending on species. Eight 

species have been tested. 

The constraints with this method are as follows. In 

the early stages after sowing, unprotected seeds are likely 

to be predated by squirrels, birds and wild boars. Fencing 

the area with barbed wire and covering the seed bed with 

a plastic sheet is thus necessary. The plastic sheet can be 

removed when the seedlings have emerged when damage 

by birds and squirrels is unlikely. 

Cryopreservation of dipterocarp seed material 

Cryopreservation generally refers to the preservation of 

material at -196°C, which is the temperature of liquid 

nitrogen (LN). The method is being examined at FRIM 

for the storage of dipterocarp seed material. At this 

temperature, all metabolically related sources of 

deterioration in the seed are greatly reduced or stopped, 

thus supporting preservation for very long periods. Work 

of this type has been carried out on some 

recalcitrant-seeded tree species of temperate climates


(Pence 1992). In addition, material from the recalcitrant 

seed of the tropical forest tree Araucaria hunsteinii can 

be cryopreserved; storage of species for four years at 

-20°C has been achieved, viability being measured in 

terms of callus production (Pritchard et al. 1995). 

Growth occurred from the radicle end of the embryo axis. 

Some results achieved by Krishnapillay and 

colleagues (unpublished) are described. Studies were 

conducted on the recalcitrant-seeded dipterocarp species 

Hopea odorata and Dryobalanops aromatica. Embryos 

were first subjected to cryoprotection treatment using 

sucrose and dimethyl sulphoxide; following this, the 

embryos were partially dried to a moisture content near 

14-15%. The temperature of the material was then taken 

to -30°C at the rate of 1°C per minute, finally being 

reduced to -196°C by plunging into LN. After one week 

the embryo axes were removed, thawed at 40°C and 

evaluated for survival. About 5% showed signs of 

swelling and/or the emergence of growth initials. These 

post-thawing changes were observed in the epicotylar 

region; no development was observed in the radicle region 

and none of the embryonic axes were able to grow into 

whole plants. Improvements to the protocol are being 

sought. A total of 50 excised embryo axes were used in 5 

replicates for each study and for each species. 

Cryopreservation has also been used for whole seeds 

of Dipterocarpus alatus and D. intricatus (Krishnapillay 

and Marzalina, unpublished). However, these species are 

basically orthodox in storage physiology (Tompsett 

1987). Cryopreservation is not recommended for species 

of this storage physiology type because of the comparative 

practical benefits of using conventional seedbank storage 

at -20°C (Pritchard 1995). 

The greatest proven uses of this approach have been 

with small pieces of tissue. Complete success in the 

production of entire seedlings after freezing of tissues 

may require the development of in vitro culture methods 

(see Chapter 3). In addition, nursery techniques for 

weaning the developed plantlets are required. 

Considerable investment of research time and resources 

may thus be needed to assess if the method can be useful 

in practice for recalcitrant-seeded material. 

Summary of Seed Handling Methods 

For South and Southeast Asian Dipterocarpaceae, the 

following current seed handling recommendations have 

been made (Tompsett and Kemp 1996a, b). 

Collection Recommendations 

Check a small sample of seeds before collecting, since 

insect infestation may be excessive. Collect seeds from 

the tree when the wings are turning from green to brown. 

Collection is best accomplished by shaking or plucking 

branches; a climber may be needed where branches are 

inaccessible from the ground. Plan the collection to 

minimise the period of time (preferably a maximum of 

three days) between harvest and either nursery sowing or 

short-term storage at the seed centre. 

Transport Recommendations 

Recalcitrant and OLDA seeds are considered separately. 

Recalcitrant seeds should be transported moist and in 

ventilated containers; they should be kept as cool as 

possible but not below 18°C. If the wings are left intact, 

a reservoir of air is created which provides oxygen for 

respiration. This method reduces both the imbibition of 

moisture in the container and the accumulation of heat 

produced by respiration, thereby limiting the chance of 

germination during transport. Possible containers include 

open, folded-over polythene bags, closed polythene bags 

with small ventilation holes, and open-weave sacks. 

Where greater rigidity is required, the bags or sacks should 

be enclosed in cardboard or wooden boxes with 

ventilation holes. Care should be taken to avoid overheating 

by exposure of the containers to direct sunlight. 

Additionally, seed should be retained above its lowest-safe 

moisture content 

For species with OLDA storage physiology seeds, 

collections may sometimes need to be made from the 

ground with moisture contents at or below 12%. Dry seed 

of this type should be transported as follows. For use in 

the short-term, transport the seed at a cool temperature 

above 2°C; for use in the long term, transport material at 

as low a temperature as possible, but not below -20°C. 

Retain the dry seed in sealed containers during transport. 

For moist OLDA seed, follow the methods described for 

transport of recalcitrant seed. 

Processing Recommendations 

Remove wings for ease of handling and to reduce storage 

bulk for all species. 

Other processing applies to OLDA species. Seeds of 

this type will dry well in 20°C or higher with a low relative 

humidity. Material should be transferred to the appropriate 

storage conditions as soon as the desired moisture content 

is reached. Retaining seeds in a monolayer in a flow of


air will ensure rapid drying, thereby reducing the risk of 

seed ageing. Careful removal of the calyx can further 

reduce bulk for dry storage. However, this procedure is 

time consuming and may only be economic for longerterm 

(conservation) storage. 

Storage Recommendations 

Procedures are recommended separately for small and 

large quantities of recalcitrant seeds and for OLDA 

seeds. 

For larger quantities of recalcitrant seeds, material 

should be kept at near the harvest moisture content and 

in media such as sawdust and perlite. Seed moisture 

content should be checked at the start and then 

periodically during storage; any wide fluctuations 

observed should be counteracted by increasing or 

decreasing the moisture content of the medium. This 

careful moisture content monitoring and management 

can reduce the rate of pre-germination. Excess moisture 

in the medium causes the seeds to become anoxic, whilst 

too little moisture lowers seed moisture content and 

leads to desiccation damage. Suitable containers include 

open-weave sacks or bags. Storage in a high-humidity 

room at 18 o C is recommended. 

The optimal condition for storage of smaller 

quantities of recalcitrant seed is retention within inflated 

polythene bags in a 99% relative humidity incubator at 

18 o C and at a moisture content near that of the seed at 

harvest. Polythene bags with rib-channel closure provide 

suitable packaging; alternatively, loosely-tied, thin-gauge 

polythene bags may be employed. Insertion of a rigid 

object helps to maintain an air space. Ventilation at least 

weekly is essential; use of air at a high relative humidity 

would be desirable for this purpose. Moisture content 

should be checked at the start and then periodically 

during storage; fluctuations should be counteracted by 

increasing or decreasing the relative humidity around the 

seed, if possible. 

Different handling is required for OLDA species; the 

summarised methods for collection, processing and 

transport described above must be followed closely if 

the three storage methods below are to be effective. 

a) Before sowing in the nursery, seed storage of OLDA 

species at about 40-50% moisture content is suitable 

over very short periods of about fifteen days. Moist 

storage avoids the risk of partial loss of viability on 

drying. Temperatures employed should be no lower 

than 18 o C. The seed containers described above in 

relation to recalcitrant species for smaller and larger 

seedlots are suitable. 

b) In the case of longer-term storage any initial, partial 

loss on drying may be outweighed by the improved 

final longevity achieved. Storage for conservation is 

possible if seed is dried to approximately 12% 

moisture content; material should be sealed in 

suitable rigid containers (e.g. Kilner jars) and retained 

at -20 o C. Further relevant information is given in the 

summarised processing method above. 

c) For medium-term storage periods (between 2 weeks 

and 24 months), OLDA seed should be retained at 

2°C with other conditions as prescribed for longerterm 

storage. 

Germination Recommendations 

Remove seed wings prior to sowing in order to ensure 

good contact with the germination medium and germinate 

at approximately 26 o C -31 o C. In the case of dry, decoated 

OLDA seeds (de-coating is recommended for 

conservation storage), slow imbibition is essential. This 

can be achieved by retaining the material in 100% 

humidity for 24 hours before sowing. 

Research Priorities 

Changes are occurring in relation to reforestation and 

afforestation programmes in the regions where 

dipterocarps are grown. The emphasis is now on the use 

of indigenous species in combination with exotics. It 

follows that suitable planting material will be 

increasingly in demand. Hence, there is a necessity to 

increase research in the areas described below: 

1. Optimising methods for the collection, processing, 

storage and germination of forest seeds so that seed 

storage life is maximised, taking into account the 

need for retaining the viability of seeds that germinate 

during storage. More detailed suggestions are given 

in Chapter 3. 

2. When the seed storage physiology is known, other 

information is required. In particular, practical 

methods for large-scale drying are required in the 

case of OLDA species, and methods for the storage 

of bulky recalcitrant material need improvement. 

3. Identification of seed-predating insects leading to 

assessment of their behaviour, especially in the 

seasonal zones, is desirable. This would complement 

studies undertaken already in the aseasonal areas.


4. Means to store germinated seeds as seedlings should 

be assessed. 

Relevant Institutions with their 

Strengths and Potential Contributions 

A national pattern of involvement in seed handling 

activities is given below, using the example of Peninsular 

Malaysia. This structure may not apply in other 

dipterocarp countries, but elements of it may be relevant. 

To carry out the tasks of procuring material for the 

planting programmes, there is the need for the close 

networking of the particular agencies involved. These can 

include the Forest Department of Peninsular Malaysia 

(FD), FRIM, and the Malaysian Timber Council (MTC). 

Other relevant agencies are the Agricultural University 

of Malaysia (UPM), the ASEAN Forest Tree Seed Centre 

(AFTSC) and the private sector. 

Institutions with involvement in dipterocarp seed 

research in other countries are referred to in the Seed 

Physiology chapter. 

Forest Department of Peninsular Malaysia 

The FD consists of the Headquarters in Kuala Lumpur 

and 10 State Forest Departments located throughout the 

territory. The headquarters is responsible for planning, 

operational studies and development of the forestry 

sector as well as provision of technical advice and 

services and the provision of training facilities for the 

forest industry. The State Forest Departments are 

entrusted with the management of the forest in the 

respective states. 

The role of the Forest Department in the seed and 

plant procurement programmes may include: 

1. assessing plant demand for the each planting activity; 

2. providing areas in the forest for seed collection; 

3. providing manpower to be trained in carrying out 

phenological studies, monitoring development and 

collection of seeds at maturity; 

4. allowing upgrade of state nurseries for large-scale 

plant production; and 

5. providing the manpower for planting and subsequent 

maintenance of the planted areas. 


FRIM is a statutory research body with the mandate to 

promote and improve the sustainable development of 

forest resources and their industrial uses through 

research, development and application activities. 

The purpose of FRIM is to develop appropriate 

knowledge and technology for the conservation, 

management, development and utilisation of forest 

resources. Excellence in scientific research and 

development, and technology transfer to the forestry 

sector is also pursued. 

FRIM is the research arm to the Forest Department; 

its role in the seed programme is: 

1. providing technical expertise in tree selection, phenological 

monitoring, and seed collection; 

2. seed handling, nursery techniques (including vegetative 

propagation techniques), setting up of seed orchards, 

seed testing, and the documentation and certification 

of storage details; 

3. assisting in the development of a programme for seed 

and plant procurement; and 

4. making available its international contacts for the 

improvement of the seed and plant procurement programme. 

Malaysian Timber Council 

MTC has the mission to promote the development of 

timber-based industry and to facilitate trade in timber 

within Malaysia. Among the objectives of MTC are to 

promote the rehabilitation of degraded forests and to 

encourage the reforestation of logged-out areas. 

The role MTC can play to enhance the seed 

programme is in: 

1. becoming an investment arm of the seed and plant 

procurement programme; and 

2. assuming a role in the rehabilitation of degraded forests 

on a privatised basis. 

Agricultural University Malaysia 

The Faculty of Forestry of UPM is responsible for 

producing trained manpower in all aspects of the forestry 

sector. 

The Faculty of Forestry could, as a teaching and 

research unit, contribute to the seed programme by: 

1. providing scientists trained in the fields of seed and 

plant procurement; 

2. disseminating research information related to seed 

programmes; and 

3. providing training (short courses in forestry) to upgrade 

the skills of those involved in the seed programme.


ASEAN Forest Tree Seed Centre 

This centre in Thailand serves the needs of all ASEAN 

countries in relation to forest seed problems common 

to all the ASEAN countries. 

The AFTSC could assist in the seed programme by: 

1. raising funds and acting as host for the training of 

seed research personnel; 

2. providing relevant technical support through shortterm 

consultancies; and 

3. disseminating knowledge and technology gained in 

seed programmes. 

Private Sector 

The private sector is at present not significantly involved 

in the plantation forestry programmes in Peninsular 

Malaysia. However, there is interest being expressed 

by the large plantation holdings to go into forest 

plantation in support of the Malaysian government’s 

aspirations to produce timber from sustainably managed 

forests. 

With its long and successful experience of rubber, 

cocoa and oil palm plantation, the private sector could 

contribute to the seed programme by: 

1. contributing experience in establishing large-scale 

plantations of forest species; 

2. managing large-scale nurseries; and 

3. becoming investors in forest plantations. 

Institutes with Resources Relevant to Insect 

Research 

The central laboratory in the Royal Forest Department 

in Thailand has a programme of research on forest tree 

insects. In the UK the Natural History Museum (British 

Museum) also has resources and relevant experience in 

relation to insect research. 

Conclusion 

In this chapter, current knowledge on handling of 

dipterocarp seeds has been outlined and areas indicated 

where further work is required. The potential exists to 

overcome difficulties in producing planting material, 

but the collaboration of several agencies is required. A 

suggested framework has been provided for Peninsular 

Malaysia. While the individual organisations may not 

be entirely the same in other countries, equivalent 

groups will need to collaborate to attain the objectives. 

Acknowlegements 

We thank the Forest Research Institute Malaysia, the 

Royal Botanic Gardens Kew and the Center for International 

Forestry Research for providing facilities and financial 

support. 

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Seedling Ecology of 

Mixed-Dipterocarp Forest 

M.S. Ashton 

Introduction 

Successful reproduction depends on the completion of a 

sequence of events starting with flower bud initiation and 

ending with the establishment of a young seedling (Smith 

1986); failure of any single stage in this sequence can 

have catastrophic consequences for the regeneration of a 

new stand. Several stages of the sequence considered in 

this chapter are i) the dispersal of fruits; ii) germination 

of seed; iii) early survival; and iv) the establishment of 

seedlings. These stages comprise a period of 

reorganisation and initiation of a new forest stand after 

which composition and structure depends mainly upon 

competition and self-thinning. These stages provide an 

opportunity in silviculture for promoting the desired 

composition and stocking of the future stand. To quote 

from Smith (1986) ‘Many of the successes or failures of 

silvicultural treatment are preordained during stand 

establishment. Physicians bury their worst mistakes but 

those of foresters can occupy the landscape in public view 

for decades’. 

South and southeast Asia boast a rich history of forest 

research. The mixed-dipterocarp forest 1 of this region has 

been studied more than any other tropical forest type 

primarily because of its importance for producing timber. 

This chapter reviews the state of knowledge on the 

seedling ecology of regenerating mixed-dipterocarp forest 

and suggests future avenues of research. However, it is 

not an exhaustive review of the literature and in most 

cases cites widely available papers. There is much 

information on seedling dipterocarp ecology that remains 

unpublished or is only available at local research institutes, 

or university and government departments. This 

information in its own right deserves documentation, 

compilation and synthesis. Also, though this account 

concentrates on a review of the literature of the seedling 

ecology of dipterocarp species it emphasises the need to 

obtain information about the seedling ecology of non- 

Chapter 5 

dipterocarp species in mixed-dipterocarp forests. Often 

silvicultural management of mixed-dipterocarp forests has 

concentrated on the regeneration autecology of a few 

commercial dipterocarp species without an understanding 

of their interaction with other species, and their role in 

the successional dynamic of the whole forest. This has 

led to a silviculture that has focused on only the current 

commercial species and has tended to simplify, and in 

many instances degrade, the dynamic and structure of 

mixed-dipterocarp forests (Ashton et al. 1993). 

Dispersal and Germination 

Early studies on mixed-dipterocarp forests were done on 

seed phenology and dispersal mechanisms and the 

categorisation of tree species by dispersal agent (Ridley 

1930). Subsequent work has been done in more detail on 

the role of seed dispersal by animals (Medway 1969, 

Leighton 1983, unpublished data); and on dipterocarps 

in particular (Fox 1972, Kochumen 1978, Dayanandan 

et al. 1990). However, these studies are few and much 

more long-term phenological information on seed 

dispersal needs to be gathered on representative guilds 

of species within mixed-dipterocarp forest. Future studies 

should also focus on the amount and distribution patterns 

of seed dispersed from parent trees and germination. This 

will lead to a better understanding of the spacing and 

period of time required for the retention of a residual 

overstorey to ensure satisfactory stocking of seedlings. 

This kind of information is essential for the development 

of natural regeneration methods. 

1 Mixed-dipterocarp forest is defined here as that lowland and hill 

rain forest where the Dipterocarpaceae are predominant amongst 

the canopy and emergent trees of mature forest. The majority of 

tree species are non-dipterocarp. The soils are weathered in situ 

and would be classified as belonging to either oxisols or ultisols 

(USDA 1975). The climate is warm and humid with high rainfall 

that has little seasonality.

Seedling Ecology of Mixed-Dipterocarp Forest 

Work has been done on dipterocarp germination 

mostly during the 1970s and 80s. Many studies of 

dipterocarp species reported them to be recalcitrant 

(Jensen 1971, Tang 1971, Tang and Tamari 1973, Tamari 

1976). Chapter 3 gives a more detailed review of 

dipterocarp germination. Other studies suggested that 

many non-dipterocarp species, mostly pioneers, had 

dormant seed buried in the soil that germinated with a 

marked increase in radiation at the ground surface (Liew 

1973, Aminuddin and Ng 1983, Raich and Gong 1990). 

However, unlike the neotropics (see work by Vazquez- 

Yanes and Orozco Segovia 1984, Garwood 1996), no 

critical experiments have focused on this buried seed 

phenomenon. Also, relatively few studies have 

comprehensively evaluated patterns of germination for 

the whole forest in relation to successional status and 

taxonomy (Ng 1983). This is perhaps because past 

research has focused on the autecology of individual 

dipterocarp species. Future work should focus on 

clarifying the germination mechanisms of mixeddipterocarp 

forest tree species in general and the role 

dipterocarp species play within it. Little work has focused 

on the competitive interactions between dipterocarps and 

non-dipterocarps and yet they are of direct relevance to 

the maintenance of dipterocarps in a managed forest. 

Early Survival and Establishment of 

Dipterocarps 

The main research objective early in this century was to 

develop a method for evaluation of regeneration stocking 

before logging (Wyatt-Smith 1963). The survey 

techniques that developed were usually based on line 

transects that assessed stocking by measures of 

dipterocarp seedling distribution and number. Surveys 

revealed that the abundance of regeneration was 

associated with certain dipterocarp species and sites. In 

many circumstances regeneration was absent particularly 

on the slopes of hill forests and where competing 

understorey palms, shrubs and herbs were present 

(Burgess 1975, Wong 1981, Kusneti 1992). Though 

measures of distribution are important to gauge adequate 

and even coverage of seedling establishment within a 

stand, measures of seedling number and density do not 

necessarily predict successful establishment. A measure 

that incorporates an estimate of seedling vigour is needed. 

More recent studies have used different size classes and 

estimates of leaf area to gauge vigour, promoting survey 

90 

techniques that discard seedlings in the ‘less vigourous 

classes’ for a representation of regeneration stocking 

(Ashton 1990). These can be useful measures for most 

dipterocarp species because they have poor ability to 

sprout. Measures of their above-ground performance can 

therefore be used to predict future growth and survival. 

Studies by Nicholson (1960) and others (Fox 1972, 

1973, Liew and Wong 1973, Tomboc and Basada 1978, 

Appanah and Manaf 1994) elucidated the cyclic nature 

of population recruitment and survival in the groundstorey 

of a closed forest and demonstrated the importance of 

advanced regeneration in the form of a seedling bank for 

the successful establishment of new forest stands. 

Conceptual models of the regeneration dynamic have 

been developed that explicitly suggest the importance and 

reliance of mixed-dipterocarp forest on advance 

regeneration (see Fig. 1). This reliance is not only for 

dipterocarps but also for late successional canopy trees 

that are non masting, subcanopy trees and shrub species. 

Forest management should therefore focus on advanced 

regeneration of dipterocarp trees and similar associates. 

These are the trees that are the canopy dominants during 

the mid and late stages of forest succession. They, 

therefore, create the basic forest structure beneath which 

other strata exist, and reflect the changes in composition 

associated with differences in site quality. 

Studies have also shown that dipterocarp species 

could be broadly categorised as shade-tolerant or lightdemanding 

based on differences in frequency of 

recruitment and rate of seedling death. Shade-tolerant 

dipterocarps can have seedlings established beneath 

closed canopied forest for long periods of time (> 10 

years). Mast years for shade-tolerant dipterocarps can 

therefore be fewer than relatively more light-demanding 

dipterocarps but still provide adequate advance 

regeneration establishment (Wyatt-Smith 1963, Fox 1972, 

Gong 1981). In general, however, all dipterocarps have 

cohorts of seedlings that continually replenish the seedling 

bank from successful mast years. Over time, seedlings 

die primarily from the very low light regimes of a closed 

forest canopy (Ashton 1995). Groundstorey levels of 

photosynthetically active radiation (PAR) beneath the 

canopy of a mixed-dipterocarp forest have often been 

recorded as less than 1% of that received in the open 

(Torquebiau 1988, Ashton 1992a). 

Other studies have also suggested the importance of 

an increase in amounts of PAR that promotes only partial 

shade for dipterocarp germination and early survival


Figure 1. Regeneration recruitment frequency and stand canopy dominance of ecological species groups over different successional stages of stand 

development for a mixed dipterocarp forest. Examples of species are given for each ecological group along with codes denoting their structural position 

within the stand over time. Note the periodic recruitment of seedings for tree species belonging to the late-successional canopy dominants. (modified after 

Ashton 1992a). 

PHASES OF STAND DEVELOPMENT 

BUILDING MATURE 

GAP 

Disturbance 

Approximate point of canopy closure 

Approximate peak in canopy exclusion 

Initiation of groundstory Canopy break-up 

PIONEERS OF GAP PHASE 

(Macaranga, Trema) 

PIONEERS OF BUILDING PHASE 

(Albizia, Alstonia) 

LATE-SUCCESSIONAL DOMINANTS 

(Dipterocarpus, Dryobalanops, Shorea) 

LATE-SUCCESSIONAL NON-DOMINANTS 

(Durio, Ficus, Mangifera) 

LATE-SUCCESSIONAL SUBCANOPY 

(Calophyllum, Garcinia) 

LATE-SUCCESSIONAL UNDERSTORY 

(Gaertnera, Psychotria) 

EARLY SUCCESSIONAL MID-SUCCESSIONAL 

LATE SUCCESSIONAL 

Juveniles recruited under canopy light conditions and considered as advanced regeneration (seedlings, seedling sprouts, root and stem 

suckers). The breadth of the bar represents amount of regeneration relative to other ecological species groups. 

Juveniles recruited under open conditions of full sun (buried seed, seed dispersed by wind or animals into opening after disturbance). 

91 

Stand canopy dominance. The breadth of the bar represents the degree of dominance in relation to other ecological species groups.


(Nicholson 1960, Ng 1978). However, it was shade house 

investigations (Mori 1980, Sasaki and Mori 1981, Ashton 

and de Zoysa 1989) that clearly demonstrated that most 

dipterocarp seedlings require greater amounts of radiation 

than that received at the groundstorey of a closed canopy 

dipterocarp forest, but less than the amount of radiation 

received when exposed to full sun. 

Field research on disturbance regimes of mixeddipterocarp 

forest supports evidence from shade house 

experiments and studies of seedling population dynamics 

in the forest. Natural disturbances documented in mixeddipterocarp 

forest are varied but most are of a kind and 

scale that promote the survival and release of an existing 

seedling groundstorey. Disturbance types include 

lightning strikes (Bruenig 1964), insect defoliation of 

canopy trees (Anderson 1964), single and multiple tree 

falls (Anderson 1964), and cyclones (Whitmore 1974, 

1989). All allow the groundstorey vegetation to remain 

largely unharmed. Disturbances in mixed-dipterocarp 

forest that have been observed to destroy groundstorey 

vegetation include landslides, flooding and fire (Day 

1980, Leighton and Wirawan 1986, Tagawa and Wirawan 

1988). This might be one reason why dipterocarp 

regeneration does not establish well in parts of a forest 

landscape subject to lethal disturbance of the groundstorey 

such as steep slopes, flood plains and swidden agriculture. 

Current investigations focus on refining our 

understanding of the regeneration microenvironment of 

tree species in mixed-dipterocarp forest. Compared to 

other tree species of mixed-dipterocarp forest, 

dipterocarps have some general autecological 

characteristics that allow for their categorisation in the 

same regeneration guild (Table 1). Although dipterocarps 

have the same general autecology there are also 

differences among them, however, these differences are 

small compared to other regeneration groupings. 

Important questions are: what degrees of difference exist 

and why do they occur among species belonging to the 

same congeneric group. The answers are particularly 

relevant to understanding dipterocarp dominance in 

mixed-dipterocarp forests and will provide the silvical 

information for the fundamental treatments imposed on 

these forests for management purposes. 

One such topic that merits attention is the site 

specialisation of dipterocarp regeneration. How site 

specific is advanced regeneration of dipterocarp species? 

Recent field studies demonstrate that forest gaps of 

different size exhibit considerable spatial (Ashton 1992a, 

92 

Table 1. Silvical characteristics of canopy tree species 

belonging to genera assemblages (e.g. Shorea) that dominate 

the mature phase of mixed-dipterocarp forest. These 

characteristics should be interpreted broadly as exceptions 

will exist (Ashton 1992b). 

Reproduction 

• Pollination vectors are small insects 

(hymenoptera, hemiptera) 

• Seed is with storage tissue 

• Seed is dispersed by gravity (often aided by 

territorial animals such as rodents) 

• Fruiting time is more or less supra-annual with 

distinctly different amounts of seed at each fruiting 

(masting) 

• Seed shows no classical dormancy 

Establishment and Growth 

• Seed requires partial shade protection for 

germination and early survival 

• Seedlings require an increase in light (as 

compared to understorey conditions) for 

satisfactory establishment and growth 

• Seedling survival and establishment is usually site 

specific, according to particular biotic, 

microclimatic and edaphic characteristics 

Brown 1993) and temporal (Raich 1989, Torquebiau 

1988) variation in forest groundstorey microclimate. 

Changes in size of small canopy openings can greatly 

influence the overall amount of radiation received at the 

groundstorey of the opening centre (Brown 1993). 

However, larger canopy openings provide a greater range 

of microclimates at the groundstorey of the opening 

(Ashton 1992a). Studies that monitored pre-established 

seedlings and new recruits (Raich and Christensen 1989, 

Brown and Whitmore 1992) showed that there were 

significant differences among dipterocarp species in 

survival and growth at these different microsites. Studies 

by Ashton et al. (1995) that controlled age and spacing 

of dipterocarp seedlings supported these findings. 

However, investigations by Turner (1990a, b), who 

monitored pre-established seedlings, suggested mixed 

results of dipterocarp seedling survival and growth in 

relation to light availability at the scale of the microsite. 

Studies are now investigating the competitive relationship 

between species for regeneration growing space through 

the monitoring of long-term self-thinning trials located 

on different sites and within different


microenvrionments (Gunatilleke and Ashton, 

unpublished). 

These studies have focused investigations on the 

spatial availability of light at the forest groundstorey and 

its relationship to seedling survival and growth. Findings 

also suggest that the seasonal variation in soil water 

availability, scaling up from different groundstorey 

microsites to across the landscape (ridge to valley), can 

be another factor that affects the survival and growth of 

dipterocarps. Ashton (1992a), Brown (1993) and 

Palmiotto (1993) observed seasonal periods of water stress 

that may play a critical role in determining seedling 

composition of canopy gap regeneration. Transplant 

experiments suggested soil water availability, related to 

topography (slope, ridge, valley etc.), affects the survival 

and growth of dipterocarps. The transplant experiments 

also suggested seedling survival and growth allocation 

was affected by interaction between soil water availability 

and radiation. For example, some species showed fivefold 

decreases in root mass between seedlings growing 

in the understorey of a ridgetop site as compared with 

those seedlings in the understorey of a valley site. 

Although understorey PAR was comparable between the 

two sites the poor root development on the ridge 

predisposes these species to drought. Studies by Brown 

and Whitmore (1992) and Ashton et al. (1995) suggested 

that seedlings of more light demanding dipterocarp 

species have larger leaves and that more shade tolerant 

species have smaller leaves and are more sensitive to heat 

stress. This is contrary to most other literature for example, 

Givnish (1988) which has mostly described the sun shade 

dichotomy for mature trees that are from temperate forest 

regions. There are only a few studies (Ashton and Berlyn 

1992, Strauss-Debenedetti and Berlyn 1994) that have 

investigated the sun shade dichotomy for seedlings of 

the moist tropics. 

Other studies are also providing evidence that 

dipterocarps are affected by soil characteristics related to 

the underlying parent material. Surveys by Baillie et al. 

(1987) and Ashton and Hall (1992) suggest both 

concentrations of total and available magnesium and 

phosphorus to be particularly important in determining 

species-site associations. However, no fertiliser studies 

of seedlings using field experiments have clearly 

demonstrated that these factors affect the establishment 

stage of forest development (Turner et al. 1993, Burslem 

et al. 1995) although some studies that are in progress 

are suggesting differences may occur (Gunatilleke et al. 

93 

1996, Palmiotto, in preparation). In these experiments 

different soils are being investigated to understand 

nutrient use efficiency of dipterocarp species whose 

distribution is restricted to very different levels of soil 

fertility. These kinds of studies are beginning to provide 

the basis for the development of new silvicultural 

regeneration methods and the refinement of currently used 

methods. These studies on light, soil moisture and fertility 

are providing knowledge for a better mechanistic 

understanding of regeneration dynamics of forests. In 

some cases they have contradicted previous understanding 

of forest dynamic patterns based only on observation and 

census methodologies. An example would be the recent 

findings that show discrete differences in the sitespecialisation 

among species of Shorea section Doona. 

These species were formerly assumed to be very similar 

in their site requirements and therefore their silvicultural 

treatments were the same. 

In addition, there are many biotic interactions that 

can moderate or accentuate patterns in the establishment 

of seedlings within the physical environment. For 

example, although no studies substantiate this, host 

specific ectomycorrhizae could accentuate the differential 

exploitation of soil nutrient resources among closely 

related assemblages of dipterocarp species. Studies by 

Becker (1983) and Smits (1983) suggest that 

ectomycorrhizae can play important roles in dipterocarp 

seedling establishment and growth. Mycorrhizal infection 

was found to be greater for seedlings located in small 

clearings than for those seedlings located beneath forest 

canopy. These results suggest that seedling regeneration 

of dipterocarps will respond more vigourously to 

overstorey removal if pre-release treatments create higher 

light environments in the understorey. In addition, Lee 

and Lim (1989) found that foliar phosphorus contents of 

Shorea seedlings growing on either phosphorus deficient 

or phosphorus rich soils were the same - indicating a 

difference in uptake efficiency that was attributed to 

ectomycorrhizae (for more detail see Chapter 6 on 

nutrition and root symbiosis). 

Herbivory is another biotic effect that has had little 

investigation. Becker’s (1981) studies of seedling 

populations found less herbivory on the leaves of a late 

successional, more shade-tolerant Shorea species as 

compared to more light-demanding Shorea species. 

However, no studies followed up on this work. More 

investigation should be done, particularly on the role of 

non dipterocarp tree species in mixed-dipterocrap forest.


Does the simplification of mixed-dipterocarp forest by 

the frequent use of various silvicultural release 

treatments (weeding, cleaning, liberation) favour so few 

commercial tree species that this may lead to greater 

susceptibility to disease and/or herbivory of the forest? 

Questions such as these need to be further tested. 

Growth in Relation to Physiology and 

Structure of Dipterocarps 

Recent seedling experiments have focused on separating 

the various abiotic and biotic factors that influence 

seedling establishment and growth under controlled 

conditions. Many studies have been investigating light 

and the different effects of light quality, quantity and 

duration. These experiments reinforced findings from 

the earlier shade house studies but demonstrated that 

forest understorey light quality can accentuate the poor 

growth and survival of seedlings in deep-shade 

conditions (Kamaluddin and Grace 1993, Lee et al., 

unpublished manuscript). Experiments that simulated 

quality and intensity of light environments of a rain forest 

also demonstrated that Shorea species allocate dry mass 

proportions to roots, stems and leaves in different 

amounts (Turner 1989, Ashton 1995). These results show 

that the more shade-tolerant Shorea species allocate 

proportionately more dry mass to root development than 

to stem and leaves in forest understorey environments 

whereas the reverse is true for more light-demanding 

Shorea species. 

The process of photosynthesis requires 

photosynthetically active radiation, water and carbon 

dioxide . The adaptations a seedling leaf can make to its 

surroundings must accommodate all three. The 

relationship among all three factors is so closely linked 

that many of the leaf adaptations and adaptation responses 

to environmental change are the same. Heat and 

desiccation of leaves exposed to the full radiation of the 

sun can promote leaves that have similar physiological 

and anatomical adaptations as leaves that are droughtenduring. 

Leaves that have grown in the shade often 

resemble those of drought intolerant leaves. Alhough 

much work has been done elucidating differences in leaf 

anatomy and morphology between species of different 

cladistic or successional groups for other forest regions 

(Wylie 1951, 1954, Jackson 1967 a, b, Givnish 1988, 

Lee et al. 1990), little has been done that examines these 

relationships for mixed-dipterocarp forests. However, 

there is some evidence that suggests the same leaf 

94 

anatomical and morphological trends exist for mixeddipterocarp 

forest. 

For species belonging to the same cladistic group 

or regeneration guild work has been equally negligible 

in mixed-dipterocarp forest. In a seedling study of 

Shorea by Ashton and Berlyn (1992) data show that 

differences in net photosynthesis (P N ), transpiration (E), 

and stomatal conductivity (g) can be associated with 

differences in the anatomy of Shorea species. General 

trends indicate that in experimentally controlled 

conditions maximum P N rate was a good measure of the 

light tolerance of Shorea. The shade tolerant species had 

maximum P N rates at relatively lower light intensity 

compared to that of more light demanding species. Ratios 

between rates of P N and E of species at their maximum 

P N light intensities can also suggest trends in water-use 

efficiency. This can reveal some indication of species 

order in relation to drought tolerance in controlled 

environments. Differences in physiological attributes 

also suggest that the greatest plasticity of response to 

differences in availability of light was exhibited by the 

most light-demanding species and the least by the most 

shade-tolerant. At a regional scale, Mori et al. (1990) 

showed similar patterns with dipterocarps. Those from 

more seasonal climates having greater rates of P N and E, 

and higher levels of plasticity than dipterocarps from 

aseasonal everwet climates. 

An array of anatomical characteristics can, in 

combination, partly determine the physiological light and 

drought tolerance of Shorea species in relation to their 

associates. Patterns suggest stomatal frequency is a 

factor differentiating Shorea species, with the most 

tolerant having fewer and smaller stomates than the most 

intolerant forms. Differences in thickness of the whole 

leaf blade and the leaf cuticle among species appear 

similarly related to both light and drought tolerance; with 

sun loving species having thicker dimensions of both 

characters than shade tolerant or demanding species. 

These results elucidate some of the relationships between 

the distribution patterns of Shorea species across the 

topography and their differences in light and drought 

tolerance. They also show that an important period 

determining site specialisation of a dipterocarp species 

occurs during regeneration establishment. Another area 

of study related to the anatomy and physiology of 

seedlings is tissue chemistry (foliar nutrients, secondary 

compounds). Although little work has examined tissue 

chemistry, investigations along these lines would tie in 

closely with studies on soil fertility, seedling herbivory


and seedling physiology that have been done at larger 

scales or from other disciplinary perspectives. 

In summary, much more work has yet to be done 

that clarifies relationships among similar or related 

species such as the dipterocarps. This work should also 

strive to link structure and physiology to seedling growth 

and mortality to gain a better mechanistic understanding 

of regeneration establishment. Research on seedling 

ecology of mixed-dipterocarp forest is substantial 

compared to other tropical forest regions. However, our 

knowledge of dipterocarp seedling ecology is 

fragmented and poor compared to other commercially 

important timber families such as Fagaceae (oak, 

chestnut, beech) where knowledge is fairly 

comprehensive for most Fagaceous forest regions. We 

have a long way to go! 


I would like to thank Peter Becker (Universiti Brunei 

Darussalam) and Ian Turner (National University of 

Singapore) for comments and suggestions for the improvement 

of this chapter. 

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Root Symbiosis 

and Nutrition 

S.S. Lee 

At present dipterocarps are gaining much attention, this 

volume being testimony to it. Since large tracts of 

dipterocarp forests in tropical Asia have become 

overlogged and/or degraded, interest in planting 

dipterocarps either in plantations or by underplanting in 

poor forests has gained momentum. With this move, 

research on mycorrhizas and their association with 

dipterocarps has gained a high profile. 

Mycorrhizas are the symbiotic association between 

specialised root-inhabiting fungi and the roots of living 

plants. Harley and Smith (1983) recognise seven 

mycorrhizal types but only two, the ectomycorrhizas and 

the vesicular-arbuscular mycorrhizas (VAM) (now more 

popularly referred to as arbuscular mycorrhizas) occur 

in the Dipterocarpaceae. Dipterocarps are predominantly 

ectomycorrhizal but a few species have been reported 

to form both ectomycorrhizas and VAM (Table 1). Unlike 

some members of the Leguminosae, the dipterocarps are 

not symbiotic with nitrogen fixing bacteria. 

Typical dipterocarp ectomycorrhizas are short, 

pyramidal or racemously branched and variously 

coloured (e.g. brown, black, white, yellow). A fungal 

sheath (mantle) characteristic of the fungal partner 

surrounds the host root. Underneath this sheath lie the 

often radially elongated epidermal cells between which 

are located the hyphae of the Hartig net (Alexander and 

Högberg 1986, Lee 1988). The surface of the sheath may 

be smooth but often bears hyphae or hyphal strands which 

radiate out into the substrate. 

The role of mycorrhizas in increasing the absorptive 

efficiency of roots is well known. The growth of 

mycorrhizal hyphae into the surrounding soil effectively 

shortens the distance over which the slowly diffusible 

ions, such as phosphate, must travel before being 

absorbed and the association has proven particularly 

Chapter 6 

beneficial to the host in soils of low available phosphorus 

concentrations. Ectomycorrhizas are also seen to play a 

role in minimising nutrient losses from the nutrient cycle 

through leaching (Read et al. 1989). The production of 

a potent acid carboxypeptidase by some ectomycorrhizal 

fungi such as Amanita and Boletus (Read 1991) indicates 

that these fungi have the potential to mobilise the plant 

growth limiting nutrient, nitrogen, from protein. This 

implies that such ectomycorrhizal infected trees are no 

longer dependent upon the activities of a separate group 

of decomposer fungi for the release of nitrogen in the 

form of the ammonium ion for plant uptake. 

Ectomycorrhizas are also known to be able to increase 

the tolerance of trees to drought, high soil temperatures, 

organic and inorganic soil toxins, and very low soil pH. 

The sheath has been shown to have important storage 

functions, not only for phosphorus but also for other 

absorbed nutrients and carbon. The sheath also protects 

the root from pathogens, and is thought to be able to 

reduce water loss and allow rapid rewetting, thus 

lengthening root life and thereby increasing mineral 

uptake and retention (Janos 1985). It has also been 

suggested that the key role of the mycorrhizal symbiosis 

under natural conditions is to enable seedling persistence 

rather than rapid growth (Abuzinadah and Read 1989). 

The presence of ectomycorrhizas in the 

Dipterocarpaceae has led to several hypotheses regarding 

the role they might play in dipterocarp biology. Ashton 

(1982) suggested that the clumped distribution of the 

dipterocarps might be reinforced by their 

ectomycorrhizal associations as the mycelia persist and 

gradually spread with the ever dispersing and coalescing 

clumps of the dipterocarp trees themselves. He suggested 

that his observation of the association of two different 

groups on soils of different soil phosphorus levels could

Root Symbiosis and Nutrition 

Table 1. Dipterocarp species reported to be ectomycorrhizal based on root examination. (Only the first report for the 

species in each location is given). 

Genera Species Location Vegetation Reference/Source 

Anisoptera 

A. costata Korth. *(VAM also) Thailand Dry deciduous forest Aniwat (1987) 

A. laevis Ridl. Pen. Malaysia Lowland rainforest Singh (1966) 

A. marginata Korth. Kalimantan Lowland rainforest Smits (1987) 

A. oblonga Dyer Pen. Malaysia Rainforest Mohd. Noor (1981) 

A. scaphula (Roxb.) Pierre " " " 

A. thurifera (Blco) Bl. Luzon Rainforest Zarate et al. (1993) 

Cotylelobium 

C. malayanum Sloot. Pen. Malaysia Dipterocarp arboretum Hong (1979) 

C. scabriusculum Brandis Sri Lanka Lowland rainforest de Alwis & Abeynayake (1980) 

Dipterocarpus 

D. alatus Roxb. Thailand Semi-evergreen forest Aniwat (1987) 

D. baudii Korth. Pen. Malaysia Rainforest Mohd. Noor (1981) 

D. chartaceus Sym. Pen. Malaysia Dipterocarp arboretum Hong (1979) 

D. confertus Sloot. Kalimantan Lowland rainforest Smits (1992) 

D. cornutus Dyer " " Bimaatmadja in Hadi et al. (1991) 

D. costatus Gaertn. f. Thailand Semi-evergreen forest Aniwat (1987) 

D. costulatus Sloot. Pen. Malaysia Rainforest Mohd. Noor (1981) 

D. elongatus Korth. Kalimantan Lowland rainforest Smits (1992) 

D. gracilis Bl. " " " 

D. grandiflorus (Blco) Blco " " Smits (1983) 

D. hasseltii Bl. " " Smits (1992) 

D. hispidus Thw. Sri Lanka Lowland rainforest de Alwis & Abeynayake (1980) 

D. humeratus Sloot. Kalimantan " Smits (1992) 

D. indicus Bedd. India Wet evergreen forest Alexander & Hogberg (1986) 

D. intricatus Dyer. Thailand Dry deciduous forest Aniwat (1987) 

D. kunstleri King Sarawak Kerangas Alexander & Hogberg (1986) 

D. oblongifolius Bl. Pen. Malaysia Lowland rainforest Singh (1966) 

D. obtusifolius Teysm. ex Miq. Thailand Dry deciduous forest Aniwat (1987) 

D. sublamellatus Foxw. " " " 

D. tempehes Sloot. Kalimantan Lowland rainforest Smits (1992) 

D. tuberculatus Roxb. Thailand Dry deciduous forest Aniwat (1987) 

D. verrucosus Foxw. Pen. Malaysia Rainforest Mohd. Noor (1981) 

D. zeylanicus Thw. Sri Lanka " de Alwis & Abeynayake (1980) 

Dryobalanops 

D. aromatica Gaertn. f. Pen. Malaysia Lowland rainforest Singh (1966) 

" Kalimantan Lowland rainforest Smits (1992) 

D. keithii Sym. " " " 

D. lanceolata Burck Java Dipterocarp arboretum Nuhamara et al. in Hadi et al. 

(1991) 

" Sabah Lowland rainforest Unpublished data 


D. oblongifolia Dyer Pen. Malaysia Dipterocarp arboretum Hong (1979) 

D. oocarpa Sloot. Kalimantan Lowland rainforest Bimaatmadja in Hadi et al. (1991) 

100


Table 1. (continued) Dipterocarp species reported to be ectomycorrhizal based on root examination. 


Hopea 

H. bancana (Boerl.) Sloot. Java Dipterocarp arboretum Nuhamara et al. in Hadi et al. 

(1991) 

H. dryobalanoides Miq. Kalimantan Lowland rainforest Smits (1992) 

H. ferrea Laness. Pen. Malaysia Rainforest Mohd. Noor (1981) 

" Thailand Semi-evergreen forest Aniwat (1987) 

H. ferruginea Parijs Pen. Malaysia Lowland rainforest Singh (1966) 

H. iriana Sloot. ? ? Ashton (1982) 

H. jucunda Thw. Sri Lanka Lowland rainforest de Alwis & Abeynayake (1980) 

H. mengerawan Miq. Kalimantan Lowland rainforest Smits (1992) 

H. montana Sym. Pen. Malaysia Rainforest Mohd. Noor (1981) 

H. nervosa King " " " 

" Sabah Lowland rainforest Unpublished data 


H. nudiformis Thw. Java Dipterocarp arboretum Setiabudi in Hadi et al. (1991) 

H. odorata Roxb. Pen. Malaysia Rainforest Mohd. Noor (1981) 

" Thailand Semi-evergreen forest Aniwat (1987) 

" Java Dipterocarp arboretum Nuhamara et al. in Hadi et al. 

(1991) 

H. parvifolia (Warb.) Sloot. S. India Wet evergreen forest Alexander & Hogberg (1986) 

H. plagata (Blco) Vidal Luzon Rainforest Zarate et al. (1993) 

H. sangal Korth. Kalimantan Lowland rainforest Julich (1985) 

Marquesia 

M. acuminata Gilg. Zambia Miombo Hogberg & Piearce (1986) 

M. macroura Gilg. " " " 

Monotes 

M. africanus (Welw.) A.D.C. Zambia Miombo Hogberg & Piearce (1986) 

M. elegans Gilg. Tanzania Miombo Hogberg (1982) 


(Balanocarpus) 

N. heimii (King) Ashton Pen. Malaysia Lowland rainforest Singh (1966) 

Parashorea 

P. densiflora Sloot. & Sym. Pen. Malaysia Lowland rainforest Mohd. Noor (1981) 

P. lucida (Miq.) Kurz. " " " 

P. malaanonan (Blco) Merr. Sabah Lowland rainforest Unbubl. data 

Pentacme 

P. contorta (Vidal) Merr. & Rolfe Philippines ? Tupas & Sajise (1976) 

P. siamensis (Miq.) Kurz. Pen. Malaysia Rainforest Mohd. Noor (1981) 

Shorea 

S. academia (?) Kalimantan Nursery Ogawa (1992a) 

S. acuminata Dyer Pen. Malaysia Rainforest Mohd. Noor (1981) 

S. affinis (Thw.) Ashton Sri Lanka Lowland rainforest de Alwis & Abeynayake (1980) 

S. assamica Dyer Pen. Malaysia Rainforest Mohd. Noor (1981) 

S. balangeran (Korth.) Burck Kalimantan Lowland rainforest Smits (1987) 

101




S. bracteolata Dyer Pen. Malaysia Rainforest Mohd. Noor (1981) 

" Kalimantan Logged over forest Suhardi et al. (1992) 

" *(VAM also) Pen. Malaysia Lowland rainforest Norani (pers. comm.) 

S. compressa Burck Java Dipterocarp arboretum Nuhamara et al. in Hadi et al. (1991) 

S. curtisii Dyer ex King Pen. Malaysia Lowland rainforest Singh (1966) 

S. dasyphylla Foxw. " " Lee (1992) 

S. faguetiana Heim Kalimantan Lowland rainforest Smits (1992) 

S. foxworthyi Sym. Pen. Malaysia Rainforest Mohd. Noor (1981) 

S. glauca King " " " 

S. guiso (Blco) Bl. " " " 

S. henryana Pierre Thailand Semi-evergreen forest Aniwat (1987) 

S. hypochra Hance Pen. Malaysia Rainforest Mohd. Noor (1981) 

S. javanica K. & V. Indonesia Agroforestry area Nuhamara in Supriyanto et al. 

(1993a) 

S. johorensis Foxw. Kalimantan Lowland rainforest Smits (1992) 

S. laevis Ridl. Pen. Malaysia Rainforest Mohd. Noor (1981) 

" Kalimantan Lowland rainforest Julich (1985) 

S. lamellata Foxw. " " Smits (1992) 

S. lepidota (Korth.) Bl. Pen. Malaysia Lowland rainforest Berriman (1986) 

S. leprosula Miq. Pen. Malaysia Lowland rainforest Singh (1966) 

" Kalimantan Lowland rainforest Bimaatmadja in Hadi et al. (1991) 

S. macrophylla (de Vriese) Ashton Sarawak ? Chong (1986) 

S. macroptera Sloot. Pen. Malaysia Lowland rainforest Singh (1966) 

S. maxwelliana King " " Becker (1983) 

S. mecistopteryx Ridl. Indonesia ? Hadi et al. (1991) 

S. obtusa Wall. Thailand Dry deciduous forest Aniwat (1987) 

S. ovalis (Korth.) Bl. Pen. Malaysia Lowland rainforest Singh (1966) 


S. ovata Dyer ex Brandis Pen. Malaysia Rainforest Mohd. Noor (1981) 

S. palembanica Miq. Java ? Hadi et al. (1991) 

S. parvifolia Dyer Pen. Malaysia Rainforest Mohd. Noor (1981) 


S. pauciflora King Pen. Malaysia Lowland rainforest Singh (1966) 


S. pinanga Scheff. Java Dipterocarp arboretum Nuhamara et al. in Hadi et al. (1991) 


S. platyclados Sloot. ex Foxw. Pen. Malaysia Rainforest Mohd. Noor (1981) 

S. polyandra Ashton Kalimantan Lowland rainforest Smits (1992) 

S. robusta Gaertn. f. India Moist deciduous forest Bakshi (1974) 

S. roxburghii G. Don Thailand Dry deciduous forest Aniwat (1987) 

S. scabrida Sym. Sarawak Kerangas Alexander & Hogberg (1986) 

S. selanica (Lamk.) Bl. Java Dipterocarp arboretum Nuhamara et al. in Hadi et al. (1991) 

S. seminis (de Vriese) Sloot. " " Hibau in Hadi et al. (1991) 

S. sericeiflora Fisher & Hance Pen. Malaysia Rainforest Mohd. Noor (1981) 

102




S. siamensis Miq. Thailand Dry deciduous forest Aniwat (1987) 

S. smithiana Sym. Kalimantan Lowland rainforest Bimaatmadja in Hadi et al. (1991) 

S. stenoptera Burck Java Dipterocarp arboretum Setiabudi in Hadi et al. (1991) 


S. sumatrana (Sloot. ex Thor.) Sym. Pen. Malaysia Rainforest Mohd. Noor (1981) 

S. talura Roxb. " " " 

S. teysmanniana Dyer ex Brandis " " " 

Vatica 

Vatica sp. 1 Kalimantan Lowland rainforest Smits (1992) 

V. chartacea Ashton " " " 

V. papuana Dyer ex Hemsl. Pen. Malaysia Lowland rainforest Singh (1966) 

V. rassak (Korth.) Bl. Kalimantan Lowland rainforest Smits (1992) 

V. sumatrana Sloot. Java Dipterocarp arboretum Hadi & Santoso (1988) 

V. umbonata (Hook. f.) Burck Kalimantan Lowland rainforest Smits (1992) 

Vateria 

V. indica L. S. India Wet evergreen forest Alexander & Hogberg (1986) 

Vateriopsis 

V. seychellarum Heim Aberdeen greenhouse Potted plant Unpublished data 

be consistent with the theory that dipterocarps are 

ectomycorrhizal. Smits (1983) suggested that the 

clumped distribution of dipterocarps was due to an 

ecological adaptation between suitable fungi and selected 

dipterocarp species on the different sites. He (Smits 

1994) has also suggested that dipterocarp mycorrhizas 

contribute to speciation amongst the Dipterocarpaceae 

through enhanced isolation. Janzen (1974) speculated 

that dipterocarps which shed their litter containing large 

amounts of phenols and other secondary compounds 

required ectomycorrhizas to avoid self-toxicity. Other 

researchers speculate that the high mortality of outplants 

and lack of success in vegetative propagation of 

dipterocarps may be due to a lack of or death of 

ectomycorrhizas (Ashton 1982, Becker 1983, Smits 

1983, Noor and Smits 1987). Lee et al. (1966) however, 

have shown that outplanted seedlings of Hopea nervosa 

and Shorea leprosula survived better and had higher 

levels of ectomycorrhizal infection in logged forest than 

in undisturbed forest. 

Although dipterocarps were known to form 

ectomycorrhizas since the 1920s (Van Roosendael and 

Thorenaar, and Voogd, cited in Smits 1992), it was only 

103 

in the last ten years that research into applied aspects of 

the dipterocarp root symbiosis, in particular its role in 

plant establishment and nutrition, has intensified, with 

the growing need for rehabilitation and reforestation. 

Mycorrhizas are also viewed as ‘biofertilisers’, an 

alternative to chemical fertilisers for infertile tropical 

soils where reforestation is being carried out (de la Cruz 

1991). This chapter discusses the current state of 

knowledge of dipterocarp nutrition and root symbiosis, 

and identifies priorities and needs for future research. 

Mycorrhizal Fungi and Dipterocarps 

Mycorrhizal Associates of the Dipterocarps 

Ectomycorrhizas are usually formed by members of the 

Basidiomycetes and Ascomycetes but in some cases may 

also be formed by Zygomycetes (species of Endogone) 

(Trappe 1962, Harley and Smith 1983). In the 

Dipterocarpaceae most observations implicate 

Basidiomycete genera. In Malaysia over fifty different 

agarics and boleti, four earthballs and a new species of 

Pisolithus have been found associated with dipterocarps 

(Watling and Lee 1995). The dominant fungi were


species of Amanita, Boletus and Russula with members 

of the Russulaceae being most numerous. Other 

researchers have also reported species of Amanita, 

Russulaceae, Boletaceae and Sclerodermataceae as 

mycorrhizal associates of dipterocarps in Malaysia 

(Becker 1983, Lee 1992). Species of Amanita, Russula, 

Boletus and Scleroderma were reported as dominant 

ectomycorrhizal fungi of dipterocarps in Indonesia (Hadi 

and Santoso 1988, Ogawa, 1992a, Smits 1994). Similar 

associations have also been reported from Sri Lanka (de 

Alwis and Abeynayake 1980). Over a six year observation 

period in Kalimantan, Indonesia, 172 fungi species from 

36 genera were found associated with 23 dipterocarp 

species with species of Amanita, Boletus and Russula 

being the dominant fungi (Yasman 1993). In the 

Philippines 32 species of ectomycorrhizal fungi from 

11 families were found associated with dipterocarps, 

with species of Russula and Lactarius predominating 

(Zarate et al. 1993). In Thailand, Aniwat (1987) reported 

species of Russula, Lactarius, Boletus, Amanita, 

Pisolithus, Tricholoma and Lepiota as the most common 

genera of ectomycorrhizal fungi in dry deciduous 

dipterocarp forest and semi-evergreen dipterocarp 

forest. Such information on the identity and diversity of 

the mycorrhizal fungi would assist in the development 

of a base for understanding the relationship between 

mycorrhizal fungi and forest function. 

It is an established fact that several different fungi 

can form morphologically different mycorrhizas on the 

root system of a single plant. Although some 

ectomycorrhizal fungi show some host specificity at the 

host genus level (Chilvers 1973), most ectomycorrhizal 

fungi generally have broad host ranges. In a recent 

experiment Yazid et al. (1994) showed that a strain of 

Pisolithus tinctorius isolated from under eucalypts in 

Brazil could form perfectly functional ectomycorrhizas 

with two species of Malaysian dipterocarps, Hopea 

helferi and H. odorata. Various studies with dipterocarps 

have shown that several different ectomycorrhizas may 

be associated with the roots of any one plant and very 

often the same mycorrhiza may be associated with 

different host species and even genera (Becker 1983, 

Yusuf Muda 1985, Berriman 1986, Lim 1986, Julich 

1988, Hadi et al. 1991, Lee 1988, 1992, Smits 1994, 

Lee et al. in press). 

Until the late 1980s there were only two published 

reports of successful isolation of indigenous 

ectomycorrhizal fungi associated with dipterocarps into 

104 

pure culture (Bakshi 1974, de Alwis and Abeynayake 

1980). However, recently successful isolations of 

several indigenous dipterocarp ectomycorrhizal fungi 

species were obtained in Indonesia, Malaysia, the 

Philippines and Thailand, from various dipterocarp hosts 

(FRIM unpublished data, Sangwanit 1993, Supriyanto et 

al. 1993a, Zarate et al. 1993). 

Inoculation and other Studies 

In most studies of the effects of mycorrhizal inoculation 

on dipterocarps reported thus far, seedlings have been 

inoculated with soil inoculum, chopped dipterocarp root 

inoculum or chopped fruit bodies. Such uncontrolled 

inoculation studies are self-limiting and non-repeatable. 

Controlled inoculation experiments with identified and 

definite fungal strains or species, in particular indigenous 

ones, are needed so that we can better understand 

dipterocarp mycorrhizal physiology and explore their 

potential for application in forestry. Spore inoculum in 

the form of capsules, tablets or powder of 

ectomycorrhizal fungi collected from the wild have also 

been used for inoculation of dipterocarps (Fakuara and 

Wilarso 1992, Ogawa 1992b, Sangwanit 1993, 

Supriyanto et al. 1993b), but these remain on a small 

scale and are dependent on fungi which fruit frequently 

and produce spores in abundance. Some progress has also 

been made in the development of controlled inoculation 

techniques for dipterocarps using mycelial pure cultures 

(Sangwanit 1993, Lee et al. 1995) but much fundamental 

research needs to be carried out before the development 

of appropriate delivery systems is explored. 

The few reports of controlled dipterocarp 

mycorrhizal inoculation and synthesis have been 

conducted with exotic fungi, mainly Pisolithus tinctorius 

(Smits 1987, Sangwanit and Sangthien 1991, 1992, Hadi 

et al. 1991, Lapeyrie et al. 1993, Yazid et al. 1994), and 

Cenococcum (Sangwanit and Sangthien 1991, 1992). In 

Indonesia, successful mycorrhizal synthesis has been 

reported between a local isolate of Scleroderma 

columnare and seedlings of Shorea stenoptera, S. 

palembanica, S. selanica, S. leprosula, Hopea 

mengerawan and H. odorata (Santoso 1989). 

Successful synthesis with Astraeus hygrometricus on 

Dipterocarpus alatus in Thailand (Sangwanit and 

Sangthien 1991, 1992) and unnamed species of Russula, 

Scleroderma and Boletus on five dipterocarp species in 

Indonesia (Hadi and Santoso 1988) has been implied on 

the basis of a growth response in inoculated seedlings.


However, no data on the nature or level of mycorrhizal 

infection were presented. The report by Louis and Scott 

(1987) of mycorrhizal synthesis in root organ cultures 

of Shorea roxburghii can be discounted as their 

illustrations and descriptions do not show 

ectomycorrhizas but hyphal invasion into root cells. 

Moreover the fungus they used, Rhodophyllus sp. was 

from a genus not normally considered to be 

ectomycorrhizal. 

The effects of various environmental factors on 

dipterocarp ectomycorrhizas and their subsequent 

effects on plant growth have been the subject of some 

recent studies. Smits (1994) suggested that the obligate 

nature and temperature sensitivity of the dipterocarp 

ectomycorrhizal relationship are the determining factors 

for good dipterocarp seedling performance. Yasman 

(1995) stated that light intensity influenced 

ectomycorrhizal formation in dipterocarp seedlings but 

that the effects varied between different host species. 

The physiology of how light regulated ectomycorrhizal 

formation was, however, not examined. According to 

Yasman (1995), neither light nor soil conditions 

represented the main factors for successful dipterocarp 

regeneration under a closed canopy; dipterocarp seedling 

survival was mainly related to the presence of 

mycorrhizal inoculum and the support of the seedlings 

by their ectomycorrhizal connections to roots from 

mother trees that had well illuminated emergent crowns. 

However, this may be an oversimplification as different 

species of dipterocarps have different light requirements 

(Mori 1980, Sasaki and Mori 1981). Lee et al. (in press) 

found high levels of ectomycorrhizal infection (60%) 

on seedlings of Hopea nervosa and Shorea leprosula 

under heavy shade in undisturbed forest supporting 

Yasman’s (1995) hypothesis, but also found that S. 

leprosula which is a light demanding species had poor 

survival compared to H. nervosa which is a shade 

tolerant species. 

Mineral Nutrition 

It must be emphasised that very few studies have been 

conducted on the very important aspect of mineral 

nutrient requirements of dipterocarps. Fertiliser trials 

have been conducted for several dipterocarp species but 

the information presently available is far from 

conclusive. Although a preliminary guide for the 

diagnosis of nutrient deficiency of tropical forest trees 

105 

has been developed (Fassbender 1988), its applicability 

and suitability for dipterocarps has to be tested more 

extensively. 

Sundralingam (1983) found that NP fertilisers 

generally improved growth of potted Dryobalanops 

aromatica and D. oblongifolia seedlings. In another 

experiment, Sundralingam and her co-workers (1985) 

found that nitrogen rather than phosphorus was the most 

important fertiliser required for improved growth of 

potted Shorea ovalis seedlings. Madius (1983) found 

that potted Shorea bracteolata and S. parvifolia 

seedlings had improved growth and increased nutrient 

uptake at higher fertiliser levels, particularly when 

moisture supply was abundant. Turner et al. (1993), 

however, found that potted Shorea macroptera seedlings 

did not respond to fertiliser application. However, they 

found that the extent of mycorrhizal infection on S. 

macroptera seedlings was correlated with seedling dry 

mass in the unfertilised control. Similarly, Burslem et 

al. (1995) working with potted Dipterocarpus kunstleri 

seedlings, found no positive growth response to nutrient 

additions although addition of P increased the 

concentrations of K and Ca in the leaves. Burslem and 

his co-workers (1995) caution that results of pot 

bioassay experiments may be dependent on factors such 

as pot size, irradiance and soil moisture conditions and 

that conclusions drawn from such experiments need to 

be tested by field fertilisation experiments. 

Turner et al. (1993) also reported that naturally 

regenerated seedlings of Shorea curtisii and Hopea 

beccariana did not show any significant response to 

fertiliser application in the field. They suggested that 

addition of nutrients to promote higher growth rates in 

regenerating seedlings in dipterocarp forests is unlikely 

to be a silvicultural practice although ensuring adequate 

infection could be beneficial. Aminah and Lokmal (1995) 

reported that outplanted rooted, stem cuttings of H. 

odorata showed a significant increase in stem diameter 

and height only nine to 24 months after application of 

granular compound fertiliser. No growth response was 

recorded when plants were measured earlier. Nussbaum 

et al. (1994) found that nutrient availability was the major 

factor limiting the establishment of Dryobalanops 

lanceolata and S. leprosula seedlings on degraded soils 

in Sabah rather than low soil moisture or high soil 

temperature. Preliminary results of the effects of NPK 

fertilisers on the growth of D. lanceolata and S. 

leprosula on a large-scale enrichment planting project


in Sabah showed that increasing concentrations of 

fertiliser resulted in increased growth rates but that 

growth was reduced when 2000 mg of NH 4 NO 3 was 

applied (Yap and Moura-Costa, in press). It will be 

interesting to see the final outcome of this large-scale 

field experiment. 

Lee and Lim (1989) found that foliar P concentration 

in naturally regenerated seedlings of Shorea curtisii and 

S. leprosula growing in a logged over forest site with 

low levels of available P (5.8 to 7.1 ppm) was significantly 

correlated with the extent of ectomycorrhizal infection. 

Lee and Alexander (1994) working with Hopea helferi 

and H. odorata found positive growth responses to 

mycorrhizal infection but variable responses to nutrient 

treatments. They also reported the first direct 

experimental evidence that ectomycorrhizal infection 

improved P uptake and growth of a dipterocarp species, 

H. odorata. Scleroderma dicstyosporum and S. 

columnare were reported to increase levels of nitrogen, 

phosphorus and potassium in seedlings of Shorea 

mecistopteryx (Supriyanto et al. 1993b) but these results 

may have been misinterpreted. Increased plant height 

growth, diameter and dry weight as well as uptake of Fe, 

Mn, Cu, Zn and Al by seedlings of Shorea compressa, S. 

pinanga, S. stenoptera, H. odorata and Vatica 

sumatrana inoculated with chopped fruit bodies of 

Russula sp., Scleroderma sp. and Boletus sp. have been 

reported in Indonesia (Santoso 1989, Santoso et al. 

1989). However, it is not clear whether ectomycorrhizas 

were formed by the test fungi or by contaminants. 

In a study of site characteristics and distribution of 

tree species in mixed dipterocarp forests in Sarawak, 

Baillie and co-workers (1987) considered phosphorus 

the most critical nutrient while magnesium was thought 

to be important because of possible effects on the 

efficiency of the dipterocarp mycorrhizal root systems. 

Some species of dipterocarps, e.g. Shorea parvifolia 

were consistently associated with sites of high P status 

while others like S. quadrinervis were associated with 

sites of low P status. Amir and Miller (1990) found 

potassium to be the primary limiting nutrient in two 

separate forest reserves in Peninsular Malaysia. Burslem 

et al. (1994), however, are of the opinion that any of the 

macronutrients and micronutrients can become 

potentially limiting to plant growth when the primary 

limitation by P is overcome. From a study of soils under 

mixed dipterocarp forest in Brunei, Takahashi et al. 

(1994) stated that logged over forests are suitable for 

106 

enrichment planting with dipterocarps since loss of soil 

nutrients and degradation of nutrient status would be small 

because of nutrient accumulation in the deeper horizons. 

It is known that different tree species have differing 

site requirements reflecting their differing abilities to 

take up nutrients from intractable soil sources due to 

differences in root system architecture and in the 

particular differences in the mycorrhizal relationships 

between species (Miller 1991). Yasman (1995) found 

that light demanding Shorea leprosula seedlings could 

form many more ectomycorrhizal types than shade 

tolerant Dipterocarpus confertus seedlings. Mineral 

nutrition, plant light requirements and mycorrhizal 

infection are very intimately related but it is only recently 

that the importance of this relationship has begun to 

receive recognition. Newton and Pigott (1991a) working 

with oak and birch found that application of fertilisers 

could reduce the number of mycorrhizal types and their 

relative abundances. Lee and Alexander (1994) found that 

full nutrient application prevented ectomycorrhizal 

formation in Hopea odorata but not in H. helferi. This 

may indirectly affect the drought tolerance of the host 

plants and consequently have implications on forest 

management. Burslem et al. (1994) suggested that 

mycorrhizas play an important role in enabling 

Melastoma to grow on very nutrient poor soils despite 

being highly nutrient demanding. They suggested that for 

mycorrhizal plants, limitation by the major cations may 

prove more significant than limitation by P. In a more 

recent study, Burslem et al. (1995) suggest that shade 

tolerant seedlings of lowland tropical forest which 

possess mycorrhizas are not limited by P supply because 

the mycorrhizas effectively relieve them of P limitation 

and/or because such plants have a low demand for 

nutrients for growth at low irradiance. 

It is clear that there is an urgent need for more 

integrated studies on dipterocarp mineral nutrient 

requirements and that such studies must take into 

consideration the role of the dipterocarp mycorrhizal 

association and the effect of different light regimes. 

While such studies are more easily conducted in the 

nursery with potted plants, there is also a need to test 

the conclusions of such experiments in the field. 


The need for more research into the dipterocarp 

mycorrhizal association is already well recognised and


is actively being pursued in Southeast Asia. However, the 

same cannot be said of research into dipterocarp mineral 

nutrition requirements. With the present interest in 

establishing plantations of dipterocarps, fertilisers are 

being applied with the hope of producing enhanced or 

more rapid growth without a clear understanding of 

dipterocarp mineral nutrition requirements. This very 

important aspect of dipterocarp silviculture needs to be 

studied in much more detail. This is reflected in the 

current state of knowledge discussed above and in the 

identification of research priorities discussed below. 

A word of caution before discussing future research 

priorities: results of many of the dipterocarp mycorrhizal 

studies carried out in this region, for example, the BIO- 

REFOR proceedings, are often difficult to interpret or 

not verifiable because of poor experimental design, lack 

of statistical analysis, or incomplete monitoring and 

reporting. Experiments need to be more carefully 

planned, controlled and monitored, to ensure that the 

observed effects are genuinely due to the inoculated 

ectomycorrhizal fungi and not from other contaminants. 

In view of the multi-faceted and some yet unknown 

aspects of dipterocarp mycorrhizas and nutrition, and the 

current efforts to establish dipterocarp plantations in the 

region, the following research priorities have been 

identified. Many paraphrase the recommendations made 

by Malajczuk et al. (undated) in their Annex 1 - 

Recommended Research Programme on Mycorrhizal 

Management, as these are found to be very relevant to 

dipterocarp mycorrhizal research. The following should 

be the future research priorities for dipterocarp 

mycorrhizas and nutrition: 

1. There is a need for more integrated studies on 

dipterocarp mineral nutrient requirements and 

mycorrhizal infection for seedling/cutting 

establishment in the field. 

Most fertiliser trials carried out thus far have ignored 

the role of mycorrhizas. They have a significant role 

to play in plant mineral uptake and are being 

considered in some quarters as possible fertiliser 

substitutes/supplements. Results from pot 

experiments have limited applicability in field 

conditions especially if plants in the field are 

interconnected by mycorrhizal links. These intact 

mycelial networks constitute the main source of 

inoculum when seedlings are grown near an 

established tree (Newton and Pigott 1991b, 

107 

Alexander et al. 1992, Yasman 1995) as is likely to 

occur in reforestation of selectively logged 

dipterocarp forests. 

2. The mycorrhizal dependency of dipterocarps for 

reforestation should be determined for each species 

at various ages in various habitats (different light 

regimes, soil nutrient levels, water retention, organic 

substrates). 

Mycorrhizal fungi like vascular plants may vary in 

their ecological and physiological requirements and 

under given circumstances, some fungi may benefit 

particular hosts more than others. The ability of a 

particular mycorrhizal fungus to enhance the foliar 

nutrient content of the host may not be indicative of 

the isolate’s ability to improve seedling growth and 

subsequent outplanting performance (Mitchell et al. 

1984). Surveys and identification of ectomycorrhizal 

fungi associated with dipterocarps should be 

continued and the results shared among workers in 

the region. 

3. Field studies should be conducted to determine the 

influence of nutrition and mycorrhizal infection on 

dipterocarp seedling survival, and their roles in 

determining forest composition. 

It has been suggested that the ‘nursing’ phenomenon 

(Read 1991), i.e. regeneration of seedlings in the 

vicinity of parent trees whereby they become 

incorporated into a mycelial network, reduces tree 

species diversity (Alexander 1989). It is believed that 

because mycorrhizal fungi have a great influence on 

plant survival in new and reclaimed sites, tree health 

and site quality, they are the cornerstone to proper 

establishment of functional forest ecosystems 

(Malajczuk et al. undated). 

4. Isolation and pure culture of indigenous 

ectomycorrhizal fungi should be intensified, and 

species associated with the desired host plant species 

both in unlogged and logged over forest requiring 

rehabilitation should be determined. 

There is evidence that some of the easily manipulated 

exotic mycorrhizal fungi such as P. tinctorius may 

be out competed by indigenous (co-evolved) 

mycorrhizal fungi in the field (see Chang et al. 1996). 

Moreover, fungi which are beneficial to the host in 

the natural forest may not be adapted to the degraded 

site where reforestation will be carried out. It has 

been suggested that successful establishment of 

indigenous ectomycorrhizal trees is limited to areas


where inoculum already exists (Alexander 1989). 

However, Smits (personal communication) reported 

that dipterocarps have been successfully established 

on a large-scale in heavily burned secondary forest 

at Longnah, East Kalimantan. 

5. The mycorrhizal fungi should be compared for effects 

on hosts in different soils under controlled 

conditions and for adaptability to handling in nursery 

inoculation processes and to nursery cultural 

practices. 

Brundrett et al. (1996) have comprehensively 

discussed the differential effect of various soil 

attributes on mycorrhizal fungal growth which have 

implications for tree establishment. 

6. Host specificity and compatibility of selected 

ectomycorrhizal fungi should be determined in pot 

experiments with selected host species and genera. 

7. Efforts on the selection of mycorrhizal fungi for 

inoculation of seedlings should be continued. This 

should be based on a set of criteria which would 

include satisfactory vegetative growth or abundant 

sporulation for production of large quantities of 

inoculum, adaptability to inoculation manipulation, 

ability to form mycorrhizas with a broad range of host 

species, and ability to enhance growth of the host 

tree (Trappe 1977, Marx et al. 1992). 

8. Inoculation experiments should be conducted with 

identified or known and preferably indigenous 

mycorrhizal strains. 

This is to ensure that results are repeatable and 

verifiable and for development into practical application 

techniques for field use. This is important for the 

sustained production of effective mycorrhizal inoculum. 

Current Mycorrhizal Research Groups 

and Needs 

Presently dipterocarp mycorrhizal research is most 

actively being pursued in Indonesia and Malaysia and to 

a lesser extent in Thailand. Some research has also 

recently begun in the Philippines. 

Indonesia 

Among the Southeast Asian nations, Indonesia has the 

most numerous researchers and research institutes 

engaged in dipterocarp mycorrhizal research. The main 

institutes are BIOTROP and Bogor Agricultural 

University in Bogor, Gadjah Mada University in 

108 

Yogyakarta, Universitas Mulawarman and the 

TROPENBOS Project in East Kalimantan which includes 

the Association of Forest Concession Holders. A variety 

of topics are being investigated but most of the results 

are published in local Indonesian journals in Bahasa 

Indonesia (see Supriyanto et al. 1993a) and often are 

very brief with details of experiments missing. This 

situation is slowly changing with the emergence of 

collaborative projects funded by foreign agencies such 

as the European Economic Community (EEC), Overseas 

Development Authority of the U.K. (ODA), the Dutch 

TROPENBOS and the Japanese government, and as more 

international symposia on mycorrhizas are organised. 

However, there appears to be some lack of coordination 

and communication between the various research groups, 

with each group appearing to work in isolation. It has 

also been pointed out that many of these groups conduct 

research in nurseries or in small experimental 

dipterocarp plantations outside the area of their natural 

occurrence (Smits 1992). Consequently not all the 

results may be of equal importance for an understanding 

of the functioning of dipterocarp mycorrhizas under 

natural conditions. Comprehensive reports of the status 

of mycorrhizal research in Indonesia are given in Hadi 

et al. (1991) and Supriyanto et al. (1993a). 

Malaysia 

In Malaysia dipterocarp mycorrhizal research is 

presently only being conducted at the Forest Research 

Institute Malaysia (FRIM). Considerable progress has 

been made towards the understanding of the biology and 

ecology of some dipterocarp mycorrhizas, and 

techniques are being developed and improved for 

controlled inoculation of dipterocarp planting material. 

The research has largely benefited from collaboration 

with researchers from Europe under a joint FRIM- 

Commission of the European Communities collaborative 

project. The survey and identification of mycorrhizal 

fungi are actively being pursued under another 

collaborative project with the Royal Botanic Garden, 

Edinburgh, funded by the ODA. Results have been 

published in several international journals. 

Thailand 

There are two institutes conducting dipterocarp 

mycorrhizal research in Thailand, these being the Faculty 

of Forestry at Kasetsart University and the Royal Thai 

Forest Department. Most of the research has


concentrated on surveys and the effectiveness of 

ectomycorrhizas in promoting growth of seedlings under 

adverse conditions. Presently dipterocarp mycorrhizal 

research is not very active and progress has been slow 

due to the limited number of researchers and funds 

available (Sangwanit 1993). 

Philippines 

Work on dipterocarp mycorrhizas in the Philippines 

started about five years ago at the University of Los 

Baños, Laguna (de la Cruz 1993) with attempts to 

combine dipterocarps propagated by cuttings/tissue 

culture and mycorrhizal inoculation. Results will be 

reported in a forthcoming publication (de la Cruz in 

press). Considerable research has been focused on the 

development of mycorrhizal inoculum delivery systems, 

mainly for use with pines and eucalypts. Some of these 

systems may be effective for dipterocarps but tests need 

to be carried out, especially under field conditions. 

Recently a survey of ectomycorrhizal fungi associated 

with pines and dipterocarps was undertaken with funding 

from the EEC (Zarate et al. 1993). 

Other Groups 

Some preliminary research on dipterocarp mycorrhizas 

has also been carried out in Sri Lanka (Abeynayake 1991). 

However, such work is not given much emphasis as 

reforestation of degraded lands with dipterocarps has not 

been successful and Sri Lanka is presently not using 

dipterocarps for reforestation on a large scale 

(Abeynayake 1991). In India some research was 

conducted on ectomycorrhizal fungi associated with 

Shorea robusta in the early 1970s (Bakshi 1974) but 

since then there have been no new reports of mycorrhizal 

research on dipterocarps. 

Mycorrhiza Network Asia 

Mycorrhiza Network Asia was established at the Tata 

Energy Research Institute, New Delhi on 1 April 1988. 

This network serves as a point of reference for 

mycorrhizal scientists in Asia and provides various 

services such as literature searches, a directory of Asian 

mycorrhizal researchers, a germplasm bank, organisation 

of meetings and symposia, and the publication of a 

quarterly newsletter, Mycorrhiza News. Mycorrhizal 

researchers from the various Southeast Asian countries 

are members or are aware of the existence of this network 

109 

and meet from time to time at the Asian Conference on 

Mycorrhizae (ACOM); the Third ACOM was held in 

Indonesia in April 1994. Previous meetings were held in 

India (1st ACOM) and Thailand (2nd ACOM). 

However, rapid progress on dipterocarp mycorrhizal 

research in the Southeast Asian region is constrained by 

several factors: 

1. Insufficient numbers of suitably trained and active 

mycorrhizal researchers in most Southeast Asian 

countries. For example, there are only two scientists 

actively working on dipterocarp mycorrhizas in 

Malaysia and Thailand respectively. 

BIOTROP has conducted several training courses on 

mycorrhizas for participants from the ASEAN 

countries but it is unfortunate that most trainees do 

not engage in mycorrhizal research upon returning 

to their own countries. A slightly different problem 

is encountered in the Philippines where many trained 

researchers leave the country for better opportunities 

abroad. In Indonesia an encouraging situation has 

recently developed where practising foresters were 

sent by their employers, the various concession 

holders, to attend a two-week local training course 

on mycorrhizal techniques conducted by BIOTROP. 

2. Insufficient budget to undertake such research. 

Most local governments do not allocate sufficient 

funds for basic research including mycorrhizal 

research. De la Cruz (1993) pointed out that much 

of the productive mycorrhizal research came from 

external grants. 

3. Lack of regional collaboration. 

Much has been spoken about the need for regional 

research collaboration in many fields, including 

mycorrhizal research, but to date no concrete 

proposals have materialised for regional mycorrhizal 

research. 

4. Lack of expertise in some fields of mycorrhizal 

research, such as identification of ectomycorrhizal 

fungal associates, culture and propagation of 

mycorrhizal inoculum. 

A local or regional flora of potential ectomycorrhizal 

fungi is needed as baseline information for many 

studies. A start has been made in several Southeast 

Asian countries to collect and collate such 

information. However, most of the research is only 

possible because of the collaboration of foreign 

experts working on short-term projects.


5. Limited access to regional research results. 

Results of many studies are reported only in local 

publications to which other researchers in the region 

have no access. Presently the most important 

channels of information are regional and international 

symposia or conferences where researchers have an 

opportunity to discuss their findings. Researchers 

should be encouraged to publish their findings in 

refereed journals or in publications with a wider 

circulation so that their results may be shared with 

others. 

Joint collaborative projects involving active 

dipterocarp mycorrhizal researchers, plant 

physiologists, and soil scientists from the various 

countries in the region and experienced scientists 

from the developed countries would be one approach 

to advancing research in this field. Training relevant 

personnel who would be likely to put their training 

into practice would also help overcome some of the 

problems encountered. It is envisaged that agencies 

such as the Center for International Forestry 

Research and the European Union can play pivotal 

roles in this respect. 

Funding Requirements 

Funding is required for a multi-lateral collaborative 

project involving scientists from related disciplines in 

the various Southeast Asian countries and experienced 

mycorrhizal researchers from the developed countries 

to conduct research into some, if not all, of the priority 

areas identified. Funding should at least be for an initial 

period of three years and should include components of 

training for local scientists and field personnel. Local 

scientists who will be directly involved in the research 

should receive relevant training in the first year of the 

project. 


I would like to thank Willie Smits (The International MOF 

TROPENBOS – Kalimantan Project) for comments and 

suggestions for the improvement of this chapter. 

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Pests and Diseases of 


C. Elouard 

Introduction 

There has been relatively little research on the pests 

and diseases of dipterocarps. Most investigations have 

been directed to forest products commensurate with 

their economic value. Now that dipterocarps are being 

established by enrichment planting in forests or in 

extensive plantations, more attention will have to be 

directed to the pests and disease problems of living 

trees. 

Pests and diseases on dipterocarps affect seeds, 

seedlings, saplings, trees and their products. A large 

proportion of earlier studies catalogued dipterocarps’ 

pests and diseases. Little is known about their ecology, 

natural enemies, management and control. The only 

well-studied species is Shorea robusta, an important 

timber species in central and northern India and grown 

in plantations. Pests have been investigated on forest 

trees only when mortality resulted in economic loss, 

as for Shorea robusta in India. There has been 

considerable work on pests of Indian dipterocarps 

(Stebbing 1914, Beeson 1941, Bagchee 1953, 1954, 

Bagchee and Singh 1954, Bhasin and Roonwal 1954, 

Bakshi 1959, Mathur and Balwant Singh 1959, 1960a, 

b, 1961a, b, Mohanan and Sharma 1991). Dipterocarp 

diseases are mainly recorded as fungal diseases. The 

only record of bacterial disease is Agrobacterium 

tumefaciens, causing leaf gall formation on saplings 

(Ardikosoema 1954, Torquebiau 1984, Smits et al. 

1991). An alga, Cephaleuros virescens, is reported 

causing leaf disease (Mittal and Sharma 1980, Elouard 

1991). 

The establishment of forest plantations and the 

enrichment planting of logged-over forests with local 

species such as dipterocarps requires collection of 

fruits, seed storage and raising of seedlings in nurseries. 

Chapter 7 

Thousands of seedlings growing in a confined place can 

lead to the development and proliferation of nonspecific 

and even specific pathogens and pests. A timber 

trend study in India (Anon. in Bakshi et al. 1967) shows 

that combined loss in forest wealth due to causes like 

fire, decay, insects and windfall is 13 per cent. This 

emphasises the need for proper integrated pest and 

disease management to protect investments. 

Pests and pathogens are present in forest 

ecosystems at all stages and take an active part in their 

ecological balance and dynamics. Though pathogen and 

pest damage is kept controlled at non-epidemic levels 

in natural forests (Augspurger 1990), logging activities 

change the natural balance of the forest ecosystems, 

and can favour proliferation of pests and pathogens. 

Moreover, enrichment planting and forest plantations 

can be a dramatic source of pest and disease 

propagation, particularly on monospecific plantations 

such as the case of the leaf blight (Microcydus ulei) of 

rubber in South America. The major epidemics recorded 

on dipterocarps are caused by insects on Shorea 

robusta, e.g. Hoplocerambyx spinicornis 

(Cerambycidae), an important heartwood borer in India 

and Pakistan, and the mealybug Drosicha stebbingi 

(Coccidae) which have caused considerable damage 

(Beeson 1941). 

The main constraints to research on dipterocarp 

pests and diseases are shortage of trained staff, lack of 

cooperation among scientists and institutions in Asia, 

inadequate funding and infrastructure facilities, high 

cost of pest and disease identification, lack of 

information on the economic effects of pests in 

plantation forestry, and need for more contacts between 

researchers, foresters and staff of timber companies.

Pests and Diseases of Dipterocarpaceae 116 

Pests 

Seeds 

Dipterocarp seeds are produced irregularly and sparsely 

in some species, and fruit production varies in quantity 

and quality from year to year. Mass fruiting appears to 

favour seed predators, but it can also be a strategy to 

escape complete seed destruction (Janzen 1974). Seed 

predation can be very high, and the crop can be 

completely wiped out. Curran and Leighton (1991) 

reported that the 1986 crop was entirely destroyed 

(100,000 seeds/ha) in the lowland forest of West 

Kalimantan. The major losses are caused by insect pests. 

Natawiria et al. (1986) observed weevils (Curculionidae) 

damaged 40-90% of the seeds of Shorea pauciflora, S. 

ovalis, S. Iaevis, S. smithiana and Dipterocarpus 

cornutus. Daljeet-Singh (1974) reported that weevils 

were responsible for more than 80% of the total seed 

damage in all case studies except Shorea macrophylla, 

in which the most important pests were the Colytidae. In 

1991, 70% of Dryobalanops aromatica seeds were 

damaged by weevils in Malaysia (Elouard, unpublished). 

While insects are the major seed pests, there is 

destruction by birds and mammals. Wild pigs (Sus scrofa), 

squirrels (Callosciurus prevostii and C. notatus) and 

monkeys (Presbytis rubicunda) caused damage to the 

crops of some species (Kobayashi 1974, Natawiria et 

al. 1986, Curran and Leighton 1991). Kobayashi (1974) 

observed that 80% of the mature seed crop of Hopea 

nervosa was damaged by squirrels. Parrots (Psittacula 

sp.) have been observed feeding on dipterocarp seeds 

(Natawiria et al. 1986). However, monkeys and squirrels 

prefer to eat other available fruit and seeds (Curran and 

Leighton 1991). Dipterocarp resin contains a high 

percentage of alkaloides and can be repellant to 

mammals. Neobalanocarpus heimii seeds are hardly 

eaten by mammals, but losses are due to the destruction 

of a part of the seed tasted by the rodents and then 

rejected (Elouard et al. 1996). 

Over 80 species of seed pests have been described 

on various dipterocarp seeds, with both pre- and postdispersal 

insect pests. The former attack the fruits on 

the tree before dispersal, whereas the latter attack fruits 

on the ground. The pre-dispersal fruit pests are weevils 

(Curculionidae) and Lepidoptera, and the post-dispersal 

ones are Lepidoptera (Toy 1988). It is rarely possible to 

distinguish between pre- and post-pests of Lepidoptera. 

The mode of attack of the weevils and Lepidoptera on 

dipterocarps is described by Daljeet-Singh (1974). The 

weevils come at the early development of the fruits, 

pierce the pericarp and deposit a single egg. The larvae 

feed on the cotyledons throughout the period of growth. 

The pupal chamber is made of larval frass. Usually, the 

fruit drops to the ground before pupation and the adult 

weevils remain within the fruit for a few days before 

emerging. They are sexually immature at emergence. The 

Lepidopteran predators lay their eggs on the dipterocarp 

fruits. On hatching, the larvae bore into the fruit, feed on 

the cotyledons and pupate. Prior to pupation, the larva 

attacks the pericarp leaving only a thin covering that the 

newly emerging adult can break. 

Toy (1988) observed in Malaysia that species of 

Nanophyes (Curculionidae) were generic specialists and 

some species appeared to be even sub-generic 

specialists. The existence of insect pests which have a 

‘familial specialisation’ raises questions on the function 

of mass-flowering as a pest satiation strategy (Janzen 

1974). The survival of these insects between fruiting 

events are ascribed to three hypotheses: i) they have either 

alternative hosts in non-dipterocarp families; ii) 

dormancy; or iii) maintain more or less continuous 

generation of pests developing on sporadically flowering 

trees (Toy 1988). In a study of Nanophyes shoreae 

survival in Shorea macroptera, Toy observed that a 

maximum 1.8% of insects survived during sporadic 

events, thus dispersal of the insect is not probable. He 

suggested generalist feeding of adults is the key to their 

persistence between fruiting events. 

Seedlings and Saplings 

Few records exist on pests of seedlings and saplings in 

nurseries, though some reports are available for natural 

forests. Insects are the main source of damage as leaf 

feeders, borers, suckers and in gall formation. The other 

pests recorded are wild boars, rodents and nematodes. 

There are few reports of leaf damage to seedlings 

and saplings (Becker 1983, Tho and Norhara 1983) and 

the defence properties of essential oils in mature leaves 

were discussed by Becker (1981). Galls causing leaf 

damage were reported on dipterocarp species in 

Singapore, Malaysia and India (Anthony 1972, 1977, 

Mathur and Balwant Singh 1959), mortality and setback


in growth by attacking the young shoots and twigs of 

Dryobalanops aromatica saplings over 1 m tall (Anon. 

in Tho and Norhara 1983). 

Shoot and root borers were recorded on various 

dipterocarp species (Beeson 1941, Chatterjee and Thapa 

1970, Daljeet-Singh 1975, 1977, Sen-Sarma and Thakur 

1986, Shamsuddin 1991, Smits et al. 1991). Shoot 

boring does not generally cause mortality (although it 

was recorded as the major factor of die-back of Shorea 

teysmanniana seedlings (Shamsuddin 1991) but rather 

induces the formation of multiple leaders after 

destroying the main shoot (Daljeet-Singh 1975, 1977, 

Smits et al. 1991). Therefore, shoot boring insects are a 

problem for reforestation programmes. Planting trials 

with Shorea ovalis, S. leprosula, S. acuminata and S. 

parvifolia were conducted in Malaysia, where 50% of 

S. acuminata and 7.3-16.5% of the other Shorea 

seedlings were attacked by shoot borers (Daljeet-Singh 

1975). 

Insect borers and nematodes can destroy roots. The 

lepidopteran root borer Pammene theristhis 

(Eucosmidae) has emerged as the most serious pest of 

the seedlings and young shoots of Shorea robusta (sal) 

in all areas where it is grown in India. It probably plays a 

prominent role in the regeneration-failure in sal. It has 

been closely associated with the dying-off of new sal 

regeneration in the submontane belt of Uttar Pradesh 

(Beeson 1941; Chatterjee and Thapa 1970). The borer 

has more than three generations a year: the first generation 

lays eggs on the seeds on which the larvae feed; the 

second one bores into young growing shoots of coppice 

or regeneration of sal up to sapling stage with the 

resultant die-back of leaders; and the third generation 

attacks and kills the young seedlings by hollowing the 

tap root and a part of the stem (Beeson 1941, Sen-Sarma 

and Thakur 1986). Nematodes were recorded feeding on 

rootlets of Hopea foxworthyi and Shorea robusta 

(Catibog 1977, Mathur and Balwant Singh 1961a). 

Wild pigs (Sus scrofa) can completely destroy 

seedling regeneration (Becker 1985, Elouard 

unpublished). Rodents can be significant as pests of 

germinating seeds and the cotyledons of young seedlings 

(Wyatt-Smith 1958) and deer browsing was partly 

responsible for mortality of Shorea robusta seedlings 

and saplings in India (Davis 1948). 

Trees 

Tree pests were recorded in Malaysia, Thailand, 

Indonesia, India, Pakistan and Burma. Most of them are 

insects belonging to Coleoptera and Lepidoptera, causing 

defoliation and leaf damage, wood boring and root 

sucking. 

The extent of the damage and the economic losses 

due to defoliation, essentially caused by insects, has 

seldom been estimated. Over 130 species of insects 

cause leaf damage, mostly belonging to the families 

Geometridae, Lymantriidae, Noctuidae, Pyralidae, 

Tortricidae (Stebbing 1914, Beeson 1941, Bhasin and 

Roonwal 1954, Ghullam Ullah 1954, Mathur and Balwant 

Singh 1959, 1960a, b, 1961a, b, Anderson 1961, 

Torquebiau 1984, Pratap-Singh and Thapa 1988, Messer 

et al. 1992). 

Defoliators in India, Pakistan, Malaysia, Indonesia, 

Thailand and Philippines, at times cause important 

damage, e.g. Shorea robusta trees in Assam, India were 

entirely stripped of all green leaves over a very large 

area by species of caterpillars of the genus Lymantria 

(Stebbing 1914). Defoliation can lead the trees to an 

extremely weak state which makes them attractive and 

highly receptive to a lethal infestation from borers such 

as Hoplocerambyx spinicornis (Pratap-Singh and Thapa 

1988). Successive defoliations can kill trees, e.g. 

Lymantria mathura on Shorea robusta in Assam and 

north India (Beeson 1941). Following defoliation, the 

physiology of the tree is affected by the loss of 

photosynthetic activity: Shorea javanica trees, tapped 

for resin in Sumatra, Indonesia, stopped their resin 

production (Torquebiau 1984). The attack by insects in 

Shorea robusta forests of Bangladesh appeared to be 

minor (Ghullam Ullah 1954). According to the author, 

this may be due to the presence of large colonies of the 

brown ant, Oecophylla smaragdina, known to destroy 

all kinds of caterpillars (except the hairy species) and to 

drive away beetles and bugs, thus preventing oviposition 

in the latter case. Ghullam Ullah noted all the Shorea 

robusta defoliating larvae are hairy caterpillars which 

are not destroyed by ants. 

The borer-fauna of Dipterocarpaceae is very 

extensive, and has been mostly recorded in India. 

According to Beeson (1941), only one species, the 

heartwood borer Hoplocerambyx spinicornis 

(Cerambycidae), is capable of killing healthy trees. The 

other borers, or secondary borers, attack sickly trees, 

possibly hastening death by a year or two. 

Hoplocerambyx spinicornis is widely distributed in 

Asia (Burma, Bhutan, India, Indo-China, Indonesia, 

Malaysia, Nepal, Papua New Guinea, Pakistan,


Philippines, Singapore, Thailand). It is a pest of 

Parashorea robusta, P. malaanonan, P. stellata, 

Shorea siamensis, S. assamica, S. obtusa, S. robusta, 

Anisoptera glabra and Hopea odorata. This insect is a 

principal pest in the Matang Forest Reserve of Sarawak, 

Malaysia, and causes severe damage in central and 

northern India on Shorea robusta. Outbreaks of this 

insect have been recorded periodically since 1897 in 

Chota Nagpur, India. The grub feeds on and destroys the 

bast layer eventually killing the tree, and it tunnels down 

into the heartwood spoiling it for commercial purposes. 

This cerambycid has the habit of destroying the trees in 

patches (Stebbing 1914). It produces characteristic 

symptoms: i) dying-off from the crown downwards by 

sudden withering of the foliage in autumn or spring; and 

ii) profuse exudation of resin at points where the first 

stage larvae bore through the bark. 

The biology of H. spinicornis, the damage caused by 

the insect and its control have been studied by Stebbing 

(1914), Beeson and Chatterjee (1925), Atkinson (1926), 

Beeson 1941, Bhasin and Roonwal 1954, Roonwal 1952, 

1976, 1977, 1978, Mathur and Balwant Singh (1959, 

1960a, b, 1961a, b), Mathur (1962), Chatterjee and Thapa 

(1964, 1970), Sen-Sarma et al. (1974), Singh et al. 

(1979), Mercer (1982), Singh and Mishra (1986), 

Bhandari and Pratap-Singh (1988) and Baksha (1990). 

The borers prefer large, mature trees, where there is more 

chance of completing the life cycle. But during 

epidemics this borer is capable of infesting every tree 

above 0.3 m girth and and is not confined to mature or 

over-mature trees. It then affects thousands of hectares 

of Shorea robusta (Sen-Sarma and Thakur 1986). The 

emergence of the adult beetle is closely synchronised 

with rainfall (June/July). The beetles lay eggs in the bark 

and sapwood and a heavily infested tree may contain as 

many as 900 living larvae. Full grown larvae tunnel into 

the heartwood and riddle it with galleries, making it unfit 

for marketing as timber (Sen-Sarma and Thakur 1986). 

Stebbing (1914) and Mathur (1962) described a method 

of trapping the insect called the ‘tree-trap’ system. During 

outbreaks, one tree per hectare is felled, and the log 

beaten to expose the inner bark. The adults, attracted by 

the inner bark, are collected by hand and destroyed. This 

method has been used since then and is successful in 

monitoring and controlling the beetle populations 

(Chatterjee and Thapa 1970, Roonwal 1978, Bhandari 

and Pratap-Singh 1988). A beetle can locate a freshly 

felled tree of S. robusta at a maximum distance of 2 km 

(Pratap-Singh and Misra 1981). 

Many of the secondary borers attack freshly felled trees, 

but can occasionally attack moribund trees and hasten 

their death. They also attack young growth in sickly 

condition due to some abiotic factors (frost or drought) 

or biotic factors (e.g. infestation by defoliators) or kill 

the trees (e.g. Massicus venustus) by mass-attack 

(Beeson 1941). Most borers are not a threat for the tree 

itself but make it useless for construction purposes and 

reduce the market value of the timber. 

Suckers, belonging to Cicalidae and Coccidae were 

reported damaging roots (Hutacharern et al. 1988) and 

leaves (Mathur and Balwant Singh 1961a). Lacifer lacca 

(Coccidae), the insect involved in the production of lac, 

is a sap sucker of Shorea talura, Shorea spp. and 

Dipterocarpus tuberculatus (Mathur and Balwant Singh 

1959, 1961a, b). 

Termite attacks have been reported on living 

dipterocarp trees (Wyatt-Smith 1958, Nuhamara 1977, 

Sen-Sarma and Thakur 1980, Smits et al. 1991). Smits 

et al. described termite attack on living Shorea 

polyandra in Kalimantan: the trees shed their leaves, 

while the crown became lighter and the death of the tree 

was manifested by the exudation of many clumps of black 

resin from the stem base. 

Forest Products 

Damage on logs and timbers are mainly caused by 

termites and beetles. Since it is a field of economic 

importance, many studies have been conducted on the 

identification of the pests, their biology and control 

methods. 

Termites attacking logs and wood were studied in 

Malaysia, Indonesia, India, China (Mathur and Balwant 

Singh 1960a, b, 1961a, b, Becker 1961, Sen-Sarma 1963, 

Abe 1978, Sen-Sarma and Gupta 1978, Hrdy 1970, Said 

et al. 1982, Ping and Xu 1984, Dai et al. 1985, 

Quiniones and Zamora 1987, Hutacharern et al. 1988), 

but also in European and even Saudi Arabian laboratories 

(Alliot 1947, Badawi et al. 1984, 1985). Tests on the 

resistance of wood to termite attacks were widely 

conducted (Alliot 1947, Becker 1961, Sen-Sarma 1963, 

Schmidt 1968, Sen-Sarma and Gupta 1978, Hrdy 1970, 

Dai et al, 1985, Badawi et al. 1984, 1985). Pentacme 

suavis, Shorea guiso, S. robusta, S. obtusa, S. 

stenoptera, Vatica astrotricha, Hopea hainanensis, 

Dipterocarpus sp. proved to be resistant to termite 

attack. In other studies wood from Dipterocarpus spp.


was particularly susceptible to termite attack (Alliot 

1947) and that of Vateria indica was preferred by 

Microcerotermes cameroni (Hrdy 1970). Heavy 

hardwoods, Neobalanocarpus heimii and Vatica sp., 

were the least susceptible species to termite attack. 

Wood of Neobalanocarpus heimii, Shorea ovalis and 

Shorea spp. contain repellants against Cryptotermes 

cynocephalus (Said et al. 1982). 

Ambrosia beetles (pin-hole borers) infest logs and 

wood timber (Browne 1950, Bhatia 1950, Anon. 1957, 

Anuwongse 1972, Fougerousse 1974, Garcia 1977, 

Hutacharern et al. 1988). Browne reported the 

susceptibility of Shorea leprosula logs to attack by 

ambrosia beetles, more particularly Xyleborus 

pseudopilifer which usually attacks only dipterocarps, 

and X. declivigranulatus which is polyphagous. Shot and 

pin-hole borers attacked barked-logs of Parashorea 

malaanonan more severely than unbarked ones, as well 

as logs left in the shade (Anon. 1957). 

Insecticide trials against termites (Mathur et al. 1965, 

Said et al. 1982, Schmidt 1968) found BHC, aldrex and 

chlordane were effective. Preservatives, such as copperchrome-arsenic, 

increased wood resistance to attack of 

Coptotermes curvignathus. 

Studies of treatment against insect damage on logs 

and wood have been mainly conducted in India. 

Insecticides such as BHC, fenpropathrim, fenvalerate, 

permethrine, telodrine, diedrex, gammexane and to a 

lesser extent chlordane were effective against beetles 

such as Lyctus brunneus (Lyctidae), Cerambycidae, 

Bostrichidae, Platypodidae and Scolytidae (Browne 

1951, Menon 1954, 1958, Francia 1958, Thapa 1970, 

Ito and Hirose 1980, Chatterjee and Thapa 1971, 

Nunomura et al. 1980, Daljeet-Singh 1983). Thapa 

(1970) showed that BHC offered a satisfactory 

protection when sprayed on logs of Parashorea 

tomentella against cerambycids and more particularly 

Dialeges pauper and Hoplocerambyx spinicornis. 

A minimum of 3 months immersion of Shorea 

robusta poles in water gives protection against bostrichid 

attack, most probably due to the leaching of sugars during 

soaking (Anon. 1946). Fresh water and marine borers 

have damaged boats and poles (Shillinglaw and Moore 

1947, Anon. 1947, Edmonson 1949, Premrasmi and 

Sono 1964, Mata and Siriban 1976, Chong 1979, 

Santhakumaran and Alikunhi 1983, Chen 1985). Most 

records concern marine borers, though nymphs of 

species of mayfly (Ephemeroptera) burrow into and 

damage boats and submerged wooden structures in fresh 

water in Thailand (Premrasmi and Sono 1964). 

The durability and resistance of dipterocarp timbers 

and poles against marine borers, mainly in the genera 

Martesia, Teredo, Nausitora, Dicyathifer, 

Bactronophorus, Baukia, Nototeredo and Limnoria, 

were studied by Shillinglaw and Moore (1947) and Mata 

and Siriban (1976). Anisoptera polyandra in New 

Guinea (Shillinglaw and Moore 1947), 

Neobalanocarpus heimii and Shorea maxwelliana 

(Chong 1979) had good natural resistance to shipworms 

and other marine borers. A. polyandra is therefore 

recommended for piling in new marine structures. 

Shorea laevifolia has been reported as being resistant 

to Martesia and Teredo species (Anon. 1947). In China, 

Chen (1985) demonstrated that the resistance to marine 

borers of hardwood is higher than that of softwood, and 

heartwood is superior to sapwood. Edmonson (1949) 

reported Martesia sp. destroyed rapidly apitong 

(Dipterocarpus sp.) and Shorea sp. in the Philippines. 

According to Santhakumaran and Alikunhi (1983), 

Shorea robusta and Dipterocarpus indicus had a very 

heavy attack whereas D. macrocarpus had a medium 

attack and D. turbinatus and Hopea parviflora had a 

moderate attack in 7-8 months by Martesia and Teredo 

species. Some treatments with creosote proved to be 

effective (Chong 1979, Mata and Siriban 1976). 

Diseases 

Seeds 

Bacteria, viruses and especially fungi cause loss of seed 

viability. Infection takes place on the tree, during the 

flowering and/or development of the fruit, on the ground 

at the fruit fall, and during the period from harvesting to 

sowing in the nursery. During these stages, seed 

contamination can occur with organisms causing diseases 

in the nursery or serving as primary inocula for decay 

organisms specific to seedlings (Mohanan and Sharma 

1991). Seeds collected from the forest floor are more 

liable to be infected by decay organisms. Fungal infection 

also occurs during seed storage, where large quantities 

of seeds in containers and high moisture are propitious 

conditions for fungal development. 

Over 100 species of seed fungi have been identified 

in Malaysia (Hong 1976, 1981a, Lee and Manap 1983, 

Elouard and Philip 1994), in Thailand (Pongpanich 

1988), in Indonesia (Elouard 1991), and in India (Mittal


and Sharma 1982, Mohanan and Sharma 1991). Most of 

these fungi belong to Fungi Imperfecti 

(Deuteromycetes). Though a large number of species are 

recorded on dipterocarp seeds, their disease transmission 

and seed degradation is not well documented. In general, 

poor seed storage conditions affect seed quality and 

facilitate fungal infection and spread of fungi (see 

Chapter 4). There have been few fungicidal studies on 

stored dipterocarp seeds and there is a need for seed 

pathology research to establish suitable control methods 

for fungal infection both during storage and in nurseries. 

Two categories of seed fungi can be identified, the 

storage fungi and the seedborne fungi. The first category 

includes saprophytic fungi growing on the seed testa, and 

the second refers to pathogenic fungi developing from 

the internal part of the seed. Both cause significant 

damage during storage. 

Storage fungi 

Storage fungi grow fast, developing from the ever-present 

spores in the air or on the seed testa. They rapidly invade 

the embryo, causing damage and decreased germination 

(Neergard 1977). These saprophytes do not feed on the 

seeds, but their excessive development leads to the 

rotting of the seeds. The most common species belong 

to the genera of Aspergillus, Penicillium, Pestalotia, 

Pestalotiopsis, Gliocladium, Fusarium, 

Cylindrocladium and Lasiodiplodia. Most of these fungi 

produce enormous quantities of spores spreading rapidly 

and infecting the whole seed stock. 

Aspergillus niger was widely recorded on 

dipterocarp seeds (Pongpanich 1988, Singh et al. 1979, 

Mittal and Sharma 1981, 1982, Hong 1976, 1981a, Lee 

and Manap 1983, Hadi 1987, Elouard and Philip 1994). 

In India, fungicidal trials were conducted on fungi 

infecting Shorea robusta seeds, namely Aspergillus 

niger, Penicillium albicans, P. canadense, 

Cladosporium cladosporioides, C. chlorocephalum 

and Rhizopus oryzae (Mittal and Sharma 1981). 

Brassical, Bavistin and Dithane-45 proved effective. In 

Malaysia, Elouard and Philip (1994) tested fungicides 

on Hopea odorata seeds, and Benlate 50 and Thiram 

were effective without preventing germination or 

affecting seedling development. 

Seed-borne fungi 

Seed-borne fungal infection most probably takes place 

during the flowering period or at the early stage of 

fructification. The infection occurs through spores 

present in the environment or through inoculation of 

spores or mycelium by pollinating insects or predispersal 

insect predators while laying their eggs. Seed-borne 

fungi feed on living tissues, destroying the embryo and 

the cotyledons. The mycelium develops inside the seed 

and eventually covers the whole fruit. In natural stands, 

seed destruction is mainly caused by seed-borne fungi. 

The most common seed-borne fungi belong to the 

genera Fusarium, Cylindrocladium, Lasiodiplodia, 

Colletotrichum, Curvularia and Sclerotium (Hong 

1976, 1981a, Lee and Manap 1983, Charlempongse et 

al. 1984, Pongpanich 1988, Mohanan and Sharma 1991, 

Elouard 1991, Elouard and Philip 1994). The 

Basidiomyceteae Schyzophyllum commune has been 

observed on several dipterocarps (Hong 1976, Vijayan 

and Rehill 1990, Elouard and Philip 1994), developing 

on the cotyledons and embryo and ultimately covering 

the whole seed and producing carpophores. Infection 

leads to high levels of mortality: 70% of Shorea 

leprosula and S. ovalis seeds were rotted by a Fusarium 

species and 90% of Shorea glauca seeds were destroyed 

by Schyzophyllum commune (Elouard and Philip 1994). 

Seedlings and Saplings 

Over 40 species have been identified causing seedling 

diseases. The most common are in the genera 

Colletotrichum, Cylindrocladium, Fusarium and 

Lasiodiplodia, which are responsible for damping-off, 

wilting, root and collar rots, cankers, leaf diseases, thread 

blights and gall formation. 

Damping-off is the rotting of seeds and young 

seedlings at soil level (Hawksworth et al. 1983) and is 

in most cases caused by seed-borne fungi (Hong 1981a, 

Lee and Manap 1983, Pongpanich 1988, Elouard l991, 

Elouard and Philip 1994). Collar rot, root rot and wilting 

(loss of turgidity and collapse of leaves (Hawksworth et 

al. 1983)) are mainly caused by Fusarium species 

(Foxworthy 1922, Thompson and Johnston 1953, Hong 

1976, Lee and Manap 1983, Elouard, l991, Elouard and 

Philip 1994). 

A canker is a plant disease in which there is sharplylimited 

necrosis of the cortical tissue (Hawksworth et 

al. 1983). Though most of the time stem cankers are not 

lethal, they still can be harmful decreasing the strength 

of the stem and causing it to fracture. Root and collar 

cankers can affect the vascular system of the plant and 

eventually result in plant death by wilting (Spaulding


1961, Elouard unpublished). Schyzophyllum commune 

has been reported as causing die-back of young saplings 

of Shorea robusta, cankers caused by frost or fire 

providing the route of entry. The fungus, once established, 

attacks the living sapwood killing the stem beyond the 

scars, and it progresses both up and down the stem 

(Bagchee 1954). 

Various fungi cause leaf diseases, an infection of 

leaves characterised by spots, necrosis and leaf fall 

(Hawksworth et al. 1983), and most of them belonging 

to Imperfect Fungi (Deuteromycetes). In most cases, 

growth of seedlings and saplings is not affected, except 

when large spot areas (dead and necrosed cells) 

significantly reduce the leaf area for photosynthesis. The 

weakened plant becomes more susceptible to pathogen 

and pest attacks, or is less competitive with other 

seedlings and saplings in natural stands. Ultimately, the 

leaf becomes completely necrosed and dry and falls. On 

seedlings and young saplings, the defoliation can 

eventually lead to death (Hong 1976, Mridha et al. 1984, 

Charlempongse 1988, Harsh et al. 1989, Elouard l99l, 

Zakaria personal communication, Elouard unpublished). 

Some fungi, such as Meliola sp. develop a dense dark 

mat on the leaf surface, sometimes entirely covering the 

leaf area. Though the hyphae do not penetrate the leaf 

cells, chlorophyll development is hindered (Elouard 

l991). An alga, Cephaleuros virescens 

(Trentepholiaceae), was also recorded causing leaf 

disease on seedlings and saplings in India, Indonesia and 

Malaysia and on trees in India (Mittal and Sharma 1980, 

Elouard 1991). 

Thread blights recorded on dipterocarps are caused 

by Basidiomyceteae of the genera Marasmius and 

Corticium. There are two kinds of thread blights, white 

and dark. The white thread blights are produced by the 

development of whitish mycelium sticking on the twigs, 

branches and foliar system of the seedlings and saplings. 

The black thread blights are horse hair-like and attached 

to the host by byssus. The threads do not stick to the 

host’s organs except by the byssus, but develop an aerial 

network which, when too excessive, can hinder the host’s 

development. These fungi were observed in plantation 

and natural forests in India, Indonesia and Malaysia 

(Symington 1943, Bagchee 1953, Bagchee and Singh 

1954, Spaulding 1961, Smits et al. 1991, Elouard 1991, 

Elouard unpublished). 

Gall formation on shoots of seedlings and saplings 

has been described in Shorea javanica plantations in Java 

(about 60% of the seedlings affected), man-made 

dipterocarp forests of Sumatra and on Shorea spp. and 

Upuna borneensis (100% of the plants affected in 

nursery) in Kalimantan (Ardikoesoema 1954, Torquebiau 

1984, Smits et al. 1991). This gall formation is 

commonly attributed to a bacterium, Agrobacterium 

tumefaciens. According to Smits et al. (1991), the 

youngest leaf remains smaller than the leaves developed 

before infection, subsequent leaves no longer develop 

from the top shoot and all buds in the zone with green 

leaves produce side buds. This process continues until a 

dense clump of tiny shoots is produced at the buds’ 

positions but without development of any normal shoots 

from these clumps. The plant growth is then stopped. An 

insect is suspected to be the vector for this bacteria 

(Torquebiau 1984, Smits et al. l991). 

Trees 

About 150 fungal species have been recorded on trees, 

mainly causing rots and decay. In addition, leaf damage, 

flower necrosis and cankers were also reported. 

Parasitic plants of the family Loranthaceae have severely 

damaged Shorea robusta in India. 

Leaf disease on trees is harmful if the damaged area 

covers a large area of the foliar system. The fungal leaf 

diseases are mainly caused by species of Asterina, 

Capnodium, Cercospora, Colletotrichum 

(Thirumalachar and Chupp 1948, Bagchee 1953, Bagchee 

and Singh 1954, Chaves-Batista et al. 1960, Spaulding 

1961, Bakshi et al. 1967-1972, Elouard 1991). 

Cankers and rots were recorded on various 

dipterocarp species in Peninsular Malaysia, Thailand, 

Singapore, Indonesia and India (Bagchee 1954, 1961, 

Bagchee and Singh 1954, Bakshi 1957, 1959, Bakshi et 

al. 1967, Panichapol 1968, Hong 1976, Charlempongse 

1985, Kamnerdratana et al. 1987, Corner 1987, l991, 

Elouard 1991). 

Few fungal species are able to attack healthy trees. 

Aurificaria [Polyporus] shoreae, a fungus only reported 

on Shorea robusta, is capable of infecting healthy and 

uninjured roots, causing root rot and bark and sapwood 

decay. The disease results in top die back and death of 

trees (Bakshi and Boyce 1959). Most of fungal species 

are secondary parasites infecting the trees through 

wounds and are distinguished from the primary parasites 

which produce active root and stem rot. According to 

Bagchee (1954), at least 24 species of Hymenomycetes 

behave as facultative parasites of Shorea robusta.


Infection by heart-rot fungi on hardwood trees occurs 

through initial injuries caused by human activities (e.g. 

tapping), fire, drought, frost and other mechanical causes. 

These fungi establish themselves when the trees are 

either young or overmature. Most of these fungi live as 

saprophytes in jungle slash and become parasites when 

conditions for infection are favourable (Bagchee 1954). 

Trees with heart-rot can exhibit all the outward signs of 

healthy and vigorous growth. Heartwood is progressively 

decayed with age. Heart-rot in Shorea robusta can cause 

much loss of timber (e.g., 9-13%, with nearly 73% of 

the trees infected) (Bakshi et al. 1967). Bagchee (1954) 

reported that nearly 80% of trees with deformities have 

fungus-rot in their stems. Of Shorea javanica trees 

tapped for resin in Sumatra, 10% showed carpophore 

development on their trunks, indicating advanced 

infection (Elouard 1991). The fungi entering through the 

butt-scars and causing root damage contribute to 

windthrown trees (Bakshi and Boyce 1959). Infection 

by rot fungi is more frequent in the suppressed trees in 

overcrowded forests than in the trees of thinned coupes 

(Bagchee 1954). 

Flower destruction and seed abortion may be a 

serious problem for seed production under forest 

management. However, there has been little research, and 

the only record is Curvularia harveyi on Shorea 

pinanga in Indonesia (Elouard 1991). 

Parasitic plants, belonging to Loranthaceae, were 

observed on Shorea robusta in India and Bangladesh 

(Davidson 1945, Singh 1954, Ghosh 1968, Alam 1984) 

and on S. obtusa in Thailand (Charlempongse 1985). The 

parasites caused serious damage although the trees did 

not die (Davidson 1945, Alam 1984). The trees tended 

to form epicormic branching in some of the older 

plantations. The only method of controlling infestations 

of Loranthus appears to be eradication by lopping in the 

cold weather (De 1945). 

Forest Products 

Diseases on forest products are primarily wood decay 

and staining fungi (Bagchee and Singh 1954, Banerjee 

and Sinhar 1954, Sivanesan and Holliday 1972, Hong 

1980a, b, Shaw 1984, Balasundaran and Gnanaharan 1986, 

Supriana and Natawiria 1987, Kamnerdratana et al. 

1987). Most of them belong to the Basidiomyceteae and 

can be categorised as white rot, brown rot and soft rot. 

In white rot, both lignin and cellulose are attacked. In 

brown rot, cellulose and hemicellulose are attacked while 

lignin remains unaffected. In soft rot, cellulose is 

removed like brown rot but the mechanism of action on 

cell walls is different. The fungi causing soft rot belong 

to Ascomycetes and Fungi Imperfecti and are restricted 

to hardwoods (Supriana and Natawiria 1987). Decay of 

timber occurs mostly after felling, on wood in service 

and on industrial wood products. Likewise, on logs and 

poles an important number of wood decay fungi have been 

identified and control methods investigated. Most of 

these fungi are weak pathogens, though some can also 

infect living trees, e.g., Hypoxylon mediterraneum 

recorded both on trees and wood attacking Shorea 

robusta trees and hastening their death or preventing 

recovery (Boyce and Bakshi 1959). 

Decay fungi affect boats (Premrasmee 1956, Savory 

and Eaves 1965) and wall framing (Singh 1986). One of 

the most common decay fungi is Schyzophyllum 

commune recorded in India, Indonesia, Thailand and 

Philippines (Bakshi 1953, Bagchee and Singh 1954, 

Mizumoto 1964, Supriana and Natawiria 1987, 

Charlempongse 1985, Quiniones and Zamora 1987). 

Various dipterocarp species, Shorea elliptica, S. 

hypoleuca and S. laevis are highly resistant to 

Chaetomium globosum (soft rot) and Trametes 

[Coriolus] versicolor (Takakashi and Kishima 1973) and 

Shorea siamensis is extremely durable against 

Coniophora cerebella, Trametes [Polystictus] 

versicolor and Daedalea quercina (Bavendam and 

Anuwongse 1967). Shorea guiso, Hopea parviflora and 

Vateria indica proved to be resistant to several fungal 

species (Moses 1955, Balasundaran and Gnanaharan 

1986). Veneer-faced, low-density particleboards 

including Shorea particles, tested for its resistance 

against Tyromyces palustris and T. versicolor proved to 

be resistant (Rowell et al. 1989). 

Treatments, heating, fumigants, Wolman salt, ascu and 

borax, boliden K-33 and tanalith C. were tested on various 

wood species against decay fungi. Copper-chromearsenic 

(CCA) is the most widely used preservative in 

Malaysia for wood protection, but organotins are better 

since they have a higher fungicidal activity, provide a 

higher protection against the marine toredo worm, are 

less toxic towards mammals and more easily degradable 

(Hong and Khoo 1981, Hong and Daljeet-Singh 1985). 

Wood staining fungi infect logs in logging areas and 

freshly sawn timbers in saw mills. A large amount of 

money is spent each year on preservatives to overcome 

this problem of staining (Hong 1981b). The staining does


not reduce the strength of timbers but degrades their 

quality and value (Thapa 1971, Hong 1980a, b). Stains 

can be caused by moulds, resulting in superficial staining 

easily brushed or planed-off, and sap-staining fungi 

(‘blue-stain’), producing deep penetration stains. The 

most common are Diplodia spp., Ceratocystis spp. and 

Lasiodiplodia [Botriodiplodia] theobromae (Supriana 

1976, Hong 1980a, b, Charlempongse 1985). For 

prevention and control of stain, it is best, when possible, 

to process the felled timber within 1 to 2 weeks. 

Otherwise, chemical treatment is the only way, and the 

cut ends of logs should be immediately treated. The 

chemicals most effective against black stain and mould 

include the salts of chlorinated phenols (e.g. sodium salt 

of pentachlorophenol, SPP), and organic mercury 

compounds. These chemicals, effective against stains, 

have a low efficiency on green moulds (Hong 1980a, 

1981b). 

Physiological Disorders 

Very few studies have been conducted on physiological 

disorders such as frost, drought, poor drainage and fire 

damage, except in India on Shorea robusta (Davis 1948, 

Ram-Prasad and Pandey 1987, Raynor et al. 1941, 

Griffith 1945, Anon. 1947, Bagchee 1954). 

A review of the adverse factors that probably combine 

to cause serious dieback of Shorea robusta in Uttar 

Pradesh (India) was made by Ram-Prasad and Jamaluddin 

(1985) including deficient and erratic rainfall, low 

retention of soil moisture, nutritional imbalance of the 

soil, over-exploitation, unregulated grazing, fire and 

excess of removal of fuelwood. 

Mortality of Shorea robusta seedlings and young 

saplings due to frost was mentioned (Davis 1948, Ram- 

Prasad and Pandey 1987, Raynor et al. 1941 Griffith 

1945, Anon. 1947, Bagchee 1954). Frost initiates canker 

in advanced trees usually on the border of the forest 

facing the open lands and on the banks of perennial 

streams where the precipitation is heavy as dew or hoar 

frost (Bagchee 1954). Radiation frosts, creating 

frostholes by convection currents, kill saplings, create 

cankers providing the route of entry for heart-rot fungi 

and produce a moribund type of Shorea robusta which 

ultimately becomes the object of attack by many 

parasitic fungi and pests. 

Drought is also an important cause of S. robusta 

mortality (Pande 1956, Seth et al. 1960, Gupta 1961, 

Ram-Prasad and Jamaluddin 1985, Khan et al. 1986). In 

Malaysia, Tang and Chong (1979) have reported a 

‘sudden’ mortality of Shorea curtisii seedlings due to 

moisture stress. In India, Bagchee (1954) mentioned that 

the roots of Shorea robusta must be in the region of 

permanent water zone in order to be healthy. On the other 

hand, Yadav and Mathur (1962) reported excess water 

accumulation during the rainy season caused mortality 

of S. robusta seedlings by development of white slimy 

growth on the roots and Sharma et al. (1983) reported 

deaths due to poor drainage. 

Fire, often of anthropogenic origin, can damage S. 

robusta (Joshi 1988, Ram-Prasad and Jamaluddin 1985, 

Sinha 1957, Bagchee 1954, Bakshi 1957). It results in 

deformity and other injuries to the immature trees such 

as burrs, galls, tumourous knots, cankers, and heart-rot 

fungi entering through wounds. 

Management Aspects 

There are few practical management methods directly 

available to foresters against pests and diseases attacks 

in mature dipterocarp trees. Concerning pests, the main 

record is the ‘tree-trap’ technique set up in India for 

reducing the population of Heterocerambyx spinicornis. 

Regular surveys of insect populations in forest 

plantations can help monitor the health conditions of the 

trees, and some insect species (Buprestidae, 

Bostrichidae, Cerambycidae, Scolytidae) are indicators 

of sickly trees (Stebbing 1914, Beeson 1941). So forest 

managers can identify which trees, providing shelters for 

insect breeding, should be removed to avoid a massive 

infestation of trees and logs. The infection by heart-rot 

fungi on trees can be reduced by removing the dying and 

dead trees and burning them. The danger is more 

important if the tree bears fungal fruiting bodies and is a 

source of infection (Bakshi 1956a, b). The well-known 

technique of digging trenches around the infected areas 

to isolate the infected roots and soil area can also be 

applied. 

Bakshi (1957) suggested lowering the felling age of 

the trees in forests with a high incidence of heart-rot 

and to avoid coppicing from infested stumps. Heart-rot 

in the coppice standards due to Phellinus caryophylli 

and P. fastuosus is transmitted by grafting healthy roots 

with diseased ones or with decayed woody parts 

embedded in the ground. The disposal of slash should be 

a routine measure for protection of the stand against fire


and as a special treatment against the decay organisms 

and pests which grow and breed in the slash (Bagchee 

1954). 

The infestation by mistletoes can be controlled by 

lopping before the ripening of the fruits and their 

dispersion by birds (De 1945). 

The service life of treated wood has been estimated 

to be six times more than that of untreated wood. Greater 

utilisation of preservative treated wood would lessen the 

demand for timbers. An efficient conservation 

programme could therefore be implemented (Hong and 

Daljeet-Singh 1985). 


Pest and disease problems are going to play an important 

role in enrichment planting and establishment of forest 

plantations. As forest exploitation continues, the natural 

balance of pest and diseases in the forest ecosystem will 

be disturbed. Pathogens and pests are likely to play an 

important role in a wide variety of ecological and 

evolutionary phenomena. There is a need to formulate a 

good pests and diseases management programme, both 

at national and regional levels, with identification of 

priorities and to support the development of technology 

and capacity to face pests and diseases. Forestry pests 

and diseases on dipterocarps occur in six major 

categories: seed storage, nursery problems, 

establishment problems, chronic and sporadic problems, 

wood destruction and fruiting and seedling survival in 

natural stands. 

The main constraints on dipterocarp pest and disease 

research are shortage of trained staff, lack of cooperation 

among scientists and institutions working on pests and 

diseases in Asia, inadequate funding and infrastructure 

facilities, high cost of pest and disease identification, 

lack of information on the economic effects of pests in 

plantation forestry, and the need for more contacts 

between researchers, foresters and staff of timber 

companies. 

Future research should therefore include the following 

aspects: 

1. Seed destruction and fungal infection during storage 

Although the main insect predators and pathogenic 

fungi have been identified, emphasis is needed on 

controls, their application, effectiveness and impact 

on seed germination and seedling development. 

Chemicals as well as biological controls should be 

tested. 

2. Pest and diseases in nursery 

Except for major epidemics, attacks and infections 

can be managed by chemicals and cultural practices. 

Nevertheless, control methods need more systematic 

study. Biological control can also be considered as a 

preventive method: soil-borne fungi such as 

Trichoderma and Gliocladium species can be used 

as antagonists to soil-borne pathogens and cultured 

in the seedling beds. 

3. Pest and diseases during establishment of seedlings 

and saplings in plantations and exploited forests 

Since enrichment planting and forest plantation 

involve investment, failure of establishment can be 

economically devastating. Special attention has to be 

given to pests and diseases of dipterocarp seedlings 

and saplings. Shoot destruction can become a serious 

problem for forest management as it induces the 

formation of lateral and multiple leaders. Chemical 

control is not practicable in large forest areas and 

other methods need investigation. Prevention can also 

be assisted by dipterocarp species mixture and 

diversity. 

4. Defoliation and heart-rot problems 

Damage assessment systems for defoliation and 

heart-rot and their economic impacts are required, 

as well as the study of biology and ecology of the 

pests and pathogens. Pathogens have a major 

influence over forest reforestation methods and 

breeding programmes (Augspurger 1990). Chronic 

and sporadic pest and disease problems need to be 

more systematically studied and their economic 

losses fully quantified. 

5. Fruit and seedling pest and diseases 

More studies on pests and diseases related to fruiting 

and seedling survival should be conducted to better 

understand fruiting and dispersal strategies, seedling 

survival, management and selection of the mother 

trees, and ability to resist pathogens and pests. 

6. Insect and fungal population 

Studies on insect and fungal population ecology and 

dynamics are also essential for the conception of a 

good pest and disease management programme as 

well as a search for resistant individuals (mothertrees). 

7. Revision of the insect taxonomy 

The long lists of identified insect pests in literature


refer to the old taxonomic classification. Still many 

insects have not been identified beyond genus. A 

thorough revision of the insect taxonomy needs to 

be conducted. A taxonomy training programme for 

Asian research staff will help to reduce costs and 

update laboratories’ data-bases and collections. 


I offer my thanks to the European Union for funding a 

project on dipterocarps, and the Forest Research Institute 

Malaysia for providing the facilities to carry out this 

project. Additional thanks to Mrs. Kong and her 

colleagues for their help in my bibliographic work. I also 

thank Dr. S. Appanah and Dr. J. Intachat (Forest Research 

Institute Malaysia), Dr. R. Bonnefille and Dr. Vasanthy 

Georges (French Institute of Pondicherry), Dr. G. Maury- 

Lechon (Centre National de la Recherche Scientifique/ 

Université Lyon 1), Dr. K.S.S. Nair (Kerala Forest 

Research Institute) and Dr. L. Curran (University of 

Michigan) for their suggestions on improving the final 

draft. 

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Management of 

Natural Forests 

S. Appanah 

The view that it is not possible to manage natural forests 

in the tropics for their timber has its adherents. 

Considering the widespread failures in many countries, 

such a view is conceivable. A review by International 

Tropical Timber Organization estimated that only an 

insignificant amount of the world’s tropical moist forests 

is sustainably managed (Poore 1989). Fortunately, 

numerous reports suggest otherwise for some 

dipterocarp forests of Asia. The state of tropical forest 

management worldwide is in such a quandary that any 

success, however meagre, requires close examination. 

Such a success may provide the flicker of hope that is so 

urgently needed in our efforts to save these tropical 

forests. 

The dipterocarp forests in the perhumid zone of Asia 

form the cradle for a considerable proportion of life 

forms found on Earth. It is arguable that the only effective 

way to preserve a sizable portion of this biodiversity will 

be through effective management, including production 

of timber and other valuable products. 

Fortunately, history is on the side of dipterocarp 

forests. The origins of scientific tropical forest 

management began in Asia, particularly in British India 

around the mid-19th Century. Together with teak, the 

dipterocarp forests were among the first tropical forests 

to be managed. The Indian experience formed the basis 

for management in the Malayan realm (Hill 1900). The 

conditions for management have changed considerably 

since then, but the experience and understanding gained 

form an excellent basis for developing appropriate 

management regimes for tropical forests. 

Forest Composition, Distribution and 

Structure 

Although the family Dipterocarpaceae is presently 

recognised pantropical, with three subfamilies 

Chapter 8 

Monotoideae (Africa, Colombia), Pakaraimoideae 

(Guyana) and Dipterocarpoideae (Asia), it is the last 

subfamily that is of significance as a timber group 

(Ashton 1982). The present review will be confined to 

the Asian subfamily. It comprises 13 genera and some 

470 species, distributed from the Seychelles in the west 

to Papua New Guinea to the east. In Chapter 1 more 

details on the taxonomy, distribution and diversity of this 

subfamily are given (see also Champion 1936, Symington 

1943, Ashton 1980, 1988). 

Dipterocarps are limited to tropical climates with a 

mean annual rainfall exceeding 1000 mm, with only short 

dry spells. The Asian dipterocarp forests can be divided 

into two basic zones, viz. the Moist Tropical Forests and 

the Dry Tropical Forests (Champion and Seth, 1968, 

Collins et al. 1991). Within these two basic moist and 

dry tropical forests, four forest types can be distinguished 

(Table 1). 

Our knowledge of these forests, especially the 

distribution of dipterocarps within them is incomplete. 

This is particularly the case with forests of Indochina 

and southern China. Under these circumstances, and for 

the sake of brevity, the presentation is simplified to three 

groups of dipterocarp forests, viz. the Dry evergreen 

dipterocarp forests (Dry tropical forests), Seasonal 

evergreen dipterocarp forests (Tropical semi-evergreen 

and Tropical moist deciduous forests), and the Aseasonal 

evergreen dipterocarp forests (Tropical wet evergreen 

forests). Some information on their distribution and 

structure is given below. 

Dry Evergreen Dipterocarp Forests 

These forests are found in Central and East India, Burma, 

Thailand and Indo-China. The forests are dry, with less 

than 2000 mm of annual rainfall and a dry season of 3 to 

5 months. The forests are medium in stature, with an even 

canopy and no emergents. Shorea robusta (Indian sal)

Management of Natural Forests 

Table 1. Classification of Asian dipterocarp forests (after 

Champion and Seth 1968, Collins et al. 1991). 

I. Moist Tropical Forests: 

1. Tropical wet evergreen- 

1a. Evergreen dipterocarp- 

Malaysia, Sumatra, Kalimantan, Irian Jaya, 

Maluku (part), Papua New Guinea, Sri Lanka 

(part), Peninsular Thailand, Tenasserim, 

Andamans and Nicobar (part), Philippines 

(part), Laos, Cambodia, Vietnam (part) 

1b. Secondary dipterocarp (seral)- 

Malabar coast 

2. Tropical semi-evergreen- 

North Thailand (part), Chittagong, Laos, 

Cambodia, Vietnam (?) (part) 

3. Tropical moist deciduous- 

Maluku (part), Palawan (part), Zambales 

mountains in Luzon, W. Mindanao 

Moist sal- Terai, E. slopes of W. Ghats, Chota 

Nagpur, Upper Burma, Assam (part) 

II. Dry Tropical Forests: 

4. Tropical dry deciduous (forests heavily degraded)- 

Dry sal- Western India, Burma (part) 

Indaing- Irrawaddy plains (part) 

is the well known species of this zone. Sal occurs in the 

Himalayan foothills from northwestern Himachal 

Pradesh to central Assam and south to Tripura. It also 

spreads south along the eastern part of India up to Andhra 

Pradesh. Where sal occurs, it is the only dipterocarp in 

the forest. These forests also have five other 

dipterocarps, the majority of which are confined to the 

Indo-Burma community. The Indo-Burmese species 

include S. obtusa, S. siamensis, Dipterocarpus 

obtusifolius, D. tuberculatus and D. intricatus. Many 

of them occur as single species or codominant stands. 

These dipterocarps have thick bark and are fire tolerant. 

Today most of these forests have become more open as 

a result of browsing of young regeneration by cattle and 

felling. 

Seasonal Evergreen Dipterocarp Forests 

These forests are distributed north and east of the everwet 

Malesian region of Malaya, Borneo and Sumatra. They 

are found in places that experience a short but regular 

dry season. The forests occur in western and southern 

parts of Sri Lanka, western Ghats of India, the Andaman 

Islands, eastwards from Chittagong (Bangladesh) to 

134 

southernmost Yunnan and Hainan (China), and southwards 

to Perlis, northwest of Peninsular Malaysia. In the 

eastern parts of Malesia they occur again, in parts of 

Sulawesi, the Moluccas, Bali, Lombok and New Guinea. 

Only about a 100 species of dipterocarps are found in 

this formation. They occur in the mature phase of the 

forest, with no single species dominating the canopy. 

About half of the canopy layer may consist of 

dipterocarps. They tend to be found in gregarious stands, 

and some like Anisoptera thurifera act like pioneers, 

colonising sites that were cultivated. Details of the 

species found in these forests are found in Ashton 

(1982), Champion and Seth (1968), Chengappa (1934), 

Rojo (1979), Smitinand et al. (1980), Vidal (1979), 

Johns (1976) and others. 

Aseasonal Evergreen Dipterocarp Forests 

These are the forests that occur in the perhumid climate 

of Malesia, with rainfall over 2000 mm annually, and no 

pronounced seasonal water stress. These forests are 

found all the way from southwestern Sri Lanka, 

Peninsular Malaysia, Sumatra, Borneo, and the 

Philippines. Similar but somewhat poorer forests can be 

found in Irian Jaya in the east. The vast majority of the 

dipterocarps, over 400 species, occur in this formation, 

with Borneo having the biggest share. A complete list of 

the Malesian species is given in Ashton (1982). The trees 

dominate the emergent layer of lowland and hill forests, 

but this is not the case in Irian Jaya where the dipterocarps 

mainly make up the canopy species. Besides the lowland 

and hill species, there are dipterocarp-dominated 

montane formations, as well as several species adapted 

to heaths, coastal hills, limestone cliffs, peat swamps 

and freshwater swamp forests. Dipterocarps may 

constitute between 50-60% of the emergent stratum in 

the rich lowland formations, but under optimum 

conditions, the trees may make up 80% of the emergent 

individuals and occur as gregarious or semi-gregarious 

populations. 

Natural Regeneration 

Dry Evergreen Forests 

The best known dipterocarp forests of the dry zone are 

the sal forests of India. Sal fruits annually, with heavy 

fruiting at intervals of 3 to 5 years (Champion and Seth 

1968). The flowering begins during the dry period, and 

the fruits mature with the rains. A mature sal can produce


about 4000 viable seeds in a good year (Champion and 

Pant 1931), and the seeds germinate within a few days. 

Sal seedlings are shade tolerant and establish better under 

the crowns of other trees. Seedlings are able to coppice 

and also develop a deep taproot. They are thereby able 

to withstand ground fire and cattle browsing. 

Other dipterocarps of this formation are believed to 

regenerate like sal. All flower during the dry season, and 

fruit with the onset of rains. A light ground fire before 

seed-fall assists seedling establishment. Among some 

species, seedling establishment seems rare in nature 

(Blanford 1915), and regeneration is principally by 

coppicing. Mature trees are known to coppice readily 

following injury. 

Seasonal Evergreen Dipterocarp Forests 

The dipterocarps of this formation belong to the mature 

phase of the forest. An exception is Anisoptera thurifera 

in Papua New Guinea which can establish in cultivated 

areas (Johns 1987). The regeneration of the dipterocarps 

in these forests resembles that of the sal in many ways, 

except for the role of fire. Dipterocarp populations 

flower almost annually, but flowering is only heavy at 

intervals of 3-4 years (Chengappa 1934). The fruits are 

heavily predated by insects, birds and mammals, and 

seedling survival is poor. In some genera like 

Dipterocarpus, many years may pass without a single 

seedling becoming established. They also lose their 

coppicing ability after the sapling stage. Overall, the low 

seedling survival and the early loss of coppicing ability 

makes it difficult to regenerate these forests after 

exploitation. 

Aseasonal Evergreen Dipterocarp Forests 

The regeneration of dipterocarps in these forests has 

been relatively well studied. The dipterocarps have a 

unique flowering characteristic - they flower at supraannual 

intervals of 2 to 7 years, and the event may be 

widespread covering sometimes the whole region 

(Ridley 1901, Foxworthy 1932, Ashton 1969, reviewed 

by Appanah 1985). Whole forests may burst into 

flowering synchronously. It is not limited only to the 

dipterocarps though, and many other canopy and 

emergent species also participate in the flowering. Some 

localised flowerings also occur almost every year. 

During heavy flowering years, each mature 

dipterocarp may set up to 4 million flowers, and this 

135 

results in as many as 100,000 mature fruits. Much is 

lost to insects, birds and mammals. The ripe fruit fall 

somewhat synchronously, however, the winged fruits are 

not dispersed far from the mother trees. The dipterocarp 

seeds lack dormancy, and germinate soon after falling. 

Once established, seedling populations decline slowly 

only as a result of inadequate light conditions and 

aperiodic droughts. Growth is rapid if they are exposed 

to direct light (Wyatt-Smith 1963, Fox 1973). Among 

the dipterocarps, light demanders and shade tolerant 

species can be differentiated. Both grow rapidly where 

there is higher light intensity, but the latter species can 

survive longer under poorer light conditions, and in 

general they are the slower-growing heavy hardwoods. 

In contrast to dipterocarps in the other two formations, 

coppicing ability of the species in the everwet forests 

is limited, and ceases beyond the pole stage. The 

population structure is not the typical reverse-J shape, 

with the density of sapling-and pole-size dipterocarps 

generally low in mixed dipterocarp forests. However, 

this appears to be not so in some of the dipterocarprich 

forests in the Philippines. 

Silvicultural Systems 

A number of silvicultural systems have been developed 

for the long-term management of tropical forests, many 

with dipterocarps as the main crop. The silvicultural 

systems go by a bewildering number of technical names, 

but they can be broadly divided into Shelterwood 

(monocyclic) Systems and Selection (polycyclic) 

Systems. The situation for dipterocarps forests have 

been reviewed variably (e.g. Wyatt-Smith 1963, 1987, 

FAO 1989, Stebbing 1926, Chengappa 1944, Nair 1991, 

Weidelt and Banaag 1982, and others). 

Simply stated, the Shelterwood System attempts to 

produce a uniform crop of trees from young 

regeneration through both heavy harvesting and broad 

silvicultural treatments. A new even-aged crop is 

established by applying preparatory and establishment 

cuttings to natural regeneration (i.e. seedlings and 

saplings) of the desired trees. At an appropriate time 

the remaining overstorey is removed. 

The Selection System aims to keep an all-aged stand 

through timber cuttings at shorter intervals. Many light 

cuttings are made. Seedlings will become established 

in the small gaps. Under this system, two or more less


intensive harvests are possible during one rotation, while 

in the Shelterwood System all marketable stems are 

removed at one cutting. 

A variety of silvicultural systems have been tried out 

on dipterocarp forests, depending on markets, 

technological changes, landuse patterns, harvesting, 

regeneration, labour costs, etc. These have met with 

varying success. The systems in operation in India, 

Malaysia, Philippines and Indonesia described as forest 

management practices are well documented in these 

countries. 

India 

The seasonal evergreen and dry evergreen forests have 

been managed under the Selection System. Here it can 

be summarised as selective felling of exploitable trees 

from an area at periodic intervals, under the following 

circumstances: i) in mixed forests where utilisable 

species are few; ii) in areas that are difficult to access; 

and iii) in hilly terrain where heavy logging is 

environmentally bad. 

Trees of specific girth are removed at 15 to 45 year 

cutting cycles, calculated from growth rates. Some 

safeguards are introduced such as: a 20 m minimum 

distance between trees earmarked for felling; climber 

cutting to reduce logging damage; protection buffers for 

riversides; and only harvesting dying and dead trees in 

steep areas. Treatment is carried out to assist natural 

regeneration, and planting is prescribed for understocked 

areas. Many of the prescriptions are not met for 

several reasons: plantings are inadequate and damage to 

residuals excessive (FAO 1984). Over time, felling 

cycles have been reduced, girth limits lowered, and more 

species exploited. 

Shelterwood Systems 

Shelterwood Systems were introduced when it became 

necessary to harvest more intensively some valuable 

forests, and regeneration was not assured under the 

selection system. The variants usually applied here are 

the Indian Irregular Shelterwood System, Uniform 

System and the Coppice System. 

1. Indian Irregular Shelterwood System 

Both seasonal evergreen and sal forests are managed 

under this system. First, all trees above exploitable 

diameter are removed. If advanced growth is lacking, 

mother trees are kept. Next, the underwood and 

136 

overwood are removed periodically until regeneration 

becomes established. Finally, the remaining underwood 

and overwood is removed, except those forming future 

crops. All these are done over a rotation of 120 years. In 

addition, girdling, thinning, weeding, climber cutting and 

artificial planting are carried out as needed. 

Lack of regeneration, especially for sal forests, 

appears to undermine the Irregular Shelterwood System 

(FAO 1989). Plantings have been tried at cost. This has 

not kept to schedule, and there is a temptation to reduce 

rotation length and exploitable girth limits. 

2. Uniform System 

In high value sal forests, the Uniform System has been 

tried. All overwood is removed at one clearfelling, and 

regeneration is allowed to grow up. No regeneration 

fellings are conducted, however, and so the system has 

to rely on pre-existing seedlings. The rotations are 

between 120 to 180 years for sal. But demand for timber 

is high and rotations have been shortened. 

When natural regeneration is abundant, the overwood 

is cut completely. Groups of poles are sometimes kept 

as future crop trees if regeneration is poor. Where 

regeneration has not established, suppressed trees are 

retained to control weed growth. Steep slopes and eroded 

areas are not heavily felled. Cutting and thinning are 

prescribed for improving regeneration. The system 

should work if adequate natural regeneration can be 

secured. In the event it is poor, artificial regeneration 

has been resorted to. 

3. Coppice Systems 

A few variants of the Coppice Systems have been 

introduced for sal forests. The systems depend on shoots 

emerging from the cut stumps. Coppicing vigour declines 

with age and so short rotations are necessary. It is mainly 

suitable for firewood and small timber production. To 

produce fuelwood, a rotation of 30-40 years is used. 

Felling is done before the growing season, the area is 

protected from grazing and fire, and cleaning is done to 

remove excess coppice shoots and climbers. Over time, 

with decline in coppicing vigour, stump mortality 

increases. Seedling regeneration helps to compensate 

this loss, but seedlings are scarce because of grazing 

pressure. This has led to stand degradation. Variations to 

the system involve retention of seed trees for producing 

seedlings (see Tiwari 1968). Overall, the system has 

succeeded where biotic pressure is kept low.


4. Clearfelling System 

This system is used when there is a need to change the 

composition of the crop to a more valuable species. The 

restocking is through natural or artificial regeneration, 

the latter used to introduce a new species or to change 

the forest composition. As a consequence, the more 

valuable teak is introduced into sal forests. The trend is 

to convert most of these forests into plantations, making 

the future of sal forests uncertain. 

Peninsular Malaysia 

Forest Management Systems 

Forestry in the modern sense was started in 1883 with 

the establishment of the forestry service. Prior to 

introduction of forest management, logging was very 

selective, principally limited to the heavy hardwoods 

(mainly several dipterocarp secies), and only about 7m 3 / 

ha was taken out (Barnard 1954). Silvicultural operations 

were limited to enrichment plantings of the heavy 

hardwood, chengal (Neobalanocarpus heimii), which 

failed from lack of further tendings. But the demand for 

timber increased, leading to over-exploitation of the 

select timbers. This prompted the authorities to develop 

a series of silvicultural systems. 

1. Regeneration Fellings 

In the beginning (1910-1922) Departmental 

Improvement Fellings were implemented. All species 

whose crowns interfered with the poles of any valuable 

timber species were removed. It was subsequently 

realised that such treatments had no impact on the 

immature trees. However, they resulted in profuse young 

regeneration (Hodgson 1937). The improvement fellings 

had in fact been regeneration fellings. After 1932, 

Regeneration Improvement Fellings (RIF) came in to 

vogue. Inferior species were gradually removed over a 

series of fellings. If the regeneration was verified as 

successful, final felling of the valuable species was 

carried out. This in fact resembled the classical 

Shelterwood Systems. 

2. Malayan Uniform System 

As a rule, no forests were harvested without first carrying 

out RIF. During the Japanese Occupation (1942-1945) 

many forests were clearfelled without the benefit of RIF. 

After the war, extensive surveys revealed that these areas 

contained adequate advanced regeneration without any 

137 

assistance. It was realised that if the forest had adequate 

regeneration of the fast growing dipterocarp species, a 

single clearfelling release could result in a greater 

stocking of a more uniform crop of commercial species. 

This became the basis for the Malayan Uniform System 

(MUS), which was introduced in 1948 for managing 

Lowland Dipterocarp Forests (Wyatt-Smith 1963). 

A detailed silvicultural system was developed (Wyatt- 

Smith 1963). It consists of felling the mature crop of all 

trees above 45 cm dbh, poison girdling all defective 

relics and non-commercial species down to 5 cm dbh, 

and releasing established seedlings. Seedling adequacy 

and suitable tendings underpinned the success of MUS. 

3. Modified Malayan Uniform System 

In the mid-1970s, most of the lowland dipterocarp 

forests were alienated for agricultural programmes, and 

forestry was confined to the hills and rough terrain 

unsuitable for agriculture. Under these new conditions 

it was considered difficult to apply the MUS. The 

principal problem was the lack of uniform stocking of 

natural regeneration. It was thought that enrichment 

planting could overcome this deficiency (Ismail 1966). 

This allowed all forests to be opened up for logging, 

regardless of adequate seedling stocking, a prerequisite 

with MUS. Planting up understocked areas was carried 

out in the beginning, but their performance was very 

variable and unsatisfactory. Now, artificial regeneration 

is rarely carried out, or the practice is abandoned entirely. 

4. Selective Management System 

In the late 1970s, the Selective Management System 

(SMS) was introduced. This is a simplified version of 

the Philippine Selective Logging System (see Appanah 

and Weinland 1990). The MUS was already discarded 

for working in the hillier terrain, and the modified-MUS 

proved unsatisfactory. The felling regime is formulated 

on the basis of a pre-felling inventory. All commercial 

tree species above a certain size (ideally nondipterocarps, 

45 cm dbh; dipterocarps, 50 cm dbh) are 

felled, provided a sufficient number of residuals are left 

behind to form the next cut in ca 30 years (Thang 1987). 

Therefore the SMS relies on adequacy of healthy 

residuals which will respond to the cutting release for 

the next cut some 25-30 years later. Seedling stocking 

is assumed to be present, or will be replenished by the 

maturing residuals. The SMS is regarded as more flexible 

for managing the highly variable forest in the hillier


terrain. In situations where it is not economically 

equitable for the logger, the modified-MUS is prescribed 

which imposes an arbitrary diameter of 45 cm dbh for 

felling on a rotation of 50 years. 

Sabah 

Silviculture in Sabah followed a path similar to 

Peninsular Malaysia. In the early 1930s, RIF were tried 

on a limited scale (Fox 1968). In 1949 the Selection 

Improvement Fellings were introduced, to assist the 

pole-size trees of 10 cm dbh and above in areas logged 

15 to 25 years before (Martyn and Udarbe 1976). The 

method involved poison-girdling non-commercial 

species and climber cuttings. 

In 1956 a modified version of the MUS was 

introduced for forest regeneration (Chai 1981). The 

canopy was opened after felling by poison-girdling all 

non-commercial species as well as defective trees of 

commercial species down to 15 cm dbh. The next crop 

is expected to come from seedlings, and advance growth 

will be a bonus. This system became the standard 

regeneration technique for dipterocarp forests in Sabah. 

This modified MUS underwent further changes in 

1971 to become a minimum girth limit system, the so 

called Stratified Uniform System (Chai and Udarbe 

1977). In this refinement, the advance growth for the 

next crop is kept. The main elements of the system 

include marking 25 preferred or desired trees/ha (25- 

59 cm dbh) for retention, and poison girdling unwanted 

and defective trees. Climber cutting and girdling of seedbearers 

and relics is done in the 15th year. 

Later, Chai and Udarbe (1977) expressed doubts on 

the value of the girdling practices. They argued that since 

logging intensity is high, much of the forest gets released 

anyway without further treatment. Since then, only 

climber cuttings are meant to be done. Furthermore, 

girdling of weeds or non-commercials has been stopped 

on account that such plants may become commercial in 

the future, and moreover, the operation may be harmful 

to the ecosystem. 

Sarawak 

The timber industry in Sarawak relied mainly on 

extensive peat swamp forests, and moved into the hill 

forests only in the late 1960s. Coming so late, Sarawak 

tended to follow the systems developed in Peninsular 

Malaysia (Lee 1982). At first the forests were selectively 

logged. The relics left behind were defective and inferior, 

138 

and seedlings/saplings unlikely to reach maturity before 

70-80 years. 

As a result, three UNDP/FAO projects (1974-1981) 

were started to provide interim guidelines for managing 

Sarawak’s dipterocarp forests (FAO 1981a, b). The study 

evaluated three different treatments: 

1. Overstorey removal only - All overmature non-marketable 

trees left behind during harvesting were removed 

by poison-girdling. 

2. Malayan Uniform System evaluated - Following logging, 

all other non-economical trees, which impeded 

growth of the seedlings were removed. Such a treatment 

was considered too drastic. The rough terrain 

and shallow soil conditions are vulnerable to heavy 

erosion. A modification to MUS was tried whereby 

the advance growth of the desirable species were 

saved. In this way the advance growth may be obtained 

even before the seedlings mature, giving in effect a 

polycyclic system. 

3. Liberation Thinning - Desirable species were identified, 

and liberated from competition including removal 

of the overstorey to improve their growth. No 

specific species or species groups were eliminated, 

only those that restricted the growth of the selected 

trees. Therefore, trees of non-commercial species 

were left behind if they did not appear to hinder selected 

trees. 

Mild overstorey release was insufficient to release 

the trees of desirable species. Both the Liberation 

Thinning and the modified MUS resulted in increased 

growth of the residuals (Hutchinson 1979), but the latter 

resulted in elimination of a greater number of trees which 

could have commercial value in the future. Despite the 

potential loss in the future of commercial trees, for a 

while liberation thinning held sway in Sarawak as the 

appropriate silvicultural treatment (FAO 1981b). It lost 

support subsequently, when Lee (1982) suggested that 

the boost in initial growth is not sustained, the operations 

are difficult, and cannot be kept up with the logging rate. 

Since then, Liberation Thinning is being carried out for 

a small portion (ca. 4%) of the forest logged annually 

(Chai 1984). Otherwise, the practice has reverted to 

selective felling based on diameter limits. 

Philippines 

Scientific management of dipterocarp forests began 

during the American Regime. From 1900 to 1942 

mechanised timber extraction and processing methods


were introduced. Following the Second World War, there 

was a surge in logging for rebuilding the country, and the 

only management control was a ‘diameter limit’ of 50 

cm for cutting trees. Despite the limit, mechanisation 

of logging led to almost clear-cutting due to high 

stocking. 

The above ‘diameter limit’ cuttings brought about the 

development of the Philippine Selective Logging System 

(PSLS), which is a modification of the Selection System 

used to manage old growth hardwood forest in North 

America. Under this system, 60% of the healthy 

commercial residuals in the 20-70 cm dbh classes are 

to be retained as growing stock for a future harvest 

(Reyes 1968). This has since been raised to 70% of all 

the commercial residuals in the 20-60 cm dbh classes. 

The selective logging amounts to removing mature, 

overmature and defective trees with minimum injury to 

an adequate number of healthy residuals of commercial 

species to guarantee a future timber crop. Also 

incorporated into the system is a timber stand 

improvement (TSI) guideline which consists of 

treatments before and after the major felling to ensure 

the stand attains maximum timber quality and growth 

(Uebelhoer and Hernandez 1988). The TSI appears to be 

yielding results. Preliminary results indicate that 

liberation from crown competition results in increase 

in diameter: a removal of 33% basal area, resulted in up 

to 10% increase in basal area of crop trees in ten years. 

The Philippine forests are generally very rich in 

dipterocarps. Therefore, the PSLS is regarded as the best 

silvicultural system for their forests. If logging damage 

is contained, and residual forests protected and post 

logging treatment given, another economic cut is 

possible after 30-45 years. While the system looks good, 

overcutting and bad implementation has led to 

degradation of vast areas of forests. Today, there is 

concern for the quality of the second cut. 

Indonesia 

From historical times, teak forests in Java have received 

most interest from silviculturists in Indonesia. After 

1966, changes in forest policy took place and the 

dipterocarp forests in the other islands were opened for 

large scale exploitation. At first it was merely a timber 

felling operation. Sustained management efforts began 

in the 1970s when a simplified variation of the PSLS 

was introduced for lowland dipterocarp forests 

(Soedjarwo 1975). The original version, the Indonesian 

Selective Cutting System, locally known as the TPI 

139 

(Tebangan Pilih Indonesia), relies on leaving behind an 

adequate number (25 stems/ha or more) of sound 

commercial species of 20 cm dbh and above. With this 

minimum guaranteed, everything above a certain diameter 

limit may be harvested. If the putative residuals could be 

met, the TPI system allowed for a short felling cycle of 

ca 30 years. If these were not present, the option was to 

harvest on a Uniform System rotation of ca 60 years. 

There was also a further option to clear cut and replant, 

although not necessarily with dipterocarps. 

Compared to the PSLS, the TPI is a much simpler 

system. It is therefore cheaper and easier to monitor. 

Liberation thinning is prescribed to release residuals and 

nucleus trees for reseeding. Planting of seedlings to 

enrich the stand may be carried out if followed by 

subsequent tending and liberation thinning. 

Pre-felling inventories in Indonesia however suggest 

that stands rarely have sufficient residuals of commercial 

species (Burgess 1989). Therefore, the second cut may 

have to be delayed. The TPI was subsequently modified 

to the TPTI (Tebang Pilih Tanam Indonesia) which 

resorted to the necessity of planting if the selecting 

fellings failed. This resulted from the conviction that it 

is possible to easily plant up large areas with 

dipterocarps (see Enrichment Planting). Unfortunately, 

the impression from this decision is that uncontrolled 

logging can be done without serious consequences, as 

enrichment planting can overcome the problems. Caution 

should be exercised here until evidence for the success 

of enrichment planting is clear. 

Growth and Yield 

One of the biggest difficulties for sustained management 

of dipterocarp forests is in getting reliable data on growth 

and yield. The data are a prerequisite for determining 

harvesting volumes and cutting cycles. In this respect, 

there is much scepticism about the growth rates being 

used for managing many forests in the region. A quick 

glance of the data from the everwet region, based on only 

a few sites, gives some clue to how dipterocarps are 

growing. 

From studies in Peninsular Malaysia, Sabah, Sarawak, 

Philippines and Kalimantan, the following 

generalisations can be made. In undisturbed, virgin 

forests growth rates are relatively much lower compared 

to logged ones, and the best growth is achieved in 

plantation conditions (e.g. mean growth rate (diameter 

increment) of Shorea spp. in Sarawak: primary forest,


0.82 cm/yr; logged forest, 0.93 cm/yr; plantation, 1.22 

cm/yr) (Primack et al. 1989). In any case, among the 

commercial species, dipterocarps grow much more 

vigourously than non-dipterocarps, by at least 25-35% 

(e.g. periodic diameter mean annual increment for Labis 

F. R., Peninsular Malaysia: dipterocarps, 0.85 cm/yr; 

non-dipterocarp commercials, 0.66 cm/yr) (Tang and 

Wan Razali 1981). Among the dipterocarps the light 

hardwoods grow faster than the heavy ones (growth rates 

in Peninsular Malaysia sample plantation plots: light 

hardwood Shorea macrophylla, 2.23 cm/yr; heavy 

hardwood Shorea sumatrana, 0.86 cm/yr) (Appanah and 

Weinland 1993). Growth rate, expressed in diameter 

increment, is lowest with smaller individuals, and 

culminates usually in the 50-60 cm diameter classes, 

and declines in bigger trees. This pattern of diameter 

increment has been seen in the Philippines (Weidelt and 

Banaag 1982), Sabah (Nicholson 1965), and Peninsular 

Malaysia (Tang and Wan Razali 1981). A sample from 

the Mindanao concessions in the Philippines illustrates 

the point: 

Following logging or liberation thinning, the 

residuals are known to respond to the release by 

increasing their growth rates. In general the increments 

were highest in the first years after logging, and declined 

slowly, and after about the fifth year the benefits of 

release seem to cease (Tang and Wadley 1976). 

Site\ Age Year 1 Year 2 Year 3 Year 4 Year 5 

Peninsular 

Malaysia- 

Tekam F.R. 

Peninsular 

Malaysia- 

Labis F.R. 

dbh class (cm) cm/yr 

10 

20 

30 

40 

50 

60 

70 

80 

90 

0.72 

0.63 

0.57 

0.79 

0.68 

0.71 

0.44 

0.58 

0.69 

0.78 

0.83 

0.86 

0.86 

0.85 

0.79 

0.67 

0.63 

0.57 

Besides, the above, the trees grew faster (mean annual 

diameter increment) in plots where more timber was 

harvested (plot residual basal area) after logging (Tang 

and Wadley 1976): 

n/a 

Residual basal area 

(m 2 /ha) 

10-16 

16-22 

>22 

140 

A peculiar behaviour of all tropical trees, including 

that of dipterocarps, is the extremely wide range of 

growth rates of individual trees even within the same 

diameter class. The variation coefficient may reach 70- 

100%. This is illustrated in the mean annual diameter 

increment for the minimal, maximal and median growth 

rates (cm/yr) of Shorea species in primary, liberationtreated, 

and plantation forests in Sarawak (Primack et 

al. 1989): 

Mean annual 

diameter 

increment (cm) 

Primary 

Forest 

Mean annual diameter 

increment (cm/yr) 

Liberation 

Felling 

0.44 

0.45 

0.55 

Plantation 

Minimum 0.13 0.16 0.80 

Maximum 0.82 0.93 1.22 

Median 0.30 0.43 0.86 

Next is the variation in the growth rates within one 

region, and between regions. Studies of the annual 

diameter increment (cm/yr) of dipterocarps in the 

Philippine (Weidelt 1996) and Sarawak forests (Primack 

et al. 1989) illustrate these points: 

Location Mean annual diameter 

increment (cm) 

Sarawak: 

Mersing 

Bako 

Philippines: 

Mindanao 

Visayas 

Luzon 

0.41 

0.30 

0.73 

0.48 

0.52 

It should be noted that dipterocarps on fertile sites 

in the high rainfall area of eastern Mindanao have high 

annual increments. The growth rates are consistently 

better in the Philippines than Sarawak, indicating regional 

differences. The forests in the Philippines should 

generally have better yields. Even within Sarawak, there 

are differences between the two forests, which can be 

ascribed mainly to better soil fertility at Mersing. 

Despite the existence of some good information on 

the growth of dipterocarp trees, there is a tendency to 

exaggerate their growth rates. For example, in Peninsular 

Malaysia, the generally accepted standard for growth of 

trees in logged forests is above 0.8 cm/yr diameter


increment, and hence a cutting cycle of about 35 years. 

From a quick perusal, it is obvious the realised growth is 

far below that assumed. Furthermore, the wide variation 

in growth rates between forests calls for more precise 

local growth data for determining cutting cycles, and 

national averages are inapplicable. Next, despite evidence 

that silvicultural treatments of girdling and liberation 

felling do boost the growth of the trees, this is rarely 

undertaken. This of course has to be taken into 

consideration with the costs of operations and the 

benefits of increased timber production. 

Enrichment Planting 

Enrichment planting has been a tool in dipterocarp forest 

management, and several dipterocarp species have been 

successfully planted into natural forests (Barnard 1954, 

Tang and Wadley 1976, reviewed in Appanah and Weinland 

1993, 1996). It is indeed widely and variably practiced 

throughout the Asian tropics. Such planting is considered 

when the stocking of seedlings and saplings of desirable 

species is inadequate because of poor seedling survival 

or due to destructive logging methods. With the 

modified-MUS of Peninsular Malaysia, enrichment 

planting was supposed to be a standard practice: the 

deficit in natural regeneration to be artificially 

regenerated using dipterocarp wildings. 

The success of such plantings was variable and 

planting efforts have invariably declined. There are 

several causes for this. Planting work is difficult to 

supervise, seedlings have to be regularly released from 

regrowth, and a regular supply of dipterocarp seedlings 

is needed. Wildings can be used, but individuals differ 

widely in their performance. Moreover it is costly (labour 

demanding). As a consequence, the efficacy of 

enrichment planting has been questioned (Wyatt-Smith 

1963, OTA 1984). 

Nonetheless, enrichment planting is receiving 

accelerated attention as a possible technique under the 

selective felling practices in Kalimantan (e.g. Smits 

1993, Adjers et al. 1996). Extensive areas are being 

planted up in Kalimantan with dipterocarp wildings. 

Rooted cuttings have also been developed but their 

success in the field has not been evaluated yet. Their root 

structure must hold the tree during sudden wind storms. 

Smits (in Panayotou and Ashton 1992) has in view a 

model for enrichment planting of degraded dipterocarp 

forests in Kalimantan. Such sites are to be first planted 

141 

with an over-storey of building-phase species, and a few 

years later with dipterocarps raised from cuttings and 

inoculated with mycorrhiza. The fast growing species can 

be harvested in the mid-term, and this will release the 

dipterocarps for harvest in 50 years. One major technical 

problem is the difficulty in harvesting the pioneer species 

without causing excessive damage to the mature-phase 

trees (Panayotou and Ashton 1992). There is also concern 

for the bad form of dipterocarps raised from cuttings. 

Wyatt-Smith (1963) pinpoints the conditions which 

merit enrichment planting, and the silvical characters 

necessary for species ideal for enrichment planting. The 

characters include regular flowering and fruiting, rapid 

height growth, good natural bole form, low crown 

diameter/girth breast height, wide ecological amplitudes, 

tolerance to moisture stress, and free of pests and 

diseases. But most of all, the species should produce 

timbers of high value. 

All too often, enrichment planting is done without 

consideration for the light conditions. Supervision and 

follow-up maintenance are necessary, especially canopy 

opening treatments. With care, enrichment planting 

remains promising and viable. It has been successful in 

Karnataka and several other Indian States, and Sri Lanka, 

in both moist deciduous and evergreen forests. 

While it is generally accepted that the best and 

cheapest method for regenerating dipterocarp forests is 

still using the natural regeneration, enrichment planting 

has received a new boost particularly for badly degraded 

forests. Under the ‘Carbon Offset’ Project, an American 

utility company paid for planting dipterocarps in Sabah, 

to offset its carbon dioxide emission in its power plants 

in Boston (Moura-Costa 1996). This may appear 

innovative, although its value will be confined to 

rehabilitation programmes. Planting dipterocarps may be 

viewed as a final resort, after natural regeneration 

practices have failed. 

Exploitation Damage 

Good harvesting systems are critical for sustainable 

management of natural forests. The harvesting should not 

irreversibly compromise the potential of the forest. The 

operations should never degrade it, and must also allow 

for rapid recovery of the stand. Studies of logging damage 

in dipterocarp forests begun in the late 1950s show that 

it has been increasing with mechanisation (Nicholson 

1958, Wyatt-Smith and Foenander 1962, Fox 1968). But


with properly planned and executed harvesting 

operations, not only is the damage contained, but so are 

the harvesting costs (e.g. Marn and Jonkers 1981). 

Unlike the case with uniform (Shelterwood) systems, 

selective fellings can cause considerable damage to the 

future crop, the medium sized residuals. The damage 

intensity and extent to both trees and soils vary with the 

log extraction system used. Skidder-tractors are used 

extensively. They cause more damage to the ground 

surface, increasing soil erosion and retarding 

regeneration and growth of residuals. With precautions 

and improvements like pre-determined skid trails and 

reduced vehicle movement, damage can be considerably 

reduced. Logging on steep slopes (i.e. >15 o ), which is 

very damaging, should be curtailed. 

Besides damage caused by extraction, felling damage 

too can be very intense, especially to the advanced 

regeneration (Nicholson 1979). Directional felling and 

pre-felling climber cutting reduce such damage. 

Although this practice has been recognised as beneficial, 

it is seldom carried out. Currently, several initiatives have 

been started in reducing logging damage to the soils and 

the residual vegetation under schemes called ‘Reduced 

Impact Logging ’ (RIL). These initiatives are mainly in 

Sabah (Marsh et al. 1996). In these RIL operations, 

besides cutting lianas, directional felling and pre-planned 

skid-trails, the operations are closely supervised so as 

to minimise skid trail length and blade use. A 50% 

reduction in all measures of damage was demonstrated 

compared with conventional logging for an increase of 

about 10-15% of direct logging costs. 

High-lead yarding systems have been tried in some 

concessions in the Philippines and Malaysia. They are 

costly, difficult to maintain, and require well trained 

crews to maintain them. Basically, selection fellings and 

high-lead yarding are incompatible, as the residuals are 

damaged considerably. There is also heavy damage to the 

soil when trees are dragged uphill. However, skyline 

yarding systems are beginning to show considerable 

promise. With the simple skyline yarding where two spar 

trees are used, road building is reduced. The other is the 

Long Range Cable Crane System which uses a tight 

skyline with intermediate supports and a carriage with 

the log suspended to it vertically. The carriage travels 

along the skyline and dumps the suspended log at the 

head of the spar or tower. This has been tried in the 

Philippines (Heyde et al. 1987) and Sabah (Ong et al. 

1996). The original carriage could only lift small logs, 

142 

but the new one introduced in Sabah can lift 5 tonne logs 

(personal observation). The use of a skyline system 

reduces road building considerably, and limits damage 

to the soil and residual trees to a considerable extent. 

The skyline systems hold the answer to logging of 

dipterocarp forests of Southeast Asia. 

Helicopter logging is now being tested in Sarawak. 

This system remains rather expensive and dangerous. The 

cost of keeping the helicopter in the air is high, and the 

operations have to be perfectly coordinated: trees have 

to be felled in advance, and the helicopter can only start 

its operations when a sufficient number of trees are 

available. The timber being harvested should have very 

high value. Too many accidents have happened with 

helicopter logging for it to be considered a viable 

operation. There is also the problem of illegal logging 

as it becomes much easier to steal timber using 

helicopters, and the activities are difficult to control. 

Failures in Implementation of Practices 

It is obvious from the above review of silvicultural 

practices, there is no lack of scientific methods for 

managing the variety of dipterocarp forests. While 

systematic management may be lacking (Leslie 1987, 

Wyatt-Smith 1987), some kind of management is being 

attempted for many of the forests in Asia; it is however, 

mainly in the form of area or volume control. It was 

reported that about 19% of the Asian region’s productive, 

closed broadleaf forest is being intensively managed 

(FAO 1981c). However, one can dispute if area and 

volume control is management. 

Several factors seem to hinder true management of 

these dipterocarp forests. For one, it seems better to 

cash in the timber market now than wait for uncertain 

future markets. Next, there is a mismatch between 

declared policy and implementation. Far too few 

resources are allocated for management, while the rate 

of logging is beyond what the forestry agencies can cope 

with (Wyatt-Smith 1987). Some managers have adopted 

the ‘minimum intervention’ approach on the argument 

that there are still uncertainties in the value of some 

silvicultural treatments (Tang 1987). 

Forestry agencies are unable or unwilling to 

implement the declared management policies, and 

silvicultural prescriptions are always behind schedule, 

or abandoned altogether. Panayotou and Ashton (1992) 

present several cogent reasons for this:


1. Heavy pressure from politicians to practice 

accelerated felling cycles, clear felling, re-entry, and 

leniency with regard to logging damage and illegal 

cuttings; 

2. Uncertainty of forest tenure, due to rapid conversion 

of forest lands to agriculture, uneven distribution of 

land, and short logging tenures which discourage 

private investment; and 

3. Grossly undervalued resources, with timber prices 

not including replacement or silvicultural costs and 

non-timber values. The stumpage and royalty fees are 

kept too low, and the governments do not receive the 

logging profits needed for silvicultural treatment. 

An Evaluation 

Silvicultural systems for natural forests have to ensure 

natural regeneration succeeds, and the quality, quantity 

and size of the chosen tree species are enhanced, without 

destroying the forest structure and function. Enrichment 

planting is an expensive alternative that should be 

minimised. Both the Shelterwood (monocyclic) and 

Selection (polycyclic) Systems are being purportedly 

used for managing dipterocarp forests in Asia. But how 

do the two systems stand up in real practice for managing 

dipterocarp forests? Shelterwood Systems depend 

directly on treating the desired seedlings for the next 

crop. This is a conceptually simple system which requires 

less supervision, and if done carefully, there is little 

damage to the next stand (Putz and Ashton, unpublished). 

Several workable examples of Shelterwood Systems have 

existed, the Malayan Uniform System being a well known 

one among them. 

The critical factor seems to be the ease with which 

regeneration can be secured. It is this particular feature 

of dipterocarps that makes it much easier to manage them 

compared to other forest types. In the case of sal forests, 

natural forest management seems sustainable only where 

regeneration is easy to secure. This is the case with 

Coppice Systems, provided grazing and fire are 

controlled. The MUS has also capitalised on the profuse 

seedling regeneration capacity of the family. 

Nevertheless there are elements within Shelterwood 

Systems that are discouraging: 

1. Logging has to be delayed until the regeneration is 

ensured; 

2. Rotations are long, by human terms; 

3. Heavy felling might induce weed growth, and expose 

fragile soils to erosion; and 

143 

4. Unwanted trees which were formerly girdled can now 

be exploited with improved technology and 

diversified markets. Although such canopy openings 

would have allowed the highly preferred target trees 

to maximise their growth. 

The Shelterwood Systems developed for all three 

dipterocarp forest types showed signs of success. But 

in many instances the Shelterwood Systems seem to have 

fallen victims of outside changes. Workable systems have 

thus been continuously incapacitated by the demands of 

society, rapid and unplanned landuse changes, illegal 

felling, fire and grazing, and finally our complete 

bewilderment with tropical ecosystems. The four 

examples below highlight them: 

1. The Coppice Systems in India have been clearly 

worked out, and may be the only dipterocarp forests 

sustainably managed for 3 rotations or more. But the 

demand for timber and fuelwood in India exceeds the 

production. The silvicultural response has been to 

shorten rotations. This has not been a realistic 

solution because increased frequency of removal 

results in degradation of stumps. Leaving behind 

standards to assist natural regeneration to 

compensate for the degradation was tried. This too 

proved unsuccessful because these forests are close 

to villages and the demand for grazing lands is high. 

When the demand for firewood and small timber 

exceeded biological capacity, shorter rotations were 

resorted to to enhance supply. This has accelerated 

the decline, and the areas have to be planted up as a 

consequence. 

2. In the Malayan case, the MUS which took form 

following the Japanese Occupation (1942-1945) 

could never really be put into practice. During the 

1950s Emergency in Peninsular Malaysia guerrillas 

took refuge in these very forests. It was difficult to 

work long in a forest - it was often a case of log and 

leave. The 1970s saw peace and an acceleration of 

economic growth. Large tracts of the lowland 

dipterocarp forests, for which the MUS was 

formulated, were converted to plantations of cash 

crops. Thereafter, logging was confined to the hillier 

terrain. Here the MUS was considered unsuitable and 

selective fellings have been applied. 

3. In some instances sheer confusion seems to have 

prevailed in our attempts to manage dipterocarp 

forests. In Malaya, Departmental Improvement 

Fellings of the 1930s proved ineffective on the poles


and immature trees, for they need to be repeated 

(Wyatt-Smith 1963). Following the initial burst, 

growth slows down with onset of crown competition. 

In the 1970s, such thinnings were introduced in 

Sarawak under a different name, ‘Liberation 

Thinnings’ (Hutchinson 1979). But the Department 

reduced such treatments on the basis that the 

increments are too small for the effort (Lee 1982). 

However, liberation thinnings to forests following a 

diameter limit cutting proved better (Chai 1984, 

Primack 1987). This resembles more a MUS except 

for the logging which was under diameter limits. With 

this kind of confusion, opportunities for better 

management were bypassed. 

4. In other cases, the Shelterwood Systems have 

degenerated into selective fellings. In the Indian 

Irregular Shelterwood System, uncertainty of 

regeneration led to retention of trees below a 

specified girth as part of the future crop. This has led 

to some confusion, and silvicultural treatments 

benefit neither seedlings nor poles. 

5. Most extreme is the case with Peninsular Malaysia. 

The system introduced here to manage the hill forests 

was called the Selective Management System (Mok 

1977). One of three systems was to be applied 

depending on the requirements. This included the 

monocyclic MUS, polycyclic Selection System, and 

cutting and planting. But unfortunately, the Selective 

Management System in practice became a selective 

felling. 

In contrast with Shelterwood Systems, the Selection 

System is based on maintaining the forest stand structure, 

by extracting proportionate number of trees from 

different size classes. It works well with species that can 

tolerate some shade, and small gaps suffice for their 

growth (Putz and Ashton unpublished). The system allows 

frequent timber extractions, but substantial management 

is required. Logging has to be carefully done to protect 

young trees. 

The selection systems are not truly practised in the 

dipterocarp forests although the Philippines Selection 

Felling System in theory has the necessary silvicultural 

components to qualify as one. Elsewhere, Selection 

Systems have degenerated in practice into selective 

fellings based on diameter limit. This is not a silvicultural 

system in the classical sense. Critics claim selective 

fellings cannot fulfill the requirements of a polycylic 

system (Wyatt-Smith 1987, Appanah and Weinland 

144 

1990), and that in reality it is merely a bicyclic system. 

Its major difficulties are: 

1. Seedling regeneration is not attended to, and this 

might lead to a decline in the future crops; 

2. Composition of future crops cannot be controlled; 

3. The intermediate class (residuals) which is poorly 

represented, may also be inferior, suffer much 

logging damage, and subsequently succumb. Overall 

their growth rates may also be below that forecasted; 

4. The cutting cycles are over-optimistically short; and 

5. The more frequent entries can damage the soil and 

young regeneration. 

Despite the criticisms, most of the seasonal and 

aseasonal dipterocarp forests are selectively logged at 

present. Perhaps the advantages of short felling cycles, 

fewer tendings, and freedom from limitations of 

seedling regeneration have led to such a preference. 

Supporters nonetheless argue that the Selection System 

is suitable for dipterocarp forests, many of which are 

now in steep terrain, with spotty seedling regeneration, 

and are relatively inaccessible. The weakness is in the 

implementation. The test of course is with the second 

cut, which will soon take place in Malaysia and Indonesia: 

overall, a decline in yield is expected. The true danger 

lies in temporarily overcoming the problem by reducing 

girth limits and cutting cycles. 

In the aggregate, both silvicultural systems have their 

pros and cons. But trying to apply a workable silvicultural 

system is not a simple matter. It has to ensure society’s 

needs are met by harvesting the forest without degrading 

it. Despite the many mistakes and miscalculations, more 

has been done to develop management systems for 

dipterocarp forests. Nonetheless, detractors may 

emphasise that there is very little management in reality. 

That aside, it must be stated that if ever management of 

tropical forests is possible, the best chances are with 

the dipterocarp forests. Their special attributes have 

endowed them with several advantages in terms of easy 

regeneration, fast growth, and a rich commercial timber 

stand. So the silvicultural systems employed should 

attempt to enhance and exploit the special attributes of 

these forests. 

As for the silvicultural system, no doubt we can argue 

in favour of selection fellings for the existing 

dipterocarp forests. The advantages include long 

regeneration period for seedling recruitment, enhanced 

biodiversity, guarantee of future crops from advance 

growth that is retained, and retaining of species and grades


which may become marketable in the future. But 

maltreatment of the forest has become commonplace. 

The short cutting cycles have resulted in doubling of 

coupe areas, but almost as much timber as in a 

shelterwood cutting has been harvested. So are the 

problems of re-entry to logged over coupes as timber 

scarcity develops. Next, selection felling is regularly 

abused with the removal of the best stems without any 

attempt to redress the balance by simultaneous removal 

of the poorer material that can lead to genetic 

impoverishment of the forests. Perhaps the stage has 

arrived where management in the true sense can be 

introduced. This of course requires that besides paying 

proper attention to the silvicultural systems and 

harvesting methods, management must pay heed to other 

aspects like the preservation of ecological functions, 

conservation of biodiversity, and maintaining the integrity 

of the forest. In addition, the social issues that may 

impact on the management of a forest must be given a 

higher priority. 

Good management is indispensable whatever the 

silvicultural systems. An inappropriate silvicultural 

system may mean that the maximum productivity of the 

forest has not been captured. But what really sets back 

tropical forests is poor harvesting practices. Usually, 

harvests exceed growth rates. Few of the silvicultural 

tendings are done, further delaying the growth of the crop. 

Logging using skidder-tractor systems is exceedingly 

damaging to the soil and the standing residuals. Soil 

damage, in terms of erosion and compaction is 

exceedingly heavy. The immediate need is to adopt 

harvesting practices that minimise such damage. A swing 

in that direction has begun in Sabah. Already, one forest 

reserve is being managed under tight prescriptions. 

Skidder-tractors are heavily controlled and limited to prealigned 

trails only, and are only operable on slopes below 

15 o . On steeper slopes, a long range cable crane system 

is used which does only limited damage to the soils and 

residual trees. Such developments provide us with the 

optimism so much needed in tropical forest management. 

Additional Research Needs 

1. Management systems have been applied universally 

over the landscape without regard to site and timber 

stand characteristics. This cannot be ecologically 

optimal. Intensive management procedures should be 

developed whereby silvicultural systems are applied 

145 

that are more specific to the site (site categories, 

floristic groups, etc.). 

2. Harvesting damage can be easily controlled, and the 

improvements realised will be immediate and several 

fold. Besides research to lower harvesting damage, 

standards for allowable harvesting damage should be 

drawn. 

3. There is still much uncertainty about cutting cycles 

in selection fellings. The growth data available from 

few sites are broadly applied to large areas. Not only 

should existing growth data be reviewed rigourously 

so as to derive more appropriate cutting cycles, 

additional growth plots should be set up so all the 

different forest types are included. 

4. In selection fellings, seedling regeneration and 

growth are often not given attention. Studies should 

be initiated to determine post-harvest fruiting and 

seedling regeneration characteristics, and tending 

procedures. 

5. The response of advance growth to liberation 

treatments requires further investigation. Their 

reaction to heavy isolation, injury, soil disturbances 

and water stress should be studied. Will selectively 

logged forests require further crown liberation to 

optimise growth? Will promoting dense crop 

regrowth affect soil-moisture balance? 

6. No data are widely available on regenerated stands 

managed under Shelterwood Systems. Such stands 

should be reexamined. The structure of the stand and 

regrowth composition would help illuminate the 

effect of improvement fellings and climber cutting 

treatments. 

7. The basis for sustaining long-term forest production 

depends on the soil characteristics and organic matter 

accumulation. The impacts of harvesting on the 

nutrient cycles have to be further investigated. What 

will be the impacts of whole tree harvesting on 

nutrient cycles? 

8. With increase in utilisation of lesser-known or lesserused 

species, will selective logging be the same as 

in the past? How will this affect regeneration and 

composition of future crops? 

9. With changes to the future crops likely to take place, 

gregarious and very common species may become 

even more important for management. Autecological 

studies of these species are needed to fine-tune the 

management to favour such species for future crops.


10.In most forests selective logging will soon enter into 

the second cut. Logged over forest will be the future 

source of timber in the region. Investigations are 

needed on the consequences of a second cut on future 

production, forest structure, floristic composition, 

and seedling regeneration. 

11. Selection of species for enrichment planting programmes 

is still primarily ad hoc, usually based on 

what is available. Incidental observations have suggested 

that there is a core of species among 

dipterocarps (e.g. Shorea spp. (engkabangs), 

Dryobalanops aromatica, Shorea trapezifolia, and 

others) which share characteristics including fast 

growth and regular fruiting. Such species should be 

developed for planting, and their tolerances and 

growth requirements (light, water) should be investigated 

further. 

Acknowledgments 

The idea of producing the series of reports on 

dipterocarps that eventually led to this book was first 

proposed by G. Maury-Lechon, and I am indebted to her 

for the support. Likewise, C. Cossalter (Center for 

International Forestry Research) was a constant support 

throughout. Finally, I would like to thank the two 

reviewers, F.E. Putz (Center for International Forestry 

Research) and P. Burgess, for their detailed criticism of 

an earlier draft. P. Burgess was particularly helpful in 

providing many notes that were useful in the finalisation 

of this chapter. 

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Plantations 

G. Weinland 

Introduction 

Research on establishment and maintenance of 

dipterocarp plantations has been pursued now for almost 

seventy years. Efforts were especially intensive in three 

countries: India, Indonesia and Malaysia. In India the 

research concentrated mainly on Shorea robusta because 

of its abundance and its significance for agroforestry 

systems. In Indonesia and Malaysia and some other 

countries of the Indo-Malayan region a wider range of 

dipterocarp species was investigated. The research 

covered the whole range of plantation problems, albeit 

not with the same species over the whole range. Probably 

with exception of S. robusta no other dipterocarp species 

has been so well studied for operational schemes. On the 

whole, young dipterocarp plants were considered 

sensitive, delicate, and unsuitable for even-aged 

plantations but appropriate for enrichment planting. The 

fear of over-exposing sensitive young dipterocarp plants 

to light, however, has led to frequent failures of planting 

operations. It was thought that the young plants needed 

overhead shade for survival and good growth. The wide 

tolerance variation among different dipterocarp species, 

and their changes with age, were not recognised. 

In India, the earliest plantation efforts recorded are 

for Shorea robusta in 1860 at Barielly in Uttar Pradesh 

and Hopea parviflora in 1880 in South Kanara, Karnataka. 

Hopea was underplanted in teak plantations as a possible 

second storey crop in the coastal plains. Around 1890, 

taungya systems were started in West Bengal and Uttar 

Pradesh. This still continues, but on a reduced scale as 

there is progressively less and less clear-felling of forests. 

In Uttar Pradesh the main dipterocarp species was S. 

robusta, while in West Bengal, which has a more humid 

climate with less seasonality of rainfall, S. robusta was 

mixed with Chukrassia tabularis and Michelia champaca. 

The practice was to sow seeds in lines. Around 1910, 

Hopea parviflora, Dipterocarpus turbinatus and Vateria 

indica were raised in a clear-felled area in Makut 

Chapter 9 

(Karnataka). In South Kanara district, the home of five 

Hopea spp., techniques for raising H. parviflora and H. 

wightiana were already perfected by this time. The two 

species are raised together in private woodlots by local 

people, H. parviflora for timber and poles and H. 

wightiana for fuel wood. Currently, all these species are 

being planted for restoration of degraded rain forests and 

re-afforestation of barren land. The nursery techniques 

for some of these species have been standardised (Rai 

1983) and experimental results on restoration of degraded 

rain forests have been reported (Rai 1990). In the 

Andaman Islands, the Andaman Canopy Lifting System 

was developed to secure the regeneration of dipterocarp 

species (Chengappa 1944). To ensure regeneration of 

Dipterocarpus macrocarpus in North East India, a system 

called Aided Natural Regeneration involving 

supplementary planting of dipterocarps, is popular. 

Dipterocarp plantation research or research with relevance 

to dipterocarps covered a very wide range. The majority 

of the research was devoted to Shorea robusta. Aspects 

especially investigated were seed procurement/production 

and germination (e.g., Verma and Sharma 1978, Rai 1983, 

Prasad and Parvez-Jalil 1987), soils and nutrition (e.g., 

Bhatnagar 1978), rehabilitation of degraded sites (e.g., 

Prasad 1988, Rai 1990), pests and diseases (e.g., Harsh 

et al. 1989, Sen-Sarma and Thakur 1986) and agroforestry 

(e.g., Jha et al. 1991). In situ gene conservation of Vateria 

indica is carried out in the Western Ghats (Negi 1994). 

Troup’s Indian Silviculture (1980) gives a full account 

of silviculture in India and Burma and contains in the 

second volume the complete silviculture of sal (Shorea 

robusta) including plantation silviculture. Additionally, 

it contains the silvicultural characteristics of the following 

dipterocarp species: Shorea assamica, S. talura (syn. 

roxburghii), S. tumbuggaia, Dipterocarpus alatus, D. 

bourdilloni, D. costatus, D. grandiflorus, D. indicus (syn. 

turbinatus), D. kerrii, D. macrocarpus, D. pilosus, D. 

tuberculatus, D. turbinatus, Hopea glabra, H. odorata, 

H. parviflora, H. utilis, H. wightiana, Vateria

Plantations 152 

macrocarpa, V. indica, Vatica lanceaefolia and V. 

roxburghiana. A comprehensive description of the 

dipterocarps of South Asia is contained in RAPA 

Monograph 4/85 (FAO 1985). 

In Nepal, research on dipterocarps has concentrated 

on the management of sal (Shorea robusta) forests and 

on forest seeds and nursery procedures (e.g., Napier and 

Robbins 1989). In Pakistan, Chowdhury (1955) 

described the silvicultural problems of S. robusta and 

Amam (1970) trials of direct sowing. In Bangladesh, 

systematic planting of S. robusta started last century 

(1856) within the traditional taungya system. Since the 

late 1970s there are greater efforts to improve the 

management of the dipterocarp species (Das 1982). 

Subsequently, research has been carried out on 

propagation techniques (Banik 1980, Rashid and 

Serjuddoula 1986, Haque et al. 1985, Serjuddoula and 

Rahman 1985). Jones and Das (1979) developed a 

programme for the procurement of improved forest tree 

seeds, which is now the task of the National Tree Seed 

Center established in 1986 (Mok 1994). The Species 

Improvement Programme includes the plus tree selection 

of Dipterocarpus turbinatus and Hopea odorata (Nandy 

and Chowdury 1994). Dipterocarp species under 

investigation are: Anisoptera glabra, Dipterocarpus 

costatus, D. pilosus, D. turbinatus, Hopea odorata and 

Shorea robusta. 

In the past, the plantation efforts in Thailand 

focussed on planting Tectona grandis and fast-growing 

exotic species. Plantations involving dipterocarps have 

been established since the 1980s. Consequently, research 

on dipterocarps has been intensified. A description of the 

dipterocarps of mainland South East Asia has been 

prepared by Smitinand and Santisuk (1981) and of the 

silvicultural ecology of the dipterocarps of Thailand by 

Smitinand et al. (1980). Both contain information on 

silvical aspects. Research has been concentrating on 

collection, storage and germination of seeds and on 

mycorrhizae (e.g., Khemnark 1980, Panochit et al. 1984, 

Panochit et al. 1986, Chalermpongse 1987, Boontawee 

and Nutivijarn 1991, Linington 1991, Kantarli 1993). 

Concerning dipterocarp planting stock propagation the 

ASEAN Forest Tree Seed Centre concentrates on 

vegetative propagation (Mok 1994). Dipterocarpus 

alatus, Hopea odorata and Shorea siamensis, amongst 

others, are priority species for reforestation activities and 

D. alatus and D. turbinatus are included in the gene 

conservation programme (Sa-Ardavut 1994). Species 

which have received attention are: Anisoptera costata, 

Dipterocarpus alatus, D. costatus, D. intricatus, D. 

macrocarpus, D. obtusifolius, D. tuberculatus, Hopea 

ferrea, H. odorata, Shorea henryana, S. obtusa, S. 

roxburghii and S. siamensis. 

In Vietnam, some plantation work on an 

experimental scale is carried out in Dong Nai Province, 

in the Central Highlands and in Daklak Province (Doan 

1985, Vu 1991, Dinh 1992). Several studies on the 

distribution of dipterocarps, and on the structure and 

dynamics of dipterocarp forests in Vietnam were carried 

out which contain information on silvical characters of 

the dipterocarps (e.g., Nguyen Nghia Thin 1985, Vu Van 

Dung 1985). Bieberstein et al. (1985) investigated the 

possibilities of rehabilitating areas devastated during the 

Vietnam War. Species investigated were: Dipterocarpus 

alatus, Hopea odorata and Anisoptera costata. Research 

on various other aspects was carried out on these species 

and, additionally, Dipterocarpus dyeri, D. tuberculatus, 

D. obtusifolius, Shorea obtusa, S. roxburghii, S. thorelii, 

S. siamensis and Vatica odorata. 

In Cambodia the phenology and germination 

behaviour of Hopea odorata has been investigated by 

Tixier (1973). 

In Peninsular Malaysia planting of dipterocarps 

started in 1900 when Neobalanocarpus heimii was lineplanted 

in forest reserves but was discontinued when 

Commercial Regeneration Fellings were introduced in 

1918. Between 1929 and 1941 experimental plantations 

of dipterocarps were started at the Forest Research 

Institute Malaysia. Main dipterocarp species planted were 

Anisoptera scaphula, A. laevis, Dipterocarpus baudii, 

Dryobalanops aromatica, D. oblongifolia, Shorea 

acuminata, S. curtisii, S. leprosula, S. macroptera, S. 

macrophylla, S. ovalis, S. parvifolia, S. platyclados and 

S. sumatrana. Dipterocarps were later used in enrichment 

plantings (e.g., Tang and Wadley 1976). Main species 

planted were those of the fast-growing hardwoods. 

Enrichment planting is still pursued, albeit on low scale 

(Chin et al. 1995). Barnard (1954) summarised the 

knowledge on artificial regeneration of dipterocarps 

describing the operations from planting stock 

procurement to post-planting tending. Wyatt-Smith 

(1963b) furthered the knowledge on enrichment planting 

and presented information on choice of species and 

silvicultural operations up to the tending of the established 

crop. The review on planting high quality timber species 

by Appanah and Weinland (1993) presents an overview


on silvics and silviculture of many high quality timber 

tree species that have been planted in Malaysia. On present 

knowledge, the most promising dipterocarp plantation 

species for Peninsular Malaysia are: Anisoptera laevis, 

A. scaphula, Dipterocarpus baudii, D. costulatus, D. 

kerrii, Dryobalanops aromatica, Hopea odorata, Shorea 

acuminata, S. leprosula, S. macrophylla, S. macroptera, 

S. ovalis, S. parvifolia, S. platyclados (Wyatt-Smith 

1963b, Zuhaidi and Weinland 1994, Darus et al. 1994). 

Darus et al. (1994) carried out plus tree selection for 

Shorea leprosula and S. parvifolia, identified a seed 

production area for S. lepidota and included several other 

dipterocarps in a clonal selection programme and field 

tests. 

Sarawak embarked on plantations of dipterocarps 

in the 1920s by planting Shorea macrophylla. While such 

plantings were pursued on a small scale until 1975 

(Kendawang 1995), the state commenced large-scale 

plantings of dipterocarps in 1979 after disappointing 

results were obtained from research on exotic fastgrowing 

species (Mok 1994). Dipterocarp plantations are 

established within the Reforestation Programme for 

Permanent Forest Estates on areas degraded by shifting 

cultivation (Kendawang 1995). About 4940 ha have been 

planted on an operational scale with Shorea species of 

the pinanga group, especially Shorea macrophylla 

(Kendawang 1995). These plantings are based on a 

species-site matching procedure (e.g., Butt and Sia 1982, 

Ting 1986). 

In Sabah, dipterocarp plantations have, with the 

exception of the enrichment plantings under the Innoprise- 

Face Foundation Rainforest Rehabilitation Project 

(Moura-Costa 1993, Moura-Costa and Lundoh 1993, 

1994), only been established on an experimental scale. 

Until 1994 about 700 ha had been planted within the Face 

Foundation Project. Dipterocarp species used are: 

Dipterocarpus spp., Dryobalanops lanceolata, Hopea 

nervosa, Parashorea malaanonan, Shorea argentifolia, 

S. johorensis, S. leprosula, S. macrophylla, S. ovalis and 

S. parvifolia. The plantation target is 25 000 ha. 

In Indonesia, the establishment of experimental 

plantations (e.g., Darmaga, Haurbentes, Pasir Hantap, 

Purbah Tongah and Sangau) started at the end of the 40s 

(Butarbutar 1986, Masano 1991, Masano et al. 1987, 

Masano and Alrasjid 1991, Omon 1986). Apart from these 

experimental plantations, planting of dipterocarps was 

mainly enrichment planting in the concession areas and 

regularly carried out in the state-owned concession 

INHUTANI II in South Kalimantan (Mok 1994). Now, 

the Indonesian Selective Cutting and Planting System 

prescribes reforesting all logged areas and since the 

beginning of the 90s large-scale cutting propagation is 

carried out. Research on dipterocarps has covered a wide 

field ranging from seed procurement and testing (e.g., 

Masano 1988a, b, Syamsuwida and Kurniaty 1989), 

vegetative plant propagation (e.g., Smits 1987, 1993, 

Umboh et al. 1988), plantation stock trials (e.g., Wardani 

et al. 1987 Siagian et al. 1989b), mycorrhizal symbiosis 

(e.g., Smits 1982, Santoso et al. 1989, Santoso 1991) to 

agroforestry problems (e.g., Kartawinata and Satjapradja 

1983, Sardjono 1990). It appears that no specific tree 

improvement programme for dipterocarps has been 

initiated (Sunarya 1994). Dipterocarp species which 

received attention were: Dipterocarpus grandiflorus, D. 

retusus, D. tempehes, Dryobalanops lanceolata, Hopea 

bancana, H. mengerawan, H. odorata, H. sangal, Shorea 

guiso, S. johorensis, S. leprosula, S. macrophylla, S. 

mecistopteryx, S. multiflora, S. ovalis, S. palembanica, 

S. parvifolia, S. pauciflora, S. pinanga, S. platyclados, S. 

selanica, S. seminis and S. smithiana. Recently, a manual 

for the dipterocarp light hardwoods for Borneo Island 

has been compiled by Newman et al. (1996). 

In the Philippines first research efforts on dipterocarp 

plantation problems commenced in the 30s (e.g., Caguioa 

1938, Lantion 1938). The research work continues (e.g., 

Anon. 1982, 1991). Some experimental plantations were 

established and private companies participated in the 

plantation programmes (e.g., Notonton 1985). 

Underplanting was carried out in Benguet pine 

plantations (Anon. 1960) with success. Enrichment 

planting was rarely done with dipterocarps, but with fastgrowing 

exotic trees species such as Paraserianthes 

falcataria. Underplanting and enrichment planting trials 

with dipterocarps started late (Chinte 1982, Mauricio 

1987a, Abalus et al. 1991). Emphasis of research was on 

germination trials (e.g., Basada 1979, Garcia et al. 1983), 

seedling trials (e.g., Bruzon and Serna 1980, Gianan and 

Peregrino 1986), use of wildings as planting stock (e.g., 

Lantion 1938, Penonia 1972), planting trials (e.g., 

Tomboc and Basada 1978, Miyazaki 1989). Agpaoa et 

al. (1976) provided valuable information on planting 

techniques. A tree improvement programme for 

dipterocarps has been launched which includes seed 

production area and plus tree selection, establishment of 

clonal gardens and gene conservation (Rosario and 

Abarquez 1994). Promising dipterocarp plantation species


are: Dipterocarpus grandiflorus, D. warburgii, 

Parashorea plicata, Shorea almon, S. guiso, S. 

negrosensis, S. polysperma and S. squamata. Newman et 

al. (1996) compiled a manual of dipterocarps for the 

Philippines. 

Silvics 

Silvics deals with the life history and general 

characteristics of forest trees and stands particularly 

refering to locality factors as a basis for the practice of 

silviculture. 

For tree species of the high forests (a closed forest of 

tall trees), tolerance is their ability to grow satisfactorily 

in the shade of and in competition with other trees. If 

intolerant of shade, a species is termed a ‘light demander’, 

if tolerant, a ‘shade bearer’. Discussions on how much 

light should be given for good growth and how much 

shade should be retained started early. Sanger-Davies 

(1931/1932) considered most of the commercial 

dipterocarp species as light demanders which should be 

given full overhead light and full space for maximum 

development. While larger plants need full light for good 

growth, young seedlings need a shelter either from 

existing belukar or from planted nurse crops. Indeed, 

planting of dipterocarps under a nurse crop (e.g., 

Paraserianthes falcataria) was successful in the 

experimental plantations in Indonesia (e.g., Masano et 

al. 1987) and Malaysia (Barnard 1954) and elsewhere in 

the region (e.g., Doan 1985). All shading experiments 

showed without doubt that optimal growth of dipterocarp 

seedlings is only achieved under partially shaded 

conditions (e.g., Nicholson 1960, Mori 1980, Sasaki and 

Mori 1981 and others). 

There is a wide range of shade tolerance among older 

seedlings/saplings of dipterocarp species which follows 

the known pattern of higher shade tolerance for late 

succession species and higher light demands for earlier 

succession species (e.g., Strugnell 1936a). Qureshi (1963) 

classified about 100 tree species (including. Shorea 

robusta) as tolerant, moderately and intolerant of shade 

in comparison to Acacia arabica which is intolerant of 

shade at every developmental stage. In Peninsular 

Malaysia, field experiments on light requirements were 

established early in conjunction with Regeneration 

Improvement Systems and a discussion on canopy 

manipulations over young regeneration ensued (e.g., 

Sanger-Davies 1931/1932, Watson 1931/1932b, Walton 

1936b). However, this type of experiment was abandoned 

when Regeneration Improvement Systems ceased in 

Malaya in the 1930s. JICA (1993, 1994) reported an 

underplanting trial where dipterocarps (Shorea 

leprosula, S. parvifolia, Dryobalanops aromatica and 

others) have been underplanted in Acacia mangium 

stands with different size gaps. The performance was best 

where two rows had been removed (9 m opening). 

Controlled (artificial) experiments are needed for base 

line information on the light requirements of species to 

be complemented by field trials where shade from natural 

vegetation is manipulated. More details on the light 

physiology of seedlings can be found in Chapter 3. 

Mycorrhizal symbiosis with dipterocarps has 

received great attention in recent years. Since this field 

is dealt with in detail in Chapter 6, only some practical 

aspects are discussed here. The importance of 

mycorrhizal symbiosis for the survival and growth of 

trees is not in question. Most of the investigations deal 

with the identification of mycorrhizal fungi and their 

strains/forms (e.g., Louis 1988) and Lee and Lim (1989) 

have reported mycorrhizal infection of dipterocarp 

seedlings in logged and undisturbed forests. Host 

specificity of mycorrhizal fungi was reported by Smits 

(1982) and it is concluded that the chance of a seedling 

finding the right fungus is better the closer the seedling 

germinates and grows to the mother tree. He explains 

the formation of eco-unit patterns as linked to such a 

preference. Whether host specificity is wide spread 

among dipterocarps remains to be investigated. 

Alexander et al. (1992) found that the root contact of 

seedlings with mature trees is important for the infection 

with mycorrhizae which would have a bearing on the 

design of regeneration systems. The retention of mature 

trees seems to be important for this reason. Turner et al. 

(1993) investigated the effect of fertilser application on 

dipterocarp seedling growth and mycorrhizal infection. 

The application involved 10 g m -2 N, P 2 O 5 and K 2 O to 

Shorea macroptera seedlings grown in pots of forest 

soil (nursery condition). The results showed that 

mycorrhizal infection was significantly higher for 

fertilised seedlings. Oldeman (1990) draws attention to 

the fact that mycorrhizal symbiosis occurs particularly 

on poorer, acid soils and suspects that by changing the 

chemical status of the soil through fertilisation, 

mycorrhizal functioning might be impaired. Santoso 

(1987, 1989) showed that there is an increase in shoot/ 

ratio, dry weight of leaves, roots, stem diameter, as well 

as absorption potential for nutrients among several


dipterocarps when inoculated with Scleroderma 

columnare. One dipterocarp species (Shorea pinanga) 

showed better results when inoculated with Russula 

amatic. Inoculation techniques for nurseries are 

described for example by Bakshi (1980), Khemnark 

(1980), Smits (1987) and Tacon et al. (1988). 

Mycorrhizal research has yielded practical incoculation 

techniques for nurseries. 

Site requirements of dipterocarp species have only 

been examined systematically for Shorea robusta (e.g., 

by Yadav and Mathur 1962, Bhatnagar 1966). Butt and 

Sia (1982) and Ting (1986) touch on the problem in their 

evaluation for reforestation and rehabilitation projects 

in Sarawak, however, assignment of species to site was 

not based on species-adaptation trials. Most of the 

information has still to be obtained from species 

compilations (e.g., Foxworthy 1932, Symington 1974, 

Smitinand et al. 1980, Ashton 1982) which contain 

information on the natural habitat of the species, from 

which in many cases generalised inferences to the site 

requirements under plantation conditions can be made. 

A systematic approach to this problem through species 

adaptation trials is urgently needed. Such trials would 

include the species most likely to be used for plantation 

programmes. Field operations before and during planting 

site operation change site conditions, foremost the soilphysical 

structure so site management with low negative 

impact is important for the success of a plantation. Dabral 

et al. (1984) reported impaired rooting behaviour of 

Shorea robusta in compacted soil. Kamaruzaman (1988) 

showed that bulk densities in crawler tractor tracks 

declined to 1.52 g cm -3 , at which rooting is severely 

impaired. Gupta (1955) investigated compaction, 

erodibility and other soil-morphological features in 

Shorea robusta forests and taungya plantations. In the 

latter, cultivation and continued exposure had caused hard 

pans to develop which resulted in reduced seepage and 

increased erodibility. 

When planting a species the silvicultural characters 

of the trees should be known. Stand density regimes 

depend on a clear understanding of the growth form, 

which is the characteristic shape, posture, and mode of 

growth of a tree (Ford-Robertson 1983). Troup (1980) 

describes silvicultural characters of 22 dipterocarp 

species besides Shorea robusta. Additional work 

includes that of Kadambi (1954, 1957), but, these reports 

cover only a small percentage of the total species of 

dipterocarps. Dipterocarp species differ considerably in 

terms of crown structure, branching habit, growth 

dynamics etc. Hallé and Ng (1981) worked on crown 

architecture, especially reiteration and aggregation. 

Zuhaidi and Weinland (1994) and Appanah and Weinland 

(1993) give information on growth form of some 

commercially important dipterocarp species for planting 

and mention the species: Anisoptera laevis, A. scaphula, 

Dryobalanops aromatica, D. oblongifolia, Hopea 

odorata, Shorea acuminata, S. leprosula, S. 

macroptera, S. macrophylla, S. parvifolia, S. 

platyclados and S. ovalis. Information on speciesspecific 

growth dynamics, which is required for the 

design of species mixtures, is contained, e.g., in Howard 

(1925), Edwards and Mead (1930), Griffith and Bakshi 

Sant Ram (1943), Mathauda (1953b, 1955), Ng and Tang 

(1974), Rai (1979, 1981a, b, 1989), Masano et al. 

(1987), Primack et al. (1989), Zuhaidi et al. (1994). 

Within the group of the fast-growing light hardwoods 

(e.g., Shorea leprosula, S. parvifolia, S. ovalis and S. 

macrophylla) important differences between species in 

growth dynamics seem to exist (e.g., Wyatt-Smith 1963b, 

Zuhaidi and Weinland 1994, Zuhaidi et al. 1994). 

The following characters of a tree species to be 

planted should be known to the practising silviculturist: 

(i) control of side branch development by the leader shoot 

(apical dominance), (ii) phototropic sensitivity 

(phototropism), (iii) self-pruning capacity, (iv) type of 

branch formation, and (v) growth rates and growth 

dynamics. 

In conclusion, there is a pressing need to build up 

information on the silvical and silvicultural properties 

(stress tolerance, growth form, mode of growth) of a 

defined set of the most promising species for plantations 

and on the site requirements (site adaptation) using 

standardised methods. 

Stand Regeneration and Establishment 

Regeneration of a forest is the renewal of a tree crop, 

whether by natural or artificial means. Renewal by selfsown 

seed is termed ‘natural regeneration’, by sowing 

or planting ‘artificial regeneration’. Formation of stands 

means all the operations contributing to the creation of 

a new crop up to the stage where it is considered 

established, i.e. from seed procurement, as for a nursery, 

to early tending. Establishment is the process of 

developing a crop to the stage at which the young trees 

may be considered established, i.e. safe from normal 

adverse influences e.g., drought, weeds or browsing,


and no longer in need of special protection or special 

tending, but only routine cleaning, thinning and pruning 

(definition according to Ford-Robertson 1983). 

Species Choice 

Up to now, little systematic species elimination work 

has been done on plantation species with the exception 

of Shorea robusta, around which a complete silvicultural 

and agri-silvicultural system has developed. Anderson 

(1975) proposed Shorea spp. of the pinanga group (e.g., 

Shorea macrophylla, S. pinanga and S. stenoptera) as 

an agricultural crop. Jha et al. (1991) have discussed the 

selection and evaluation of suitable tree species and food 

crops for agro-forestry systems which include Shorea 

robusta. 

In the Malaysian context Wyatt-Smith (1963b) 

presented a list of species with promise for plantation 

establishment. They were selected on the basis of 16 

criteria, for example, fruiting frequence, seed viability, 

collection and nursery handling, fast, early height growth, 

natural bole form, self-pruning capacity, timber 

properties, etc. The species proposed were: Anisoptera 

laevis, A. scaphula, Dipterocarpus baudii, D. 

costulatus, D. grandiflorus, D. kerrii, D. verrucosus, 


odorata, Shorea acuminata, S. curtisii, S. leprosula, 

S. macrophylla, S. macroptera, S. ovalis, S. parvifolia, 

S. pauciflora and S. platyclados. 

Recently, an assessment of the dipterocarp 

plantation stands at the Forest Research Institute 

Malaysia was carried out in the field and from 

phenological and plantation records (Zuhaidi and 

Weinland 1994, Appanah and Weinland 1996). The 

indicators used were: overall diameter growth rate, initial 

height growth rate, stem shape, seedling adaptation phase, 

natural pruning capacity, cutting propagation capacity, site 

specificity, natural regeneration capacity within the 

rotation age, susceptibility to diseases and mode of 

growth. The result was that the dipterocarp species 

differed considerably in some aspects, especially in 

growth form, mode of growth, site specificity and natural 

regeneration capacity. In the case of undesirable mode 

of growth, the species was nevertheless considered 

suitable for planting, if the deficiency could be corrected 

by simple silvicultural means. As a result, 15 dipterocarp 

species were chosen for immediate inclusion into 

plantation programmes (Anisoptera laevis, A. scaphula, 

Dipterocarpus baudii, D. costulatus, D. kerrii, 


odorata, Shorea acuminata, S. leprosula, S. 

macroptera, S. macrophylla, S. parvifolia, S. 

platyclados and S. ovalis), and 2 species (S. bracteolata 

and S. curtisii) were considered promising, but were not 

included because of lack of sufficient information and 

doubtful field characteristics. For the Bornean part of 

Malaysia species could be added, such as Parashorea 

malaanonan, Shorea fallax and S. smithiana, and for 

Indonesia Dryobalanops lanceolata, Shorea laevis, S. 

macrophylla and S. selanica. The most common 

plantation species in the Philippines are Dipterocarpus 

grandiflorus, Shorea almon, S. contorta, S. guiso, S. 

polysperma and S. squamata (e.g., Assidao 1950, 

Cacanindin 1983, Abalus et al. 1991). Systematic 

species/provenance elimination trials are urgently 

needed, particularly in relation to the more pronounced 

seasonality following the extensive removal of natural 

forests in many regions of the humid tropics. 

Planting Stock Production 

Seed 

Much effort has been invested in developing methods 

for seed production, collection and handling. Generally, 

dipterocarps fruit at irregular intervals and with varying 

seed yield. On top of that, seed viability declines. This 

field is reviewed in Chapter 4. Tompsett (1991) has 

reviewed the storage aspects of dipterocarp seeds. Much 

is also known about germination (e.g., Caguioa 1938, 

Jensen 1971, Tixier 1973, Chai 1973, Masano 1988a, b, 

Ng and Mat Asri 1991, and others), the effect of 

harvesting time and sowing interval on germination 

(Haque et al. 1985), the effect of fruit ripeness upon 

germination and seedling growth of Shorea ovalis 

(Kosasih 1987), the effect of fruit collection time on 

the germination of Dryobalanops aromatica (Barnard 

1954), the effect of seed size on germination of Shorea 

contorta (Basada 1979), the effect of wing colour on 

the germination of Shorea pinanga and S. stenoptera 

(Masano 1988b) and the effect of tree girth on seed 

viability and germination of Shorea robusta (Yadav et 

al.1986). Overall, the storage/germination/viability 

aspects are sufficiently covered. 

There is definitely a lack of information on the seed 

yield from trees and stands (quantity of seeds during a 

normal seed year). Such information is only available for 

Shorea robusta (Jain 1962, Sharma 1981). In Peninsular


Malaysia, flowering and fruiting are regularly observed 

over a wide geographical and climatic range and reliable 

records are available. Darus et al. (1994) proposed the 

establishment of seed production stands in the main 

climatic regions of Peninsular Malaysia and a 

corresponding tree selection and tree improvement 

programme. Similar efforts on tree improvement 

involving dipterocarps have been made in Bangladesh 

(Nandy and Chowdury 1994), India (Negi 1994), 

Philippines (Rosario and Abarquez 1994) and in Thailand 

(Sa-Ardavut 1994). Much of the improvement work in 

the region is coordinated within the Species Improvement 

Network (Anon. 1994). 

Seedling planting stock 

‘In nursery practice, a seedling is a very young tree that 

has not been transplanted, i.e. is growing where it 

germinated’ (Ford-Robertson 1983). Seedling planting 

stock for most dipterocarp species is usually potted and 

leaves the nursery after about 9 months. The seedling 

height is about 25-50 cm. 

Generative propagation is still the prevailing method 

of plant production in dipterocarps and is technically not 

a problem if seeds are planted immediately after 

collection. Timber companies involved in propagation 

of dipterocarp seedlings have the expertise to run largescale 

dipterocarp nurseries professionally e.g., in 

Indonesia or Sabah (Moura-Costa 1993). The literature 

on the propagation of dipterocarp seedlings deals mainly 

with planting stock type (e.g., Walton 1938, Barnard 

1954, Pande 1960, Joshi 1959), sowing position of 

seeds (Serjuddoula and Rahman 1985), response of 

potted seedlings to fertilisers (Kaul et al. 1966, Bruzon 

1978, 1982, Sundralingam 1983, Sundralingam et al. 

1985), controlled mycorrhization (Garbaye 1989, 

Santoso 1989, Santoso et al. 1989), the use of bare-root 

plants (Sasaki 1980b, Mori 1981). 

As far as the age of the planting stock is concerned 

Barnard (1954) found that for most of the dipterocarp 

species planting stock between 3 and 8 months old is 

the best (e.g., Dryobalanops aromatica, Shorea 

leprosula and S. pauciflora). Hodgson (1937a) 

concluded that planting stock only a few months old is 

more likely to survive than older material. Seedlings of 

Anisoptera sp., Dryobalanops aromatica and 

Neobalanocarpus heimii were planted with cotyledons 

still attached. While D. aromatica was destroyed by 

rodents, the two other species survived. Kuraishy (1942) 

transplanted 6-week old seedlings of Shorea robusta 

successfully. 

Lamprecht (1989) proposes the use of 15-20 cm 

tall planting stock for economic and handling reasons. 

Which plant size to choose, should depend on the 

condition of the planting sites, that is, those with 

intensive weed growth require larger planting stock. To 

reduce the amount of weeding it is preferable to plant 

seedlings which are large enough to overcome weed 

competition at an early stage although growth rates might 

not be better than those of smaller planting stock. Planting 

stock size is an important aspect but root:shoot ratio, 

leaf area and diameter:height ratio are as important. 

Sturdy plants with a low root-collar:shoot ratio tend to 

form roots faster and are better equipped to withstand 

drought stress. In a trial carried out by Moura-Costa (not 

dated) initial height growth rates were significantly better 

for sturdier plants. Species tested were: Dipterocarpus 

gracilis, Dryobalanops lanceolata, Parashorea 

malaanonan, Shorea johorensis, S. leprosula, S. ovalis 

and S. parvifolia. 

Type of planting stock is another factor to be 

considered. Potted seedlings proved to be superior to 

bare-root planting stock (e.g., Anon. 1948a, Barnard 

1949b). With the exception of a few hardy species, the 

survival of bare-rooted stock seems to be low (e.g., Cerna 

and Abarquez 1959). Rayos (1940) tested the survival 

of bare-rooted seedlings of Hopea pierrei of different 

sizes with their roots stored in moist sawdust before 

planting out. Survival was inversely proportional to 

storage period and seedling size. The smallest height 

tested was 10-20cm. Prasad (1988) found in a plantation 

trial on bauxite mining land that survival and growth of 

potted S. robusta plants were superior to that of plants 

from direct sowing. 

Specific treatment of seedlings, such as shoot and 

root-pruning and the effect on growth and survival have 

been investigated. Root-pruning gave better survival and 

growth of planting stock. Walton (1938), Landon (1948b) 

and Barnard (1954) showed that survival and growth of 

Dryobalanops aromatica seedlings were superior when 

seedlings were wrenched (tap root severed) compared 

to unwrenched seedlings. The effects of shoot-pruning 

and stripping of the leaves on survival were inconclusive. 

Landon (1948b) planted Dryobalanops aromatica under 

the shade of a 20-year old Fragraea fragrans stand and 

topping, partial or total stripping of leaves had no effect 

on survival. Sasaki (1980a) pruned bare-rooted seedlings 

Shorea talura and Hopea odorata (removal of all leaves, 

all young parts of the stems and the tap root) and was


able to store them in polythene plastic bags for several 

months without loss of viability. The effect of hormone 

application on the storage of potted seedlings has been 

investigated by Siagian et al. (1989b) for Shorea selanica. 

Dabral and Ghei (1961) applied gibbelleric acid to the 

shoots of Shorea robusta seedlings but failed to boost 

root development and growth. 

There has been some systematic research on 

fertilisation of nursery planting stock. An early 

investigation into morphological symptoms of mineral 

deficiencies of nursery stock of Shorea robusta was 

carried out by Kaul et al. (1966). Deficiencies in N, P, 

K, Ca and Mg caused marked symptoms in both shoot 

and root development. Deficiencies in N, P and Mg 

affected height increment especially, while root 

development was affected by deficiencies in all minerals. 

Bruzon (1978, 1982) investigated the optimal NPK 

(14:14:14) fertilisation of Shorea contorta nursery 

seedlings of an average height of 15 cm grown in a 

mixture of potting medium and forest soil. The seedlings 

were fertilised (control, 2, 4, 6 and 8g) three times at an 

interval of approximately one month. The survival was 

best in the unfertilised control and with applications of 

2g and 4g per seedling. Height and diameter growth were 

best in the 2g, 4g and 6g treatments. Survival was 

significantly reduced with application of 4 and 8g of 

fertiliser. Fertilisation with 2g NPK per plant is 

recommended. Bhatnagar (1978) tested the nutritional 

requirements of Dipterocarpus macrocarpus seedlings. 

For 1 year the potted seedlings were fertilised every two 

weeks with 450 and 900 mg NPK solution. Achieved 

height and dry weight were greatest with N and P at 900 

mg application and K at 450 mg application. 

Sundralingam (1983) investigated the best height growth 

response of below 1-year old Dryobalanops aromatica 

and D. oblongifolia seedlings by fertilising the seedlings 

in a shaded nursery with 50 mg P 2 O 5 (as superphosphate) 

and 300 mg N (applied as ammonium sulphate at 2-month 

intervals) per plant. The height growth was reduced to 

that of the control plants when the amount of phosphorus 

was doubled. In another experiment Sundralingam et al. 

(1985) tested the nitrogen and phosphorus requirements 

of Shorea ovalis seedlings in sand culture by fertilising 

seedlings with various dosages at 2-4 week intervals. 

After 8 months it was found that the optimal N dosage 

was 80 mg/plant per application and the optimal P dosage 

4 mg/plant per application. 

Another method to boost performance is through 

mycorrhizal inoculation. Garbaye (1989) reviewed the 

literature on natural and controlled mycorrhizal infection 

in tropical plantations including dipterocarp plantations. 

Santoso (1988, 1989, 1991) tested inocula of Boletus, 

Russula (3 species) and Scleroderma spp. on 45-day old 

seedlings of Hopea odorata, Shorea compressa, S. 

pinanga, S. stenoptera and Vatica sumatrana and after 

6 months growth parameters such as diameter, dry weight 

of leaves, stems and roots were increased. Responses 

were best in Hopea odorata, Shorea stenoptera and 

Vatica sumatrana with Scleroderma spp., while 

responses of S. pinanga were best with Russula (species 

2). Santoso et al. (1989) found that under the same 

experimental conditions as above all inocula increased 

the accumulation of micro-nutrients (Fe, Mn, Cu, Zn and 

Al) in leaves, stems and roots of the seedlings. Turner et 

al. (1993) investigated the effect of fertiliser application 

on mycorrhizal infection. NPK (combined N, P 2 O 5 and 

K 2 O) was applied at a rate of 10g m -2 to potted Shorea 

macroptera seedlings (potting medium: forest soil). In 

fertilised pots ectomycorrhizal infection was increased 

but the correlation between extent of infection and 

growth was closer in unfertilised seedlings, suggesting 

that seedlings may only be responsive to fertiliser 

addition when grown at very low nutrient availabilities. 

Mycorrhizal infection may be important under such 

conditions. Smits (1982, 1987, 1993) pointed out the 

importance of mycorrhizal infection in nurseries and 

described controlled inoculation 

Wilding planting stock 

‘A wilding is a naturally-grown, in contrast to a nurseryraised 

seedling, sometimes used in forest planting when 

nursery stock is scarce’ (Ford-Robertson 1983). 

Wildings were frequently used in the past and various 

trials have been carried out with them. 

Wildings have been successfully used for planting 

places lacking natural regeneration. Capellan (1961) 

tested the possibilities of Parashorea plicata and Shorea 

contorta wildings as planting stock and P. plicata had 

better survival than S. contorta. Barnard (1954) mentions 

that wildings of Shorea macrophylla, S. multiflora, 

Dipterocarpus baudii and Neobalanocarpus heimii 

were successfully planted. Gill (1970), while reviewing 

experimental enrichment planting work in West Malaysia, 

found that transplanting bare-rooted wildings of 

Anisoptera laevis, Shorea curtisii, S. leprosula, S.


parvifolia and S. platyclados is promising. Fox (1971/ 

72) investigated the performance of wilding stock of 

Dipterocarpus caudiferus, Dryobalanops lanceolata 

and Parashorea tomentella, of which D. lanceolata 

performed best. This was confirmed in a trial by Chai 

(1975). Jafarsidik and Sutomo (1988) developed a field 

guide for the identification of dipterocarp wildings for 

a production forest in West Sumatra including wildings 

of the genera Anisoptera, Dipterocarpus, Hopea, Parashorea 

and Shorea. Wardani and Jafarsidik (1988) put together 

a field guide for the identification of dipterocarp wildings 

of the genera Dipterocarpus, Dryobalanops, Hopea 

and Shorea for a forest area in West Kalimantan. Mauricio 

(1957) investigated factors which influence the performance 

of wildings of Parashorea plicata and Shorea contorta 

to determine: (i) the effect of the wilding size on survival, 

(ii) the time the wildings require to adapt to the planting 

site, and (iii) the most suitable size. In this experiment 

P. plicata had a higher survival, specially at heights of 

20 cm and less. Lantion (1938) tested the performance 

and the behaviour of wildings of Dipterocarpus grandiflorus 

and Shorea teysmanniana and smaller plants had higher 

survival. The Forest Department in Malaya had a trial 

of Dryobalanops oblongifolia and D. aromatica in the 

nursery where six month-old wildings were transplanted 

into small claypots. D. oblongifolia wildings had 76% 

survival in the nursery and about 90% survival in the 

field after six months whereas D. aromatica wildings 

had a survival of 100% in the field (Anon. 1951). 

Rayos (1940) tested the effect of storage time of 

wildings of Hopea pierrei on survival by covering their 

roots with moist sawdust. Survival was higher the shorter 

the storage time and it was greater for seedlings 10-20 

cm high than for those in other height classes. No effect 

of storage time on survival rate was found by Siagian et 

al. (1989b). Moura-Costa (1993) obtained high survival 

rates with wildings from Parashorea malaanonan, 

Shorea parvifolia and Dryobalanops lanceolata. 

Forest-pulled seedlings were watered and kept in plastic 

covered chambers with high humidity until a new root 

system had formed. Survival in the nursery was up to 95% 

in a large scale operation. Barnard and Setten (1955) used 

wildings in an investigation on the effect of planting patch 

cultivation but found no difference to planting in 

unprepared patches. Wardani (1989) found that shoot and 

root-pruning increased survival of wildings. Hormone 

treatment of wildings for growth stimulation has been 

reported for Vatica sumatrana (Masano and Omon 

1985), for Dipterocarpus retusus (Omon and Masano 

1986), for Shorea platyclados (Napitupulu and Supriana 

1987) and for Shorea selanica, (Siagian et al. 1989b). 

Increased survival rates were found for S. platyclados 

and V. sumatrana but not for D. retusus and S. selanica. 

The Forest Department Sarawak reported the 

establishment of wilding nurseries as seedling reservoirs 

(Anon. 1948c). Before a heavy seedfall, cleanings were 

made beneath fruiting trees to form natural ‘nurseries’ 

which were used later to plant forests with low natural 

regeneration or in secondary vegetation. The seedling 

yield was excellent. 

The use of wildings is not unequivocally supported. 

Wyatt-Smith (1963b) is critical about the use of wildings 

for the following reasons: (i) transplanting large forest 

seedlings is generally not successful, (ii) small forest 

seedlings suffer high mortality during the first two years, 

and (iii) the pool of young forest seedlings cannot serve 

as a continuous supply for large-scale plantations. 

Vegetative propagation 

Among the methods of vegetative propagation of grafting, 

air layering, tissue culture and cutting propagation, the 

latter is the most commonly used technique. Plant 

production from cuttings has been intensively 

investigated. Dick and Aminah (1994) have carried out a 

thorough review on cutting propagation of dipterocarps. 

Research work has been carried out on important factors 

influencing the rooting ability of dipterocarp cuttings, 

such as rooting facilities, rooting media, source of 

cutting material, type and treatment of cutting. According 

to Dick and Aminah (1994) 56 dipterocarp species have 

been tested, among them almost all of the species 

suitable for plantations. Vegetative propagation of 

dipterocarps is increasingly successful and has been 

introduced as large-scale operations in Indonesia 

(Sutisna, personal communication). Moura-Costa (1995) 

gives a detailed description of vegetative propagation 

techniques for Dryobalanops lanceolata and several 

Shorea spp. in context of plant production for large scale 

enrichment plantings of dipterocarps in Sabah. However, 

when cutting propagation is used in plantation 

programmes, it is necessary to precede such large-scale 

application by an established procedure of selection of 

superior stock plants. Clonal propagation of selected 

material from dipterocarps is in its infancy in the whole 

region (see e.g., Finkeldey and Havmoller 1994). Moura- 

Costa (1995) discusses a procedure of selecting best 

genotypic material at the seedling stage, the so-called


‘Predictive Test for Apical Dominance’. The test has not 

yet been established for dipterocarps. 

Some research has been carried out on tissue culture 

of dipterocarps. Linington (1991) grew seedlings in vitro 

from embryos of Dipterocarpus alatus and D. 

intricatus. Other in vitro experiments were carried out 

by Smits and Struycken (1983) on leaf fragments of 

Shorea curtisii, which formed callus and roots but no 

shoots, and on nodal explants of Shorea obtusa with 

axillary buds, which formed lateral shoots but no roots. 

Suspension cultures of embryonic axes of Shorea 

roxburghii, which eventually formed whole plantlets, 

were carried out (Scott et al. 1988). Umboh et al. (1988) 

described the rejuvenation of adult trees and a three step 

bud culture for Shorea pinanga. Moura-Costa (1993) 

describes trials in tissue culture techniques for in vitro 

propagation of Dipterocarpus intricatus which were 

successful. No commercially feasible procedure has 

been developed and tissue culture cannot be introduced 

on an operational scale at this stage. 

Darus and Rasip (1990) carried out both intra and 

inter-species splice grafting of Dipterocarpus baudii, 

Shorea parvifolia, S. leprosula and S. roxburghii. It was 

successful and grafts grew faster than single-rooted 

seedlings. Chaudhari (1963) tested air-layering in 

Shorea robusta and found that it is more successful if 

carried out in the months of July and August (midmonsoon) 

when there is a full flush of green, healthy 

leaves. Zabala (1986) successfully carried out airlayering 

of Anisoptera thurifera and Shorea contorta 

but was unsuccessful with Hopea foxworthyi, H. plagata 

and Dipterocarpus grandiflorus. Air layering was 

successful in Shorea palembanica and Vatica pauciflora 

(Hallé and Kamil 1981) and in Shorea selanica (Harahap 

1972). Rashid and Serjuddoula (1986) rooted branches 

from 5 to 10-year old saplings and 50 to 80-year old 

trees of Dipterocarpus turbinatus using air-layering. 

Rooting was better on branches from old trees. 

Planting stock production of the commercially most 

important dipterocarp species, whether from seeds or 

from cuttings, has largely been solved. 

Stump plants 

Stumping is used to rejuvenate over-aged planting stock 

and in some cases, for example, Tectona grandis, it is 

applied as a method of multiplication of the planting 

stock. The use of stumped plants started early. Watson 

(1931/1932d) found that Dipterocarpus spp. can be 

successfully stumped. Hodgson (1937a) showed that 

Dipterocarpus baudii, Shorea curtisii and S. 

macroptera can be stumped, but it was unsuccessful with 

Dryobalanops aromatica, S. leprosula and S. 

pauciflora. Barnard (1956) tested stumping of 

Dipterocarpus baudii, Dryobalanops aromatica, 

Hopea helferi, Neobalanocarpus heimii, Shorea 

assamica, S. foxworthyi, S. pauciflora and S. sumatrana. 

The stumping was carried out by pruning all side roots 

close to the tap root, which itself was cut to 23 cm from 

the collar. The shoot was cut to 14 cm from the collar. 

Stumping of most species was promising but 

Dryobalanops aromatica failed and the success of 

Dipterocarpus baudii was uncertain. Sasaki (1980a) 

found that bare-root stock of Shorea talura successfully 

transplanted after stripping off all the leaves and cutting 

back the leader and the tap root. In 1985, a stumping trial 

was carried out with Dryobalanops lanceolata in East 

Kalimantan, Indonesia, which was highly successful 

(Diana 1987). A trial with 2 year old bare-rooted stump 

plants of Shorea robusta carried out in West Bengal was 

also successful (Anon. 1959). Pande (1960) found 

Shorea spp. can be stumped and he carried out some 

experiments comparing stumped plants with ball 

transplants and basket plants. Basket plants did best. 

Landon (1948b) compared stumped plants of 

Dryobalanops aromatica with potted seedlings and the 

performance of potted seedlings was superior. Pande 

(1960) obtained a similar result for Shorea robusta 

when survival and growth performance of bare-rooted 

stump plants were inferior to ball-rooted transplants and 

container plants. 

Hormone treatment for growth stimulation of stump 

plants was investigated by Masano and Omon (1985), 

Omon and Masano (1986), Srivastava et al. (1986) and 

Siagian et al. (1989a) but results were inconclusive. 

Mori (1981) investigated the effect of starch reserves 

in the stem on survival and growth of stumped bare-root 

transplants of 16 dipterocarp species. Some species 

showed high mortality after stump planting, e.g., Shorea 

curtisii, S. ovalis, Hopea nervosa, H. beccariana. 

Stimulation of root and shoot growth by growth 

regulators or fertilisers was unsuccessful in various 

species and some species survival and initial growth were 

directly related to starch reserves before planting. 

Planting site 

The positive role of an initial shelter for the newly planted 

dipterocarp trees is beyond doubt (e.g., Wyatt-Smith 

1947, Chakravarti 1948, Landon 1948b, Ardikoesoema


and Noerkamal 1955, Krishnaswamy 1956, Sudiono and 

Ardikusumah 1967). Dipterocarps, usually, are not 

planted on completely cleared sites. In enrichment 

plantings they are planted on lines cut into the forest and 

in plantations the plants are usually planted under the 

shade of a nurse crop. 

Underplanting or sowing beneath a forest canopy is 

important in restocking forests with valuable species. 

Underplanting can be done in residual stands of logged 

natural dipterocarp forests, in secondary forests, under a 

planted nurse crop or in plantations where the stocking is 

poor. The Experimental Forests of West Java (Darmaga 

and Haurbentes) were established by underplanting. 

Ardikoesoema and Noerkamal (1955) give an account of 

the establishment of the Shorea leprosula stand in 

Haurbentes. Two month-old seedlings were planted under 

the shelter of 2 year-old Paraserianthes falcataria which 

was removed after 5-6 years. At the age of 15 years the 

stand had passed the pole stage. The experimental 

plantations in the area of the Forest Research Institute 

Malaysia were partially established under nurse crop, 

either secondary vegetation or planted nurse trees 

(Barnard 1954). Paraserianthes falcataria, Peltophorum 

spp. and Adenanthera spp. were found to be useful as 

nurse trees although the latter two species, which have 

smaller and lighter crowns, were better suited. 

Dryobalanops aromatica was established under a shelter 

of Fragraea fragrans (Landon 1948b) and on lines in 

secondary vegetation (Barnard 1949a). Doan (1985) 

reported a planting trial in Vietnam, where Dipterocarpus 

alatus, Hopea odorata and Anisoptera costata were 

planted under shade of Indigofera teysmanii and Acacia 

auriculiformis. Of the three species Dipterocarpus alatus 

was more light demanding. Miyazaki (1989) found that 

age of the nurse crop had an effect on survival of 

Anisoptera thurifera. Seedlings were planted under 8-10 

year old and 2-3 year old Acacia auriculiformis. Mortality 

was higher for those seedlings planted beneath the 

younger nurse crop. In a sowing experiment by Tomboc 

and Basada (1978) seeds of Shorea contorta were sown: 

under a secondary forest canopy which allowed the sun 

to filter through the canopy for 1 hour daily, and in the 

open. Survival was significantly higher under the forest 

canopy, while height growth and leaf development were 

better in the open. Wyatt-Smith (1947) reported a 

successful sowing experiment with Dryobalanops 

aromatica under a 1½-2 year old secondary forest while 

sowing in cut lines proved a failure. The ecological role 

of pioneer species in the natural regeneration of loggedover 

dipterocarp forests is discussed. Wyatt-Smith (1947) 

suggested that secondary vegetation can be cheaply 

converted by line planting beneath its canopy in 5 to 10 

years’ time (depending on the amount of soil degradation 

that has taken place), when most of the herbs and ground 

flora will have been shaded out. Rosario (1982) proposes 

silvicultural treatments that preserve pioneer species. 

These proposals are similarly valid for the treatment of 

secondary vegetation into which dipterocarps are planted. 

Other researchers have tested specific forms of site 

preparation. Maun (1981) reported a sowing experiment 

in a dipterocarp forest, where germination, survival and 

early growth of Shorea contorta was tested. The 

treatments were five different types of cover: (1) bare 

soil, (2) soil with litter, (3) soil with litter and ground 

cover, (4) soil with litter and underbrush, and (5) soil 

with intact vegetation cover. Germination was best in 

treatment (4), survival in treatment (4) and (5) and growth 

performance was better in treatment (1) and (2). Ang 

(1991) tested survival and growth of Shorea parvifolia 

on three sites: (1) secondary forest with trees of >20 cm 

girdled well in advance of planting, (2) open site (large 

opening in forest) with 30 cm top soil removed, and (3) 

open site (large opening in forest) with top soil retained. 

Survival was similar in all three sites, but growth was 

best in the open site where top soil had been retained. 

Barnard (1949b) investigated the effect of two types of 

planting site preparation on survival and growth of 

differently prepared seedlings of Dryobalanops 

aromatica. The test site was a natural forest with invasion 

of Gleichenia spp. and Eugeissona triste. Part of the test 

site was clear-felled and burnt. A control area remained 

unburnt, where Gleichenia spp. and Eugeissona triste 

were cut only. In the unburnt site all planting stock types 

established, while in the burnt site only the potted 

seedlings succeeded. Rowntree (1940) proposed grazing 

as a means to control the growth of Imperata cylindrica 

to secure the establishment of S. robusta regeneration. 

Nykvist et al. (1994) have reported the impact of forest 

harvesting and replanting on the forest site. They conclude 

that burning should be avoided in order to reduce nutrient 

loss and ensure better plantation growth. A similar view 

was already voiced by Wyatt-Smith (1949a) for the same 

reason. In the trial described by Barnard (1949b) in the 

plots prepared by burning only potted seedlings of 

Dryobalanops aromatica succeeded. As saplings they 

developed strong stems and had good height growth.


Qureshi et al. (1968) investigated the effect of soil 

working and weeding on the growth and establishment 

of Shorea robusta plantations. 

A common practice is to establish dipterocarp 

plantations by line planting into forest vegetation. Ådjers 

et al. (1995) have investigated the effect of line width, 

direction and maintenance on survival and performance 

of Shorea johorensis, S. leprosula and S. parvifolia. Line 

direction had little effect on survival or growth, although 

SE-NW line direction was best for S. johorensis. Line 

width did not affect survival, but effect on growth was 

significant. Line widths used were 1, 2 and 3 m. In the 

control, the seedlings were planted under the forest canopy 

without opening it above the planting line. Horizontal 

line maintenance was better than vertical line maintenance 

and growth of S. johorensis and S. parvifolia benefitted 

from it. Survival was not affected. Omon (1986) tested 

the strip width to be cut into secondary forest for optimal 

growth of planted Shorea ovalis seedlings. He found that 

strips 1 m wide were the best for survival and 

performance. 

Planting patterns in the context of underplanting were 

discussed by Tang and Chew (1980). Shorea parvifolia 

was underplanted in two patterns: (i) line planting, and 

(ii) group planting in groups of 4-6 trees at final spacing. 

Six months later the tree crowns overshadowing the 

planting lines or the planting patches were removed. 

Differences in growth were not significant, however, 

survival was higher for the group planting. The authors 

recommend removal of overhead shade after 6 months 

and underplanting as group planting. Abalus et al. (1991) 

recommend groups planted at a spacing of 10 x 10 m. 

In an underplanting trial at Agumbe in Karnataka, 

India, Vateria indica seedlings were planted in 1962 and 

observed until 1978. Those growing under lateral shade 

with sufficient light had grown to an average height of 

over 5 m in 16 years while those which had no light 

reaching them had survived but had grown only about 

5 cm (Rai, personal communication). 

Planting Techniques 

Outside India, Indonesia and Malaysia no large-scale 

plantations of dipterocarp species exist. Although 

experimental forests have been established in several 

regions information on the establishment techniques is 

scarce. The most complete account of artificial 

regeneration in the Malaysian context is by Barnard 

(1956). Agpaoa et al. (1976) give an overview of the 

planting techniques in the Philippines context. Most of 

the information on planting techniques is contained in 

instructions of forest services or of companies, and thus 

not always readily available. Planting techniques have 

been worked out very well for tropical conditions and 

the basics are generally valid irrespective of region, 

species or site. 

Planting methods can be classified into: (1) planting 

of potted seedlings or transplants, (2) planting of bareroot 

seedlings or transplants, and (3) planting of stumps. 

Normally, dipterocarps are planted as potted seedlings, 

when they are about 9 months old and about 25-30 cm 

tall. Size or age of planting stock has been investigated 

by various researchers. In general, potted seedlings had 

better survival (e.g., Barnard 1954, Cerna and Abarquez 

1959). The planting holes are usually prepared to a depth 

of 25 cm. The seedling or transplant is removed from the 

container (polythene bag) with the earthball undamaged. 

If broken, the beneficial effect of planting seedlings or 

transplants with undamaged roots is lost. Different pot 

types were used in the past, e.g., bamboo pots, veneer 

pots, tin cans. However, a plastic bag of 500 cc content 

(e.g., 10 cm x 15 cm and 6.3 cm diameter) is the bag size 

commonly used. Barnard (1954) tested different sizes of 

bamboo pots and larger pots gave better survival. A trial 

on varying pot sizes using Shorea polysperma was carried 

out by Bruzon and Serna (1980) and height development 

in 8 cm diameter pots was best. When planting, the upper 

part of the earthball should be slightly below the soil 

surface for successful establishment, and never above it. 

Depth of planting was investigated, e.g., by Shrubshall 

(1940), and Walton (1938) who for Dryobalanops 

oblongifolia found deep planting (collar 5 cm below 

surface) gave the best results and shallow planting caused 

75% mortality. Shrubshall (1940) also reported deep 

planting gave the best results. Earth is firmly placed 

around the plant to close the air spaces and finally, the 

young plants are mulched with organic material to 

prevent desiccation and overheating of the soil. Bareroot 

seedlings can be planted in two ways: hole-planted 

as in potted plants; and notch-planted. In notch planting a 

cone- or wedge-shape hole is made with a spade or a 

hoe. The roots of the plant are placed into the hole to the 

required depth and the soil firmed around the plant. 

Barnard and Setten (1955) reported on the comparison 

of planting trials of Dryobalanops oblongifolia in 

prepared planting patches and in notches. The 

performance of two types of seedlings were compared:


entire seedlings lifted from the soil and stripped 

seedlings where leaves were reduced to about one third 

their length. The percentage of trees surviving after one 

year was highest for entire seedlings planted in cultivated 

patches (58%). The lowest survival was found for stripped 

seedlings planted in notches (19.8%). Bare-root stock 

requires some moisture-preserving techniques to keep 

roots moist during transport and storage prior to planting 

(e.g., Strong 1939, Rayos 1940). A detailed description 

of the planting technique for bare-root stock is given by 

Agpaoa et al. (1976). Sometimes the planting stock is 

root and/or shoot-pruned or stripped partially or totally 

of leaves to initially reduce transpiration to facilitate 

establishment. Root pruning was generally beneficial 

(e.g., Walton 1938, Sasaki 1980a). Stripped seedlings 

of Shorea talura could be stored for several months 

without losing vigour and capacity for cutting propagation 

(Sasaki 1980a). Landon (1948b) found that stripping 

leaves of Dryobalanops aromatica was unsuccessful. 

Wildings are either lifted with a ball of earth or are forestpulled. 

They can be either directly planted or they are 

kept in a temporary nursery under light shade for 3 to 6 

months to recover before they are planted. Normal 

practice is to keep wildings for some months in a nursery 

until they have recovered. Very low survival rates were 

achieved by Lantion (1938) with forest-pulled wildings 

that were planted into the forest without a recovery 

period. Wildings of Dipterocarpus grandiflorus and 

Shorea teysmanniana were pulled from the forest, 

stored for three days (partly mud-puddled and partly not) 

and then planted. The average survival for mud-puddled 

wildings was 9.5% and for wildings not mud-puddled 

2.9%. Palmiotto (1993) described a direct transplanting 

trial in the understorey and a gap using wildings of Shorea 

hopeifolia, S. johorensis, S. leprosula, S. parvifolia, 

S. parvistipulata and S. pinanga. Transplanting appeared 

to have a negative effect on survival. Survival in the 

understorey was between 8 and 58% and in the gap 

between 3 and 50%. Recovery in the nursery is important, 

if a high survival percentage after transplanting into the 

field is to be achieved (e.g., Capellan 1961, Moura-Costa 

1995). 

There are clear indicators of the need to fertilise 

initially, e.g., (1) sites where deficiency symptoms occur, 

(2) sites with top soil removed, (3) sites carrying 

vegetation indicating poor soil conditions, and (4) sites 

with strong weed competition. It is, at present, still too 

early to formulate valid fertiliser regimes. Less certain 

are the fertilising requirements of the newly planted 

seedlings. Nutrient deficiencies will occur, especially, 

in plantation establishment on areas that have suffered 

degradation to some extent (e.g., clear-felled areas and 

secondary forest). Moura-Costa (1993) reported 

fertilisation in the context of large-scale enrichment 

plantings with rock phosphate (100 g) applied to the 

planting hole. On an experimental scale, the effect of 

additional fertiliser application on the establishment of 

dipterocarps is being studied. Yap and Moura-Costa 

(1994) reported on the effect of nitrogen fertilisation, 

soil texture and other factors on biomass production of 

Dryobalanops lanceolata seedlings. Nussbaum et al. 

(1995) reported a combined experiment of soil-working 

and fertilisation of tree seedlings of Dryobalanops 

lanceolata and Shorea leprosula. The treatments were: 

(1) planting into compacted soil; (2) planting into 

compacted soil + fertilisation (100 g of rock phosphate 

placed in the planting hole and 40 g of granular 12:12:17 

N:P:K + micronutrients placed in a ring of about 10 cm 

from the seedling just below the soil surface); (3) 

planting into compacted soil + mulching (pieces of bark 

which had been stripped from felled trees 1 year earlier 

were used to cover the plot); (4) planting of seedlings 

into cultivated plots (soil in the whole plot turned over 

and broken up to a depth of 30 cm 2 to 3 weeks before 

planting); (5) planting into cultivated plots + fertilisation; 

(6) planting into cultivated plots + mulching; and (7) 

planting into planting holes with soil replaced with topsoil 

from undisturbed forests. After 6 months of observation 

best diameter growth was found in treatments (2), (5) 

and (7). Crown diameter was also largest in these three 

treatments. Seedlings responded strongly to fertiliser 

application, while (with exception of soil replacement) 

response to soil working (plot cultivation or mulching) 

was less distinct. 

Sowing 

Although not a widely used technique for establishment 

of even aged stands, sowing has been tried in the past. It 

has been applied on an operational scale in India (e.g., 

Chakravarti 1948) and Pakistan (e.g., Amam 1970). Gill 

(1970) found sowing of Shorea leprosula promising in 

the context of enrichment operations. Some of the finest 

Dryobalanops aromatica stands in Malaysia were 

established by broadcast sowing into high forest (Watson 

1935). The results of direct sowing trials are not 

conclusive. Shaded, cool and moist microsites seem to 

be essential for successful germination and survival.


Tomboc and Basada (1978) tested the performance of 

Shorea contorta sown on open areas and under secondary 

growth canopy. Survival was highest under the cover of 

the forest, while growth was better in the open. Maun 

(1981) suggests that it will be necessary to germinate 

direct-sown seeds and grow the seedlings of S. contorta 

initially in shaded conditions. Later, the vegetation should 

be opened for better growth of the seedlings. Similarly, 

Strong (1939) found in a trial of direct sowing into 

cultivated areas (taungya) and into high forest that the 

germination of Dryobalanops oblongifolia and Shorea 

sumatrana failed in the cultivated areas largely as a 

result of drought and heat. The seeds were also attacked 

by insects and rodents. Sowing under the shelter of 

Paraserianthes falcataria was successful with Shorea 

stenoptera (Sudiono and Ardikusumah 1967). 

Chakravarti (1948) found direct sowing is the only 

method to artificially regenerate Shorea robusta forests 

in India. The principal adverse factor to germination and 

survival of seeds is drought and shade is essential for 

successful regeneration. Suggestions on the best type 

of nurse crop are given. Sown seeds may be attacked by 

insects or rodents. Barnard and Wyatt-Smith (1949) 

reported high mortality in their sowing trial of 

Dryobalanops aromatica in secondary vegetation 

mainly caused by rodent attacks on the germinating seeds. 

In comparison to other methods of artificial 

regeneration, the sowing method is less convincing. 

Cerna and Abarquez (1959) compared growth and survival 

of S. contorta plants that originated from transplants and 

from direct-sown seeds 11 years after stand 

establishment. Heavy mortality of seedlings resulted 

from direct sowing. S. contorta is very sensitive to bareroot 

planting and planting of balled plants was the most 

successful method. 

Stand establishment by sowing is a very wasteful 

practice because of the large amount of seeds needed 

for sowing operations, 

Stand Tending 

‘Tending, generally, is any operation carried out for the 

benefit of a forest or an individual thereof, at any stage 

of its life. It covers operations both on the crop itself, 

e.g., thinnings and improvement cuttings, and on 

competing vegetation, e.g., weeding, cleaning, climber 

cutting, and girdling of unwanted growth, but not 

regeneration cuttings or site preparation’ (Ford- 

Robertson 1983). Stands develop and grow through 

various developmental stages from seedling or coppice, 

through thicket, sapling, and pole, to the tree stage, i.e. 

to maturity, and finally to overmaturity, but sometimes 

ending in residual standards. Residual standards are trees 

that remain standing after the rest of the stand has been 

removed or has died. 

Weeding and Cleaning 

The immediate post-planting care (mainly weeding), 

which covers the time until the plantation can be 

considered established, is crucial for planting success. 

Weeding is an operation whereby mainly herbaceous 

vegetation is eliminated or suppressed during the seedling 

stage of the forest crop. It is, therefore, the first cleaning 

and aims to reduce competition within the seedling stand. 

Cleanings to eliminate or suppress undesirable vegetation 

(mainly woody including climbers) are carried out when 

the young plant is in the sapling stage (1.5 m height and 

5 cm diameter). Cleanings are carried out during the 

thicket stage of a forest crop and therefore before, or at 

latest with, the first thinning, so that better trees are 

favoured. Removal of overtopping vegetation must be 

carried out during weeding and clearing operations in 

dipterocarp plantations established either under a nurse 

crop (natural or planted) or in existing, line-planted, taller 

vegetation (e.g., secondary forest). Watson (1931/32e) 

classified trees according to their silvicultural importance. 

He, especially, distinguished between undesirable weeds 

which needed to be eradicated under nearly all 

circumstances and harmless tree species which are useful 

for shade or cover. Barnard (1954) recommended the 

removal of the overhead shade as soon as the young trees 

have recovered from the transplanting shock. He also 

found that the slightly increased light due to the cutting 

of planting lines was beneficial. Tang and Wadley (1976) 

discuss the technique of line opening and shade regulation. 

Techniques of line opening in the context of enrichment 

planting are described, e.g., by Chai (1975), Tang and 

Wadley (1976) and Lai (1976). 

A common practice is to mark planting places with 

small poles with the empty plastic bag pulled over the tip 

so that the location of the plant can be detected by the 

weeding crews. The weeding can be done for example, 

as strip or ring weeding. Normal practice is to blanket 

weed the planting lines and remove the weeds by slashing. 

However, woody vegetation grows more vigorously if 

cut, requiring additional weeding operations. Since the


young dipterocarp plants can withstand light shade it is 

not necessary to remove all non-crop vegetation. It would, 

therefore, be more appropriate to develop more selective 

procedures with less competitive weeds being left. 

Barnard (1954) gives the following general 

recommendations for weeding operations: 

• plants must be kept free of climbers; 

• freeing from climbers must be done before the plants 

have been overgrown; 

• uprooting of weeds is preferable to slashing to prevent 

vigorous regrowth; 

• grasses and young plants compete for moisture and 

nutrients and should be periodically removed by cleanweeding 

in a circle around the plant; and 

• weeding should not be done with a hoe, to avoid damage 

to the plants. 

More investigations are needed on selective weed 

control, including the development of risk categories for 

so-called weed trees and methods of suppression or 

elimination. Useful descriptions of weed vegetation in 

the Malaysian context are found in the rubber planter’s 

manual (Haines 1940). Such a manual became necessary, 

when the so-called ‘forestry’ cultivation was introduced 

in rubber plantation management. The basic idea was to 

retain an undergrowth of non-competitive vegetation so 

as to prevent erosion and maintain favourable soil 

chemical and physical properties. Naturally, only 

harmless weeds could be allowed to grow in the 

plantations. This made it necessary to categorise the 

vegetation according to noxiousness and to define the 

treatments required. Weeds particularly noxious to 

young plants have been noted, e.g., Wyatt-Smith (1949b), 

Seth and Dabral (1961), Palit (1981). Wyatt-Smith 

(1963b) listed ‘weed’ trees that had to be poisoned 

irrespective of whether competing with ‘economic’ 

species or not. The control of specific types of weeds 

has been described, e.g., Strugnell (1934), Mitchell 

(1959) for Imperata cylindrica, Kelavkar (1968) for 

Lantana camara, and Palit (1981) and Bogidarmanti 

(1989) for Mikania spp. Liew (1973) tested methods 

to eradicate climbing bamboo (Dirochloa spp.) in Sabah 

and was successful with merely cutting the bamboo near 

the soil surface. Chemical weeding was tested by Palit 

(1981) in Shorea robusta plantations against Mikania 

scandens. Seth and Dabral (1961) tested the efficiency 

of 5 herbicides based on 2,4-D or 2,4,5-T in moist 

deciduous Shorea robusta forests against trees and 

coppice of Mallotus philippinensis, Ehretia laevis, 

Ougeinia oojeinensis, Miliusa velutina, Buchanania 

lanzan, Aegle marmelos. M. philippinensis proved to 

be resistant. Chong (1970) carried out a trial on chemical 

control of the stemless palm Eugeissona triste in Shorea 

curtisii forests. In regions with distinct seasonality, 

timing of the weeding operations is important. Bhatnagar 

(1959) related the timing of the weeding operations to 

the annual height increment peaks of Shorea robusta 

seedlings. He recommended carrying out weedings 

during or somewhat in advance of these periods, so as to 

help to relieve the intense competition between the 

Shorea robusta seedlings and the weeds. In Shorea 

robusta forests the so-called rain-weeding is carried out, 

i.e., weeding during the rainy season (e.g., Rowntree 

1940, Sarkar 1941). For good growth of the young 

planted dipterocarps a good exposure to light is essential. 

In line plantings (including enrichment planting) 

overhead shade must be continuously absent from the 

planting lines. Agpaoa et al. (1976) give a comprehensive 

description of the procedure of enrichment planting and 

the corresponding tending operations. In underplanting 

under a nurse crop the overhead shade must be removed 

within a few years (e.g., Sanger-Davies 1931/1932, 

Ardikoesoema and Noerkamal 1955, Wyatt-Smith 

1963b Agpaoa et al. 1976). Small undesirable trees (up 

to about 5 cm diameter) can easily be removed with a 

bush knife or axe. Larger trees, however, are frequently 

girdled or poison-girdled using arboricides. Arboricide 

use is described e.g., Sanger-Davies (1919), Barnard 

(1950, 1952), Beveridge (1957), Nicholson (1958), 

Roonwal et al. (1960), Wyatt-Smith (1960, 1961a, 

1963c), Wong (1966), Liew (1971), Agpaoa et al. 

(1976), Chai (1978), Chew (1982) and Manokaran et 

al. (1989). Well known arboricides are 2,4,5-T, Garlon 

4E, Tordon 22K, Velpar-L and sodium arsenite. Most 

of the tests were done with 2,4,5-T and sodium arsenite. 

Thinnings 

Thinning is ‘a felling made in an immature crop or stand 

in order primarily to accelerate diameter increment but 

also, by suitable selection, to improve the average form 

of the trees that remain, without - at least according to 

classical concepts - permanently breaking the canopy’ 

(Ford-Robertson 1983). A thinning regime is 

characterised by type, grade or weight and frequency. The 

type of thinning can be a thinning from above, where 

particularly the most promising, not necessarily the 

dominant, stems are favoured and where those trees, from 

any canopy class that interfere with the promising ones,


are removed. Another type of thinning is the thinning from 

below, where particularly the dominants or selected 

dominants are favoured and a varying proportion of other 

trees is removed. Grade of thinning is a degree of thinning 

based on dominance, crown and stem classes, and the 

extent to which these classes are removed at any one 

thinning. 

With the exception of Shorea robusta no thinning 

regimes have been developed for dipterocarp plantations. 

Krishnaswamy (1953) and Mathauda (1953a) studied the 

effect of thinning intensities on height and diameter 

development, stand basal area and volume increment of 

Shorea robusta. The conclusions from this thinning trial 

were: that the thinnings should be carried out every 5 

years up to an age of 20 years and thereafter at larger 

intervals; and the maximum volume production is 

obtained under C/D-grade (heavy to very heavy low 

thinning as per standard definition of the terms adopted 

in India). In the C/D grade the dead, moribund, diseased 

trees, whips of co-dominant and dominant trees, defective 

co-dominant and dominant trees and a small proportion 

of sound co-dominant and dominant trees are removed. 

Thinning according to the C/D grade was found to be 

best for the production of both fuelwood and timber. 

Wyatt-Smith (1963a) assumed that in dipterocarp 

plantations a thinning cycle of 5 to 10 years would be 

adequate. Suri (1975a) developed a quantitive thinning 

model for Shorea robusta forests in Madya Pradesh, India. 

Based on the correlation between crown diameter and 

stem diameter a thinning model was formulated and stem 

density regimes for different crown disengagement levels 

determined. It was concluded that quantitative thinning 

grades can be developed for different species by studying 

their crown diameter/bole diameter relationship. The 

crown disengagement in younger stands was sometimes 

carried out as so-called stick thinning, i.e. starting from a 

selected crop tree any tree growing within a defined 

distance (e.g., six, nine or twelve feet) from the selected 

crop tree was removed for example, in a naturally 

regenerated, more or less even-aged stand of 

Dryonbalanops aromatica (Anon. 1948b). An important 

conclusion from this trial is that it is not advisable to make 

heavy thinnings before the overwood has been removed, 

since the young crop can be overtaken by climbers and 

secondary species benefitting from increased light. The 

heavily thinned treatments suffered severely from 

climbers and weed species, while trees damaged by the 

falling overwood had no neighbours to replace them. 

Thinning is usually done with a bush knife (smaller 

trees), an axe or a saw but if the tree is not to be utilised, 

girdling or poison-girdling may be applied. Often girdling 

alone is unsuccessful and poison-girdling is recommended 

(e.g., Wyatt-Smith 1963b, c, Agpaoa et al. 1976). The 

trees to be removed are frill-girdled and the poison is 

applied into the frill. Effective chemicals have already 

been mentioned in the section on weeding and cleaning. 

Thinning interventions require some kind of 

classification of the stems in the stand to be thinned. 

Krishnaswamy (1953) presented a detailed stem 

classification which is based on dominance position and 

within each position on vigour, soundness, crown 

development and other characteristics. It resembles the 

classification of Kraft (1884), but includes reproduction 

or regeneration and overmature trees (e.g., standards). 

Any thinning, except for schematic interventions, requires 

that all trees in the stand are judged according to their 

function. Potential final crop trees (PCT) are distinguished 

from non-crop trees (NCT). The PCT are those trees 

which owing to their straightness and evenly formed 

crowns are to be retained as crop trees and released from 

competition. NCT may have different functions. There 

are harmful trees that damage the crowns or stems of the 

PCT and should be removed. There are useful NCT which 

enhance growth form and branch-shedding of the PCT 

or have important ecological functions. There are 

individuals for which their future development and 

function is not clear and they have to be spared from 

thinning until the necessity for removal is beyond doubt. 

In the Malaysian context Watson (1931/1932e) has 

classified the most common trees in Peninsular Malaysia. 

He classified the species into the following categories: 

• quality timber trees, 

• utility timber trees, 

• subsidiary trees, 

• insignificant trees (fillers only), 

• poles, 

• cover or nurse trees, which are harmless species, and 

• weed trees, which are undesirable. 

This classification was made for natural forests and 

is not really applicable for plantations. 

Although there is no experience available on the 

tending and thinning of dipterocarp plantations outside 

India, some inferences can be made from tending and 

thinning experiments and from observations in naturally 

regenerated dipterocarp forests, which lead to more or 

less even-aged and fairly regular stands. Such stands may


have resulted from, e.g., Regeneration Improvement 

Systems or from Uniform Shelterwood Systems, as they 

were, for example, applied in Malaysia. Wyatt-Smith 

(1963b) gives a thorough review of the thinning 

experience up to that time. His recommendations for 

thinning more or less regular crops were: 

• removal of climbers of above 2.5 cm diameter, although 

the limit can be lower if smaller climbers prove 

to be damaging the crop trees, 

• removal of all weed trees; also those that are going to 

overtop the PCT until the the next intervention, 

• removal of all malformed stems of commercial species 

provided a stem of better form is adjacent, 

• removal of all wolf trees, 

• removal of co-dominants of inferior timber value, 

• selective thinning of co-dominants of equivalent 

silvicultural and timber value that compete strongly, 

and 

• thinning to a maximum basal area of about 1/2 to 3/4 

of the expected carrying capacity of the site. 

In the context of regeneration operations within the 

Regeneration Improvement Systems Durant (1940) was 

confronted with the criticism that opening the canopy 

would lead to luxuriant ‘secondary growth’ (what we 

would call today secondary forest) consisting mainly of 

Randia scortechenii, Pasania sp., Barringtonia sp., 

Girroniera nervosa, Trema ambionensis, Macaranga 

spp., Endospermum malaccense and various fast-growing 

trees of other families. It was feared that the young 

dipterocarps might be suppressed by these species and 

frequent and expensive cleanings needed. Three 

experimental plots were set up. Two plots were 

established in stands where the canopy over young 

regeneration had been removed by regeneration 

improvement fellings and one plot was laid out in an area 

where the canopy over young regeneration had almost 

completely been removed by a heavy storm. The 

treatments in the first plot were: (i) untouched control, 

(ii) cleaning (cutting back all growth other than Shorea 

spp.), and (iii) cleaning and respacing (‘thinning’ of the 

Shorea spp. to an average distance of 1.83 m leaf to leaf). 

The treatments in the second plot were: (i) untouched 

control, (ii) cleaning (cutting back everything except 

saplings of the desirable species), and (iii) mainly climber 

cutting with minimal cutting of undergrowth. In the third 

plot only a cleaning in favour of saplings and small poles 

was carried out. The objective of the first two plots was 

to investigate the effect of the secondary forest vegetation 

on survival and diameter growth of sapling-size natural 

regeneration of Shorea spp. The third plot tested whether 

larger regeneration (large saplings, small poles) was out 

of danger from its competitors. After establishment, the 

plots were left unattended for four years and then 

enumerated again. 

From Durant’s experiment, inferences were made 

concerning the regeneration of S. leprosula: 

• However severe the opening of the canopy, provided 

adequate seedling regeneration is present, S. leprosula 

can tolerate competition with other vegetation up to 

the sixth year. 

• Cleaning and thinning after the second year will secure 

an even distribution of stocking and will increase 

the growth rates. Complete omission of tending up to 

the sixth year is not fatal (which is in agreement with 

other authors e.g., Walton 1933, 1936a, Wyatt-Smith 

1949b, 1958, 1963b). 

• Serious competition from secondary forest species 

is probably due to a comparatively few species, and, if 

these can only be recognised and eliminated, a considerable 

reduction of cleaning costs should be possible. 

(The species recognised as responsible for suppression 

were Endospermum malaccense, 

Elaeocarpus stipularis, Macaranga spp., Paropsia 

varediformis and Quercus lucida). 

• With sufficient initial opening of the canopy, good 

stocking of Shorea leprosula can be expected to survive 

up to the 14th year. At this stage the crop reaches 

pole size, and adequate assistance can be given very 

cheaply by the poison-girdling of competitors around 

individual trees. 

The conclusions are important for the tending of 

young naturally regenerated and more or less even aged 

stands originating either from natural stands or from 

plantation stands under the Shelterwood System. The 

findings of Durant (1940) can, however, not be applied 

without some restrictions to young plantations of 

dipterocarps. The initial number of stems in plantations 

is usually so low that omission of early tendings 

(weedings, cleanings) will probably entail high losses 

endangering stand establishment. 

Strugnell (1936b) tried three treatments (only 

dominant trees retained; dominant and dominated trees 

retained; dominant, dominated and suppressed trees 

retained) in a young natural pole stand of Shorea leprosula 

and S. parvifolia. He found that the basal area of the 50 

largest trees/acre was highest for the medium


intervention. Sanger-Davies (1937) carried the ideas 

further and formulated a guide for the tending of more 

or less even aged stands of S. leprosula. In his technical 

recommendations, he proposed starting tending while the 

shelterwood is still standing. 

When designing research it should be kept in mind 

that the beneficiary of the thinning operation is the crop 

tree and, therefore, indiscriminate elimination of noncrop 

vegetation is unnecessary. Non-crop trees have 

beneficial ecological functions. Mead (1937) discusses 

the formation of mixed stands of dipterocarps and shadebearing 

non-dipterocarp understorey species with dense 

crowns. The species Scorodocarpus borneensis, Mesua 

ferrea, Randia scortechinii, Randia anisophylla, 

Greenia jackii etc. were planted in mixture with Shorea 

leprosula, which forms a rather open crown, to prevent 

the invasion of light demanding pioneer vegetation which 

impede the establishment of natural dipterocarp 

regeneration. Tending has, therefore, to consider also 

the secondary vegetation. Any inconsiderate felling 

should be avoided and instead it should be asked, whether 

such vegetation could assist in keeping the forest floor 

conducive to natural regeneration. 

Re-establishment by Natural Regeneration 

Embarking on plantations with dipterocarp species which 

grow relatively slowly compared with fast-growing 

exotics needs strong economic backing. Recent 

economic calculations (Kollert et al. 1993, 1994) have 

shown that it only makes sense, if at the end of the first 

rotation the new stands are established by natural 

regeneration. It is in this context that some comments 

are given on regenerating naturally even-aged planted 

dipterocarp stands, although on an operational scale this 

will be only a problem of decades from now. Systematic 

assessment of the regeneration situation and initiation 

of natural regeneration procedures are urgently needed 

for all species identified for plantation programmes and 

for which stands near rotation age exist. This should 

include research on the harvesting techniques required 

to reduce negative impacts on stand regeneration. 

The natural regeneration of even-aged planted stands 

will most likely be carried out as some kind of 

shelterwood system. Shelterwood systems are ‘evenaged 

silvicultural systems, in which, in order to provide 

a source of seed and/or protection for regeneration, the 

old crop (the shelterwood) is removed in two or more 

successive shelterwood cuttings, the first of which is 

ordinarily the seed cutting (though it may be preceded 

by a preparatory cutting) and the last is the final cutting, 

any intervening cuttings being termed removal cuttings’ 

(Ford-Robertson 1983). Where there is adequate 

regeneration the old crop may be removed in a single 

cut (e.g., Malayan Uniform System). Preparatory felling 

means removing trees near the end of a rotation so as to 

open the canopy permanently and enlarge the crowns of 

seed bearers, with a view to improving conditions for 

seed production and natural regeneration. Here, no 

adequate regeneration is on the ground. Seeding felling 

is removing trees in a mature stand so as to effect 

permanent opening of its canopy (if there was no 

preparatory felling to do this) to provide suitable 

conditions for regeneration from the seed of trees that 

are retained. Removal felling is removing trees between 

the seed cutting and the final cutting, so as gradually to 

reduce the shelter and admit more light to aid the 

regeneration crop and to secure further recruitment. This 

type of felling is carried out over adequate regeneration. 

There is almost 80 years of experience with the 

regeneration of natural dipterocarp forests. Experience 

on individual aspects of natural regeneration gained is 

with modification applicable to even-aged stands of 

dipterocarps. This does not mean regeneration systems 

for even-aged stands can be derived from the knowledge 

available now but it is possible to outline some general 

directions. 

One important aspect of the establishment of a new 

generation by natural regeneration is, whether or not the 

stands will fruit well before the rotation has ended. A 

few observations have been made. Ng (1966) concluded 

from his work on age of first flowering of dipterocarps 

that many species begin to flower and bear good seed 

before their 30th year. Tang (1978) found three trees of 

Shorea leprosula planted in a taungya stand had fruited 

at the age of 7 years. Similar early ages of flowering/ 

fruiting were reported by Lee (1980) for Shorea pinanga 

(flowering 6 years after planting) and by Suziki and 

Gadrinab (1988/1989) for S. stenoptera (fruiting 6 years 

after planting). Ardikoesoema and Noerkamal (1955) 

described a S. leprosula stand in Java that had fruited 

aged 13 years producing a moderately dense seedling 

crop. Appanah and Weinland (1996) evaluated the field 

files of the dipterocarp plantations at the Forest Research 

Institute Malaysia and fruiting was reported for Shorea


leprosula, S. macrophylla and Dryobalanops aromatica 

stands at about 20 years age. Additionally, plantation 

stands of some other species (e.g., Dryobalanops 

oblongifolia, Shorea macroptera) have established 

regeneration. However, the exact stand age at first 

flowering has not been recorded. 

Little information is available concerning 

preparatory operations. Chong (1970) reported the effect 

of Eugeissona triste (a stemless palm) control on 

regeneration of Shorea curtisii. The experiments 

showed that a pre-felling treatment with a light girdling 

and Eugeissona triste control undertaken after a heavy 

seed fall prior to felling had a beneficial effect. The 

operation not only increased the vigour of the established 

regeneration but also created conditions on the forest 

floor conducive to recruitment of new individuals. Raich 

and Gong (1990) found that seed germination 

demonstrates clear patterns of shade tolerance or 

intolerance identical to those long recognised for tree 

seedlings. Among the species tested were Dipterocarpus 

grandiflorus, Shorea multiflora and Vatica nitens. They 

germinated in the understorey as well as in the gaps 

(typically 20-30 m in diameter) but failed to germinate 

in larger clearings. So, if preparatory canopy openings 

are prepared, these openings should not exceed normal 

gap size. 

Preparatory fellings have never played an important 

role. Treatment of seed trees in the natural forests to 

improve their crowns is unneccessary because being 

emergents they have already fully developed crowns. 

More information is available on the manipulation of the 

old crop over existing regeneration (regeneration fellings 

and final fellings). Although strictly applicable only to 

natural forest conditions, the basic findings should also 

be valid for plantations. Based on closely controlled 

experiments in the Wet Evergreen Forests of Sri Lanka, 

Holmes (1945) found that canopy conditions under 

seeding fellings most conducive to regeneration seem 

to be gaps of 20-30 m diameter evenly distributed and 

separated from one another by not more than one row of 

dominant trees. While raising the canopy gradually 

upwards, an ultimate canopy density of about 0.5 will be 

achieved. Zoysa and Ashton (1991) found that the 

germination of Shorea trapezifolia seeds planted on 

forest top soil with litter was little affected by partial 

shade or exposure to full sun. Watson (1931/1932c) 

discusses ‘preparatory’ fellings (strictly speaking they 

were regeneration fellings) for fostering natural 

regeneration within plantations. He states that seedlings 

of commercial species would establish better after 

opening the forest canopy, provided care is taken to 

prevent intrusion of weed species. He recommends 

removal of the lower forest canopy layers and cleaning 

of the undergrowth. But no fellings of this kind should 

be done in the absence of natural regeneration. Based on 

experiments of girdling understorey and upper storey 

trees, it was concluded that improvement systems should 

ensure adequate regeneration while retaining the canopy 

in such a condition that the lower storey is shaded 

preventing growth of competing vegetation (Walton 

1933, 1936a, b). Only after regeneration is abundant 

should any drastic opening of the canopy be undertaken. 

The vigorous response of seedling regeneration of 

Shorea spp. to full light indicates that treatment should 

aim at removing the canopy as rapidly and completely as 

is considered safe. The extent of canopy opening, 

however, should depend on the light demand/shade 

tolerance of the species. Strugnell (1936a) investigated 

the effect of suppression on young regeneration of 

Shorea leprosula, S. parvifolia and Neobalanocarpus 

heimii. Removal fellings should not be delayed for too 

long in light-demanding species as mortality will be high 

and growth responses weak. Shade tolerant species may, 

however, react vigorously even after a long time of 

suppression. In some species sudden exposure on canopy 

opening might lead to shoot borer attack as in 

Neobalanocarpus heimii (Durant 1939). Qureshi et al. 

(1968) emphasise that, before commencing tending 

operations on the regeneration, the canopy density has 

to be reduced to ensure sufficient light for the young 

plants. This was tested on natural regeneration of Shorea 

robusta under a planted parent stand. In mixed stands 

smaller gap sizes will favour shade tolerant species and 

larger gap sizes light demanding species. This is an 

important consideration if mixed stands of shade tolerant 

and light demanding species are to be regenerated (e.g., 

Raich and Gong 1990). 

The design of the regeneration system for 

dipterocarp plantations depends, apart from the 

production goal, on several other factors, e.g., the species 

involved, the stand condition, the regeneration behaviour 

and site factors. A uniform shelterwood system could, 

for example, be applied to Dryobalanops aromatica 

stands (Zuhaidi and Weinland 1993). They usually carry 

a fairly dense regeneration that is evenly distributed over 

the stand area. The canopy of the old crop is distinctly


mono-layered. The regeneration period will be rather 

short and the resulting stand after final felling will be 

fairly regular. Species which fruit more irregularly might 

require more irregular canopy openings following the 

recruitment patches and a group shelterwood system 

applied. The regeneration period will be protracted and 

the resulting stand more irregular. 

Reforestation and Afforestation of 

Degraded Land 

Reforestation is the re-establishment of a forest crop 

on forest land. Afforestation is the establishment of a 

crop on an area from which it has always or very long 

been absent. Degradation in the pedological sense is ‘any 

significant reduction in the fertility of the soil, whether 

in the course of its natural development or by direct or 

indirect human action’ (Ford-Robertson 1983). 

There is a growing need for rehabilitation of 

degraded forest sites following destructive logging, land 

clearing or mining. Dipterocarp species are by nature 

not very well suited for rehabilitation of severely 

degraded forest land. However, in some instances, 

dipterocarp species have been used with success (Ang 

and Muda 1989, Ang et al. 1992, Nussbaum et al. 1993, 

Nussbaum et al. 1995, Nussbaum and Ang 1996). Lately, 

Nussbaum and Ang (1996) have carried out a review on 

the rehabilitation of degraded land. Bieberstein et al. 

(1985) and Thai (1991) recommended Dipterocarpus 

spp. for the reforestation of devastated and shrub areas 

in Vietnam. Mitra (1967) describes the management 

measures carried out over large areas in West Bengal 

since the acquisition of all private forest lands (which 

were mainly Shorea robusta coppice forests) by the 

State in 1953. Shorea robusta was planted in eroded 

areas (Goswami 1957) and former bauxite mining land 

in India (Prasad 1988), Hopea parviflora on bare lateritic 

soil (Dhareshwar 1946) and Dryobalanops 

oblongifolia on waste land (Landon 1941). An initial 

burn and cultivation of planting patches were found to 

be beneficial. Shineng (1994) reports using dipterocarps, 

Dipterocarpus turbinatus and Parashorea chinensis, 

for establishing plantations on degraded forest land in 

tropical China but the growth rates of both dipterocarps 

were almost the lowest among 26 tree species tested. 

Mitchell (1963) explored the possibilities of afforesting 

raised sea beaches along the east coast of Peninsular 

Malaysia. Among the species tested was Hopea nutans 

which failed (Ang and Muda 1989). Rai (1990) describes 

a successful trial to restore degraded tropical rain forests 

of the Western Ghats (India) in which Vateria indica, 

Dipterocarpus indicus, Hopea parviflora and H. 

wightiana were used. 

Agroforestry 

Not many dipterocarp species have as yet been included 

in agroforestry systems. Shorea robusta is the only 

species which has been researched intensively in the 

context of the taungya system. Taungya is an ‘agrisilviculture 

system for the raising of a forest crop (a 

taungya plantation) in conjunction with a temporary 

agricultural crop’ (Ford-Robertson 1983). 

Nevertheless, agroforestry systems involving 

dipterocarps have been practised throughout the Indian- 

Southeast Asian region. Vateria indica and Shorea 

robusta have been used in agroforestry systems in India. 

Sal (Shorea robusta) taungya is a relatively well 

developed system in India (Huq 1945, Osmaston 1945, 

Kanjilai 1945, and others). Prominent in agroforestry 

systems in Borneo are the dipterorcarp species that 

produce edible nuts (Shorea spp. of the pinanga group) 

(Seibert 1989). An agroforestry system in East 

Kalimantan which often involves Shorea macrophylla 

is called the ‘lembo’ system (Sardjono 1990). Resin 

tapping of Shorea javanica is well developed in Sumatra 

(Torquebiau 1984). Integration of farming into the 

tending and conservation of logged forests was discussed 

(Serrano 1987, Mauricio 1987b), as well as the propects 

for agroforestry to be used for the rehabilitation of 

degraded forest land in Indonesia (Kartiwinata and 

Satjapradja 1983). Watanabe et al. (1988) investigated a 

taungya reforestation method in the context of the 

Government Forest Village Programme in Thailand, 

where Dipterocarpus alatus is involved. An agroforestry 

system using dipterocarps was also tried in West 

Malaysia (Cheah 1971, Ramli and Ong 1972) but it has 

not been adopted. These are but a few examples of 

dipterocarp species used in agroforestry systems. 

Forest Protection Aspects 

The knowledge on pests and diseases of dipterocarps is 

scanty, but a more systematic account is given in 

Chapter 7. Insects attack dipterocarp fruit crops heavily 

(Daljeet-Singh 1974). By comparison, their seedlings 

are well protected (Daljeet-Singh 1975). Becker (1981) 

investigated potential physical and chemical defences of 

Shorea seedling leaves against insects. Diseases include


fungal problems, bacterial and viral infections (Smits et 

al. 1991). Heart-rot of dipterocarps has been 

investigated, and is more serious in slow-growing than 

in fast-growing species (Hodgson 1937b, Bakshi et al. 

1963). Stand management strategies to control heartrot 

have been developed for Shorea robusta (Bakshi 

1957). 

With establishment of large-scale plantations of 

dipterocarps, susceptibility to diseases and pests is bound 

to increase. The control of pests and diseases in 

nurseries is well developed and advanced. Chemical 

control is the prevailing method to fight the attack of 

biotic agents. Chemical control is, however, not 

practicable after field planting. Prevention has to be 

secured by silvicultural means, e.g., species mixtures, 

structural diversity, avoidance of damage to trees and soil, 

etc. At present, there is an urgent need to survey diseases, 

defects and damages in existing dipterocarp plantations, 

particularly the incidence and possible causes of heartrot. 

Gaps in natural forests and plantations are created 

by natural mortality, biotic and abiotic agents. Lightning 

is a major cause for the occurrence of gaps not only in 

natural forests (Brünig 1964, 1973) but also in 

plantations. While in natural forests such gaps drive the 

regeneration dynamics, in plantations such gaps, first of 

all, reduce the stocking. The effect of lightning can be 

seen clearly in the plantation area of the Forest Research 

Institute Malaysia (personal observation). 

Management Aspects 

The available information, here, can be combined under 

the following categories: silvicultural systems, biological 

production, (especially growth), thinning schedules, 

stocking aspects, silvicultural diagnostics, and 

economics. 

‘A silvicultural system is a process, following 

accepted silvicultural principles, whereby the crops 

constituting forests are tended, harvested and replaced, 

resulting in the production of crops of distinctive form. 

The systems are conveniently classified according to the 

method of carrying out the fellings that remove the 

mature crop with a view to regeneration and according 

to the crop produced hereby’ (Ford-Robertson 1983). 

Silvicultural systems are discussed in the context of stand 

regeneration of plantations after the first rotation. 

In India, various silvicultural systems have been 

applied to encompass the wide ecological variation in 

the occurrence of dipterocarps. A selection system has 

been applied in seasonal rain forests and moist deciduous 

forests. Clearfelling and artificial regeneration have been 

carried out in moist deciduous forests where frost is 

absent. Various forms of shelterwood systems were 

applied in regions where frost was experienced. Coppice 

with standards was applied to Shorea robusta in dry areas 

and simple coppice systems used in wood lots in 

Karnataka. 

While the silvicultural systems for Shorea robusta 

forests in India are clearly formulated and understood, 

there is little information in other parts of the region on 

what the silvicultural system for dipterocarp plantations 

should be. Most silviculturists would like to re-establish 

an existing dipterocarp plantation at the end of the first 

rotation by natural regeneration. In Malaya, Walton 

(1933, 1936a), Watson (1935) and others have indicated 

the possible species among the fast-growing lighthardwoods 

which can regenerate naturally in the rotation 

envisaged for plantations. Little is known about the 

capacity of plantation-grown dipterocarp species to 

regenerate naturally at a rotation of about 50 years. There 

is already some information derived from planted 

species, e.g., Shorea leprosula, Dryobalanops spp., 

Shorea spp. of the pinanga group and S. robusta, which 

can be naturally regenerated during such a rotation time. 

All the existing, mature, experimental, dipterocarp 

plantations in the region should be assessed for natural 

regeneration. 

The growth of dipterocarps under natural forest 

conditions has been observed early this century (e.g., 

Edwards and Mead 1930, Watson 1931/1932a, Rai 1996). 

The observation of the growth of dipterocarps in plantations 

commenced only later. Analysis of 29 dipterocarp species 

in trial plots at the Forest Research Institute Malaysia 

indicates rotation ages of 40 to 50 years for the best 

performing species (Ng and Tang 1974). Individual volume 

and growth plots have also been analysed (Vincent 1961 a, b, 

c, d, Zuhaidi et al. 1994). The growth curves show a very fast 

early height and diameter growth with distinct differences 

between species in growth rates. Relatively impressive 

growth rates of the light red meranti group (Shorea spp.) 

have also been recorded in the Haurbentes experimental 

plantation stands in Indonesia (Masano et al. 1987, see also 

Ardikoesoema and Noerkamal 1955): a stand of Shorea 

leprosula achieved an average height and diameter of 44.6 

m and 77 cm respectively in 35 years. Shorea stenoptera 

was similarly fast growing with an average height and


diameter of 46.3 m and 75 cm respectively in 31 years. At 

the age of 29 years, Shorea platyclados stands in Pasir 

Hantap Experimental Forest in Indonesia have an average 

height of 29 m, bole length of 17 m and an average diameter 

of 41 cm. In Sarawak, trees of Shorea species of the pinanga 

group reached diameters between 34 cm and 75 cm between 

the age of 34 and 48 years (Primack et al. 1989). Shorea 

macrophylla showed the best growth performance, and S. 

splendida the poorest. A severe depression in growth 

occurred during flowering years. There is however, the 

danger in assuming the same growth rates in operational 

plantations and over the whole range of sites. Care has to be 

taken in economic calculations not to overestimate the 

performance. 

Shorea robusta is the most intensively researched 

species concerning growth and yield. Some yield tables 

exist (e.g., Howard 1925, Griffith and Bakshi Sant Ram 

1943). The species’ growth rate under different 

treatments has been reported by Mathauda (1953a). The 

growth rates of other dipterocarp species were reported 

by Mathauda (1953b) and Rai (1979, 1981a, b, 1989). 

More recently, the long-term research sites have been 

reviewed and updated by Rai (1996). Under natural 

conditions the annual rate of diameter growth for most 

dipterocarp species is only 0.3 to 0.35 cm. 

Among the dipterocarps, the growth and yield of 

Shorea robusta has been well investigated (Howard 

1925, Griffith and Bakshi Sant Ram 1943, Krishnaswamy 

1953, Mathauda 1953b, 1958, Chaturvedi 1975, Suri 

1975b, Raman 1976). The maximum biomass production 

was 14.62 tons/ha/year during the 18th year (Raman 

1976). In thinning trials, the results showed the 

superiority of the heavy and very heavy low thinning 

treatments (Krishnaswamy 1953). In India, research in 

thinning of plantations has been carried out, while this is 

not the case in other parts of the Indo-Malayan region. 

Dawkins (1963) introduced the crown diameter to bole 

diameter relation (also called growing space index) to 

estimate basal area density. This relation has been used 

for determining stand density regimes for Shorea 

robusta (Chaturvedi 1975). Suri (1975b) developed a 

quantitative thinning model for Shorea robusta which 

considers different types of crown disengagement 

regimes. Each of these crown disengagement regimes 

has a specific sequence of growing space index values. 

Rai (1979, 1981a) has reported growth rates of Hopea 

parviflora and H. wightiana. 

Site quality has a direct influence on growth rates. 

However, little has been researched in this respect. The 

effect of elevation on height and diameter growth of 

Dipterocarpus turbinatus was investigated by Temu et 

al. (1988). The decline in height and diameter growth 

was relatively small compared to the increase in 

elevation. The cause for the decline is probably due to 

the rapid drop in the water table and leaching of nutrients 

at the higher parts of hilly terrain. 

The economics of plantations of dipterocarp species 

have hardly been investigated. Lack of a sufficiently broad 

data base may have been the reason for the delay. 

Recently, some economic assessments were made on 

plantations of Shorea leprosula, S. parvifolia and S. 

platyclados in Peninsular Malaysia (Kollert et al. 1993, 

Zuhaidi et al. 1994, Kollert et al. 1994), and the 

following conclusions were drawn. The establishment 

and management of forest plantations are uneconomical 

if valued on financial terms alone. Forest plantations of 

relatively long rotations do not produce sufficient returns 

early enough to attract investment, especially from the 

private sector. Investors avoid the long gestation periods, 

the relatively low rate of return and the relatively high 

risk of investment. The venture of forest plantations will 

become economically attractive only by end of the first 

rotation, when the age class sequence is complete and 

future stand establishment is not by clear cut and planting 

but through natural regeneration. 


It is recommended that all research is carried out with 

the same set of dipterocarps (the most promising species 

for plantations). For species/provenance tests (species 

elimination, site adaptation), which usually remain 

untreated, it is recommended to include a standard 

silvicultural treatment. 

• Silvics and species choice: Build up of information 

on silvical and silvicultural characters including site 

requirements, establishment of a site adaptation trial, 

establishment of systematic species/provenance elimination 

trials, evaluation of the existing dipterocarp 

plantations throughout the region as a basis for the 

above-mentioned trials. 

• Seed: Seed production from trees/stands, seed orchard 

technology, dysgenic shifts as a basis for strategies 

in tree selection work. 

• Planting stock production: Comparative planting stock 

production test, comparative cutting propagation trial, 

mycorrhization techniques in nurseries.


• Planting site: Site preparation techniques for lineplanting/underplanting/open 

sites, optimal planting 

stock size for line planting/underplanting. 

• Planting: Deficiency symptoms, fertilisation trials; 

if the fertilisation trials are observed over longer time 

(e.g., into the weeding and cleaning period), it is advisable 

to overlay the fertilisation trial with a tending 

trial that includes a standard treatment. 

• Tending: Selective weeding procedures, assessment 

of weed vegetation concerning risks to the plantation 

crop, investigation into control of weed growth 

through shade management. 

• Re-establishment by natural regeneration: Assessment 

of existing dipterocarp plantations near rotation age 

as to constitution, composition, canopy structure, regeneration 

status, regeneration experiments in existing 

older experimental dipterocarp plantations. 

• Reforestation/afforestation: Investigations into site 

amelioration techniques, species adaptation trials, 

plantation site preparation procedures (nurse crops, 

fertilisation soil improvement procedures). 

• Agroforestry: Research should continue to test the 

use of promising dipterocarp species as agroforestry 

crop trees, e.g., Shorea macrophylla and other members 

of the pinanga group, Shorea javanica, etc. Such 

research would concentrate mainly on selection as well 

as agri-silvicultural systems. 

• Management: Stand establishment guidelines, speciessite 

matching procedures, weeding guidelines, feasibility 

studies on dipterocarp plantations, development 

of production schemes with early financial returns, 

growth analysis of the existing dipterocarp plantations. 


The author wishes to express his gratitude to Dr. Aminah 

Hamzah (Forest Research Institute Malaysia), Dr. J. McP 

Dick (Institute of Terrestrial Ecology), Dr. D. Baskaran 

(Forest Research Institute Malaysia), Dr. B. Hahn- 

Schilling (Malaysian-German Project ‘Forest 

Mananagement Information System Sarawak’), Dr. R. 

Nussbaum (Société Générale de Surveillance), Mr. R. 

Ong (Forest Research Centre, Sandakan, Sarawak), Dr. 

D. Simorangkir (University of Mulawarman), Mr. Truong 

Quang Tam (Institute of Tropical Biology, Vietnam) for 

providing important information. Dr. S. Appanah (Forest 

Research Institute Malaysia), Dr. P. Moura-Costa 

(Innoprise) and Dr. S.N. Rai (Forest Survey of India) 

kindly read the manuscript and their useful comments 

are gratefully acknowledged. 

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Non-Timber Forest Products 

from Dipterocarps 

M.P. Shiva and I. Jantan 

Introduction 

In the last half of the twentieth century timber has become 

the most important economic product from dipterocarps, 

but it does not have much impact on rural communities. 

Instead, the non-timber forest products (NTFPs) from 

dipterocarps such as nuts, dammar, resin and camphor, 

have a larger impact on the economies of the rural people 

and forest dwellers. In the past several decades synthetic 

materials have diminished the value of some dipterocarp 

NTFPs but at the same time others are beginning to gain 

value. As a result researchers have paid little attention to 

NTFPs and there is little detailed information on them. 

Their value to rural communities would have been better 

appreciated and critical in balancing the forces favouring 

logging against other socio-economic benefits. The 

advantages of managing NTFPs, previously known as 

minor forest products, are often ignored. Unlike timber, 

they are available at more frequent intervals and their 

harvesting is usually less destructive to the tree. Their 

value can be high, and as in some cases described here, 

may even pay towards the establishment of plantations 

for their production. In this chapter, the various NTFPs 

from dipterocarps are described, and wherever possible 

additional information on the methods of extraction, 

their industrial application and economic value is given. 

Ancient Records of Dipterocarps 

Perhaps the oldest written records of dipterocarps come 

from India; records of utilisation of dipterocarp timber 

and other products exist there since ancient times. The 

birth place of Buddha was Lumbini, situated on the bank 

of the River Rohini where there were groves of Shorea 

robusta (sal), called ‘Mangala Salvana’. Sukraniti and 

Kautilya have regarded sal amongst the strongest timber 

Chapter 10 

yielding trees of the forest. Plant remains excavated from 

Pataliputra show that sal was used for a wooden palisade 

made 2000 years ago. In Southeast Asia there is a long 

tradition of the use of NTFPs from dipterocarps. Their 

trade was extensive, and from the 1st century A.D. 

Chinese and Indian traders regularly visited the Southeast 

Asian ports for these products. Marco Polo’s chronicles 

of 1299 mention the trade of camphor (from 

Dryobalanops aromatica) by Arabs since the 6th 

century. 

NTFPs From Dipterocarps 

Much of the knowledge on the use of dipterocarp NTFPs 

is concentrated in two main regions, South Asia and 

Southeast Asia (mainly Indonesia, Malaysia, and the 

Philippines). In both regions, the dipterocarp products 

are essentially the same and four broad classes are 

predominant, viz., resins, dammar, camphor and butter 

fat. Besides these principal products, other plant parts, 

such as leaves and bark, are used to derive certain 

products. In both regions the extraction methods are 

common, however, the specific species yielding these 

products vary. Despite their importance, they have not 

been systematically exploited and have remained 

undervalued. 

Resins 

The dipterocarps are an important source of resins. The 

resin is secreted in cavities, and normally oozes out 

through the bark. The resins are of two kinds. The first is 

a liquid resin which contains resinous material and 

essential oils (oleoresins), remains liquid in nature and 

has a distinct aroma. It is often referred to as oleoresin 

in literature. Commercial production is often through 

artificial wounding. The second is the hard resin which

Non-Timber Forest Products from Dipterocarps 

is called dammar when obtained from dipterocarps. This 

is the solid or brittle resin, which results from hardening 

of the exudate following evaporation of the small content 

of essential oils. However, the classification of resins 

is very chaotic, and in the trade the term ‘dammar’ is 

also used occasionally to refer to an oleoresin. 

Oleoresins 

The genus Dipterocarpus is the principal source of 

oleoresins. The genus has large trees with erect trunks, 

the wood of which yields resin similar to copaiba. Other 

genera of lesser importance are Shorea, Vatica, 

Dryobalanops and Parashorea. All Dipterocarpus 

species produce a high proportion of oleoresins which 

come under various local names such as gurjan oil (India), 

kanyin oil (Burma) and minyak keruing (western 

Malesia). A well-known oleoresin comes from D. 

turbinatus which is the principal source of ‘kanyin oil’ 

in Burma and ‘gurjan oil’ in Bangladesh and India. The 

best yielding species are Dipterocarpus cornutus, D. 

crinitus, D. hasseltii, D. kerrii and D. grandiflorus 

(Malesia), D. turbinatus and D. tuberculatus (India, 

Bangladesh, Burma), D. alatus (Bangladesh, Andamans, 

Indochina) and D. grandiflorus (Philippines). 

Method of Tapping 

During the cold weather, a cone shaped cavity is cut into 

the trunk 1m from the ground and a fire lit to char the 

surface of the wound to induce the oleoresin flow. The 

oleoresin is periodically removed and when the flow 

stops, the wounded surface is either burnt or scraped or 

a fresh wound made to induce further flow. The collection 

season is November-May and a tree of 2 m girth can 

yield 9 kg of resin in one season. This resin compares 

favourably with balsam of copaiba (Balfour 1985). 

Traditionally in Burma, oleoresin was obtained by 

cutting 2-3 deep pyramidal hollows, (the apex pointing 

towards the interior of the stem), near the base of the 

tree and by applying fire to the upper cut surface. The oil 

was collected at the bottom of the hollow which was 

emptied at 3 or 4 day intervals. Fire was applied every 

time the oil was removed and the upper surfaces of the 

hollow were rechipped 3 or 4 times in a season. About 

180 kg of oleoresin oil was collected from 20 trees in a 

season. The oil was marketed locally in the form of 

torches and also exported. Later, tree tapping was 

prohibited owing to the heavy damage to the trees. 

188 

In Bangladesh, the practice was to cut a deep hollow, 

(transverse hole pointing downwards), in the tree and 

place fired charcoal in it during the night. The oil was 

removed in the morning and the charcoal replaced. The 

process was repeated until the oil ceased to flow. Three, 

four or more such hollows were made which often killed 

the tree. In Burma the charcoal practice was not adopted. 

In India, in the western-Ghat division of Coorg, the 

oil was collected by cutting a hole into the centre of the 

tree. It is also reported that a large notch was cut into the 

trunk of the tree about 75 cm above the ground level, in 

which fire was maintained until the wound was charred 

and the liquid began to ooze out. A small gutter was cut 

into the wood to a vessel attached to receive the oil. The 

average yield from the best trees was 180 litres per 

season. At 3 or 4 week intervals the old charred surface 

was cut off and burnt afresh. Tapping occurred from 

November to February and sick trees were rested for 1 

or 2 years. 

Properties and Uses of Gurjan Oil 

The exudate is milky and faintly acidic and when 

allowed to stand separates into 2 layers - a brown oil 

which floats on the surface and a viscous, whitish grey 

emulsion below. A pale yellow oil with a balsamic odour 

is obtained (yield 46%) through steam distillation of the 

oleoresin which leaves a dark, viscid, liquid resin. 

The commercial gurjan oil is the oleoresin mixed 

with small quantities of oleoresin from Dipterocarpus 

alatus, D. costatus and D. macrocarpus. It is a viscid 

fluid, highly florescent, transparent and dark reddish 

brown in colour when seen against the light. It oxidises 

when exposed to the atmosphere. The essential oil 

consists of two distinct sesquiterpenes, alpha and beta 

gurjunene. 

The resin contains a crystallisable acid, gurjunic acid 

(C H O ), devoid of acid character as in copaiba (a 

22 34 4 

resin containing a small portion of naphtha), which may 

be removed by warming it with ammonia and 0.08% 

alcohol. It is partially soluble in ether, benzol or sulphide 

of carbon. The portion of resin, which is insoluble even 

in absolute alcohol, is uncrystallisable. A remarkable 

physical property of this oil is that at a temperature of 

130oC it becomes gelatinous, and on cooling does not 

recover its fluidity. 

The oleoresin is applied externally to ulcers, ring 

worm, and other cutaneous infections. It is a stimulant


to mucous surfaces and also a diuretic (Kirtikar and Basu 

1935, Martindale 1958). It is an ingredient of 

lithographic ink and varnish and an anticorrosive coating 

composition for iron. It is occasionally used as a 

preservative for timber and bamboo. Mixed with 

powdered dammar from Shorea robusta or S. siamensis 

it forms a dark brown paste used for caulking boats and 

water proofing bamboo baskets used for carrying water. 

Gurjan oil is a good solvent for caoutchouc 

(unvulcanised rubber) which is applied to cloth to make 

it water-proof. This cloth resists insect-attacks. 

Traditional Uses 

a) Medicine: Ancient literature reveals that gurjan oil 

was used by the Mohammedans and it was first 

mentioned in the ‘Makhzan’ Materia Medica as ‘Duhnel-Garjan’. 

Its essential oil is effective in the treatment 

of genito-urinary diseases. The Pharmacopoeia of India 

1868, officially describes it as a stimulant of mucous 

surfaces, particularly those of the genito-urinary system, 

and as diuretic (Watt 1899). However, users of 

indigenous systems of medicine in India find it less 

powerful than copaiba. It is useful in leucorrhoea and 

other vaginal discharges, psoriasis, including lepravulgaris 

and also in the treatment of leprosy (used both 

externally and internally). All varieties of gurjan oil are 

equally useful as local stimulants but red, reddish brown, 

pale or pale white varieties are best for internal use. 

The efficacy of this oil is enhanced with the addition of 

chaulmugra oil. 

An ointment is prepared by mixing equal parts of oil 

and lime water. In European medicine gurjan oil was 

mainly used as an adulterant for copaiba. 

b) Domestic and Industrial Uses of Gurjan Oil: Gurjan 

oil was used in Burma for torches, and later, as lamp 

oil. It could be used as a varnish by mixing it with some 

good drying oil or by evaporating the essential oil. The 

oil was a good substitute for linseed oil and balsam of 

copaiba and prized as a colourless varnish and for drying 

paints. 

c) Trade of Gurjan in the 19th Century: In Burma and 

Bangladesh gurjan oil was mainly used for torches but 

its trade was limited due to the cheap price of kerosene. 

However, gurjan oil from Singapore and Malaya 

was a common article of trade in Thailand. The oil produced 

in South India and Andaman Islands was traded in 

Europe for use in artworks. The price of the black or 

dark brown varieties (‘Kala gurjan Tel’) was half the price 

189 

of the red or reddish brown (‘Lal gurjan Tel’) and pale 

white (‘Sufed gurjan Tel’) varieties. 

Other Sources of Oleoresin 

Other South Asian species important for the production 

of oleoresins include Dipterocarpus alatus and D. 

tuberculatus. The former is found in Chittagong 

(Bangladesh), Andamans (India) and Burma. D. 

tuberculatus occurs in Burma, and to a restricted extent 

in India and Bangladesh. 

Dipterocarpus alatus produces an oleoresin that 

contains 71.6% volatile oil. The oil known as ‘kanyin 

oil’ in Burma is an antiseptic applied to clean wounds 

and has been used as a substitute for copaiba in the 

treatment of gonorrhoea. In Burma, it is also used for 

treating ulcers and sores in the hoof and foot disease of 

cattle. The oil is used by forest dwellers to fuel torches 

made of rotten wood and for waterproofing the oil cloth 

used for Burmese umbrellas. It has been used in the 

preparation of lithographic inks and has been tried as a 

varnish and as a substitute for linseed oil in zinc paints. 

Its bark is a tonic given for rheumatism. 

The method of tapping oleoresins from almost all 

other species resembles that of D. turbinatus. A notch 

is made into the trunk and the wound blazed to stimulate 

resin flow. Resin is collected periodically and either the 

wound is scraped for new flow or another wound made. 

The trees eventually succumb to the regular wounding, 

and the timber, unsuitable for construction work, is used 

as fuelwood. The oil and resinous thicker substance 

mixture is strained through a cloth whereby the clear oil 

separates itself from the resinous portion. Dipterocarpus 

alatus provides the wood-oil, pegu. 

Dipterocarpus tuberculatus is the principal source 

of oleoresin known as ‘In oil’ in Burma and ‘gurjan oil’ 

in India. Its exudate is thicker than ‘kanyin oil’ from D. 

turbinatus and flows freely from the wound without the 

aid of fire. Throughout the year, resin oozes 

simultaneously from several niches on a tree. The oil 

was collected 4-10 times a month from August-February 

and 300 trees yielded about 36 kg a month. At the end of 

the season the dried resin was scraped off and used to 

make torches. Freshly collected oleoresin is a pale brown 

substance with specific gravity 1.029; acid value 17.8 

and ester value 0. It yields a yellow brown essential oil 

on steam distillation. The oil is used for varnishes and 

for water proofing umbrellas and bamboo well-baskets. 

The oleoresin is used with assafoetida and coconut oil 

as an application for large ulcers (Watt 1889).


Shorea robusta or sal is another important producer 

of oleoresin in Bangladesh, India and Nepal. It yields an 

oleoresin known as sal dammar, ‘ral’ or lal dhuma’. Earlier 

tapping methods gave low and erratic yields. The method 

recently employed is to cut 3-5 narrow strips of bark 

90-120 cm above the ground. When the tree is blazed 

the oleoresin oozes out as a whitish liquid and on exposure 

it hardens quickly and turns brown. The cut is freshened 

by scraping off the hardened resin. In about 12 days the 

grooves are filled with resin. The grooves are freshened 

and resin is collected periodically in July, October and 

January. A good mature tree yields about 5 kg of resin 

annually. 

The essential oil, sal resin, on dry distillation yields 

an essential oil, known as ‘chua oil’. The yield of the oil 

varies from 41 to 68% depending upon the source of the 

oleoresin samples. The oil is light brownish yellow in 

colour and has an agreeable incense-like odour, with 

specific gravity 0.9420, acid value 4.42, saponification 

value 15.72 and saponification value after acetylation 

39.49. It consists of 96.0% neutral, 3% and 1% phenolic 

and acidic fractions, respectively. Chua oil is used as a 

fixative in heavy perfumes, for flavouring chewing and 

smoking tobacco and in medicine as an antiseptic for 

skin diseases and ear troubles. The non-phenolic portion 

of the oil has a suppressing effect on the central nervous 

system, the phenolic portion is less effective. 

Vateria indica is also an important source of 

oleoresin in India. The trade names used for the oleoresin 

are piney resin, white dammar, Indian copal and dhupa. 

The trees are tapped either using semi-circular incisions 

or a fire is lit at the base of the tree so as to scorch the 

bark, which then splits and the resin exudes. The resin is 

in three forms: i) compact piney resin which is hard, in 

lumps of varying shapes, bright orange to dull yellow in 

colour, with a glossy fracture and resembling amber in 

appearance, is called Indian dammar; ii) cellular soft 

piney resin which occurs in shining masses, having 

balsamic odour, and light green to yellow or white in 

colour, is called a piney varnish; and iii) dark coloured 

piney resin from old trees. The resin is a complex 

mixture of several triterpene hydrocarbons, ketones, 

alcohols and acids along with small amounts of 

sesquiterprenes. On distillation, the oleoresin gives an 

essential oil (76%) with a balsamic odour. The oil 

consists of phenolic constituents and azulenes, with the 

latter predominating. The essential oil has a marked 

antibacterial property against gram negative and gram 

190 

positive microorganisms (Howes 1949, Chopra et al. 

1958). The resin readily dissolves in turpentine, 

camphorated alcohol and is used in the manufacture of 

varnishes, paints and anatomical preparations. The 

liquefied resin mixed with hot drying oil makes a varnish, 

superior to copal, for carriages and furniture. The resin 

is used to make incense, for setting gold ornaments, 

caulking boats (Trotter 1940) and in rural areas, resin 

mixed with coconut oil is used as torches and candles. It 

is a good substitute for Malayan dammar and, in solution 

in chloroform, for amber in photographers’ varnish. The 

resin has medicinal value. It is credited with tonic, 

carminative and expectorant properties and is used for 

throat troubles, chronic bronchitis, piles, diarrhoea, 

rheumatism, tubercular glands, boils etc. Mixed with 

gingili (sesame) oil, it is used for gonorrhoea and mixed 

with pounded fruits, obtained from Piper longum (longpepper), 

and butter or ghee it is useful for the treatment 

of syphilis and ulcers. An ointment of resin, wax and the 

fat of Garcinia indica is effective against carbuncles. It 

forms a good emollient for plasters and ointment bases 

(Kirtikar and Basu 1935, Chopra et al. 1958, WOI 

1989a). 

In Southeast Asia the important oleoresin trees are 

Dipterocarpus cornutus, D. crinitus, D. hasseltii, D. 

kerrii and D. grandiflorus. The old method of tapping 

is by notching a hole in the trunk and blazing to stimulate 

further oleoresin flow. This is repeated at about weekly 

intervals and the yield per tree is 150 to 280 ml per 

tapping (Gianno 1986). A less brutal method has been 

developed, known as the barkchipped method 

accompanied by application of chemical stimulants, 

which is less destructive and the yield and oleoresin 

quality better (Ibrahim et al. 1990). The oleoresin is 

processed to separate the essential oil from the resin. 

The essential oil, known commercially as gurjan balsam, 

is used as a fixative or a base in perfume preparations 

and occasionally as an adulterant of patchouli and copaiba 

balsam oils. Traditionally the oleoresin is used for 

caulking the inside of boats, coating wood as a protection 

against weather, in torches, and for medicinal purposes. 

The oil is also used to make varnishes in backyard 

industries (Burkill 1935). While the biggest suppliers 

of gurjan balsam oil are Indonesia, Malaysia and Thailand, 

limited quantities are produced in India and the 

Philippines. Sumatra is the biggest producer of all, and 

in 1984 it produced about 20 tonnes of the oil (Lawrence 

1985). The oil is now becoming scarce with an


increasing demand, resulting in increased prices. The 

price is currently over US $30 per four gallon tin. The 

oleoresin is mainly collected by natives and aborigines 

and has a ready market in Singapore where it is exported 

to Europe. 

There are several other less important dipterocarps 

which are tapped for oleoresin: 

• Dipterocarpus bourdilloni, a species from Kerala, 

India, yields opaque, straw yellow, viscid oleoresin 

which on standing deposits a crystalline unsaturated 

hydroxy ketone, C H O , M.P. 125 24 42 2 o-126oC. When 

distilled with steam at 100o , 245o and 380o C it gives 

37% 65% and 76% respectively, essential oil (Anon. 

1989). 

• Dipterocarpus costatus from Burma produces a resin 

used in the treatment of ulcers. 

• Dipterocarpus gracilis found in Bangladesh and India, 

produces a good quality oleoresin used in the soapindustry 

and also as an antiseptic for gonorrhoea and 

urinary diseases. 

• Dipterocarpus grandiflorus belonging to the 

Andamans, Thailand and the Malesian region, produces 

an oleoresin which exudes as a thick fluid which 

changes into a semi-plastic mass on long exposure to 

air. The exudate has a thick honey - like consistency 

and a balsamic odour, is reddish brown in colour and 

contains 35% volatile oil and a hard, yellow, lustrous 

resin soluble to the extent of 75% in alcohol. The 

oleoresin used in varnish is dissolved in equal parts of 

linseed oil and turpentine, and dries slowly to a tough 

hard film. 

• Dipterocarpus hispidus of Sri Lanka produces resin 

that has been found to contain dipterocarpol, 

dammarenediol, and ocotillone. 

• Dipterocarpus indicus is a species of west coast, 

tropical, evergreen forests of India. Its oleoresin is 

used in the preparation of spirit, oil varnishes and 

lithographic inks. It is also used as an adulterant of 

dammar and as an application for rheumatism. 

• Dipterocarpus macrocarpus of India and Burma 

produces oleoresin that is used as a lubricant and in 

soap making. 

• Dipterocarpus obtusifolius of Burma, Thailand, 

Indochina and northern Peninsular Malaysia has 

oleoresin that yields a clear, white or yellow resin 

which burns readily (Watt 1899). 

• Dryobalanops aromatica found in W. Malesia yields 

an oleoresin that is aromatic, volatile and is used in 

191 

medicine, in the preparation of toothpaste, powders 

and as a diaphoretic and antiseptic; it is also used for 

treating hysteria and dysmenorrhoea (Agarwal 1986). 

• Parashorea stellata, a species found in Burma, 

Thailand, Indochina and Peninsular Malaysia, produces 

resin which is used as a fumigant. 

• Shorea siamensis, found in Burma, yields a red resin. 

• Shorea megistophylla, a species found in Sri Lanka, 

yields resin that contains ursolic acid, 2-alpha, 3-beta 

dihydroxyurs - 12-Cn-28-oic acid, asiatic acid and 

Caryophyllene (Bandaranayake et al. 1975). 

• Shorea obtusa from Burma produces a white resin. 

• Shorea roxburghii, a widespread species found in 

India, Burma, Thailand, Indochina and Peninsular 

Malaysia, yields a resin, which is used as stimulant and 

for fumigation (Anon. 1985a, WOI 1988). 

• Shorea tumbuggaia is a species found in India which 

yields a resin which is used as an incense and as a 

substitute in marine yards for pitch. It is also used in 

indigenous medicine as an external stimulant and a 

substitute for Abietis; Resina and Pix Burgundica of 

European pharmacopoeias (Watt 1899). 

• Vatica chinensis, a species found in India and Sri Lanka, 

yields abundant resin nearly transparent and yellow in 

colour resembling that of V. lanceaefolia and used in 

varnishes. 

• Vatica lanceaefolia from Bangladesh, Burma and India, 

yields from its bark a clean, white, aromatic oleoresin 

which turns light amber in colour on hardening and is 

used as incense. When distilled, a strong smelling 

essential oil (9.2%) commonly known as scented 

balsam or ‘chua’ is obtained. It is used to flavour 

chewing tobacco with betel leaves. It also yields a 

strong smelling balsam ‘ghunf’ used in religious 

ceremonies. Piney tallow, dupade oil, piney yennai, or 

tam, obtained from the seeds is mainly used for lamps 

and is also suitable for soap and candle making. 

• Vatica obscura, found in Sri Lanka, produces a gummy 

exudation used for caulking boats. 

• Vatica tumbuggaia, a species found in India, yields a 

good quality oleoresin. 

Dammars 

Dammar is the hard, solid or brittle resin which hardens 

soon after exudation when its small content of essential 

oil evaporates. Although all dipterocarps produce 

dammar, only a few are of commercial importance. In 

Southeast Asia, the important genera are 

Neobalanocarpus, Hopea and Shorea. The most


important Malayan varieties are ‘damar mata Kuching’ 

from Hopea micrantha and related species, ‘damar 

penak’ from Neobalanocarpus heimii, and ‘damar 

temak’ from Shorea crassifolia (Blair and Byron 1926). 

The principal dammars of India are sal dammar from 

Shorea robusta and white dammar from Vateria indica. 

H. odorata from Bangladesh, Burma and India, is the 

source of dammar, known commercially as ‘rock 

dammar’. Dammars are also produced in the island of 

Borneo, Java, Sumatra, Thailand, and Vietnam. The 

outstanding commercial variety, the Batavian dammar, 

comes from Shorea wiesneri from Java and Sumatra 

(Burkill 1935). 

Dammar is found as natural exudations, on living trees, 

in lumps on the ground beneath the trees, near dead 

stumps, or even found buried in the ground. These 

dammars are usually collected by aborigines. Natural 

exudation also occurs from trees which are unhealthy or 

damaged by the heartwood borer. Sal resin occurs in 

rough, stalactitic brittle pieces, 16-24 cm in size, pale 

creamy yellow in colour, nearly opaque with a faint 

resinous balsamic odour. It is produced commercially 

by tapping the trees. 

Dammars are used traditionally for making torches, 

caulking boats, and handicrafts. The dipterocarpaceous 

resins have also been used as adulterants for the aromatic 

resin produced by Styrax benzoin (Styracaceae) which 

is used as an incense and medicine. Sal dammar is widely 

used as incense in religious ceremonies and as a 

disinfectant fumigant. Large quantities of dammar are 

an important ingredient in ‘Samagri’ used for cremation. 

It can also be used for hardening softer waxes for shoepolish 

manufacture, carbon paper, typewriter-ribbon, and 

in inferior grades of paints and varnishes for indoor 

decorative work, and for mounting microscopic objects. 

It has been used as a plastering medium for walls and 

roofs and as a cementing material for plywood, asbestos 

sheets, etc. Tribal people in India mix the resin with bees’ 

wax and red-ochre for fastening spear and arrow-heads. 

The resin is used in indigenous medicine as an 

astringent and detergent and is given in diarrhoea and 

dysentery. It is also an ingredient of ointments for skin 

diseases and has curative properties against ear troubles, 

toothaches, sore eyes, ulcers and wounds. The resin in 

powder form is used as an ointment for wounds and sores 

(Anon. 1985a). 

More recently, the dammars are being used in many 

technical preparations, such as in the manufacture of 

192 

paints, batik dyes, sealing wax, printing inks, varnishes, 

linoleum and cosmetics. Triterpenes isolated from 

dammar have been found to exhibit in vitro antiviral 

activity against Herpes simplex virus type I and II 

(Poehland et al. 1987). 

Dammar export is mainly from Indonesia. The 

following species produce high quality resins which fetch 

a high price: Shorea javanica, S. lamellata, S. virescens, 

S. retinodes, S. assamica ssp. globifera, Hopea 

dryobalanoides, H. celebica, H. beccariana and Vatica 

rassak (Jafarsidik 1987). Indonesia exports annually 

2000 - 7000 tonnes worth US $1.6 million. The dammar 

is mainly exported to Japan, Taiwan, Singapore, Germany 

and Malaysia. 

Dammar in Sumatra is produced mainly from dammar 

gardens that are part of an agroforestry system. With the 

decline in forest areas, farmers have resorted to 

developing resinous tree plantations. However, in 

Lampung, Sumatra, man-made dipterocarp gardens have 

been established since the 19th century (Rappard 1937). 

Shorea javanica a native of the region, is grown in an 

agroforestry system with other crop trees (Torquebiau 

1984), as is Hopea dryobalanoides. Villagers tap the 

trees by cutting holes of about 10 cm wide and 15 cm 

deep into the trunk to stimulate resin flow. The resin is 

collected periodically and the holes deepened. When the 

hole reaches the centre of the trunk, a new hole is made. 

Tapping commences when the trees are about 20 years 

old, and continues for 30 years when production declines. 

A fully productive tree may produce 50 kg of resin each 

year. One hectare of dammar gardens can produce 4.8 

tonnes per year (Torquebiau 1984). 

Camphor 

Trade in camphor (known as Borneo or Sumatra camphor 

(bhimsaini-kapur, barus kapur)) is ancient. Camphor was 

used mainly in China and its source was the gregarious 

Dryobalanops aromatica (kapur) forests in North and 

East Sumatra and Johore. Other species, such as D. 

beccarii, also yield camphor but to a lesser extent. The 

camphor is found in cavities or fissures in the wood in 

the form of solid camphor, or a light fluid called camphor 

oil. The tree is felled, cut into blocks and split into wedges 

to remove the camphor. One hundred trees rarely yield 

more than 8-10 kg solid camphor. In solid form it occurs 

in white crystalline translucent fragments, sometimes 

in long, 5 kg pieces. It closely resembles the camphor 

from Cinnamomum camphora but it is heavier than


water, does not volatilise at room temperature, and 

possesses a characteristic pungent odour and burning 

taste. It is used in medicine, perfumery and organic 

syntheses. Borneo camphor is almost pure d-borneol 

(C 10 H 17 OH, M.P. 209 o C) and is highly prized in Indian 

medicine. Chinese and Japanese also attribute a higher 

medicinal value to it than the essential oil from the wood 

of Camphora officinalis. It is converted into ordinary 

camphor by heating with boiling nitric acid. (Balfour 

1985, WOI 1989a). Dryobalanops aromatica is no 

longer a major source of camphor now that 

Cinnamomum camphora is used in the chemical industry 

and camphor can be synthesised more cheaply from 

pinene. 

Butter Fat 

Another major dipterocarp NTFP in Borneo is butter fat. 

Shorea species (the Pinanga type) produce illipe nuts 

which are called engkabang and tengkawang in Malaysia 

and Indonesia, respectively. The nuts are generally 

collected in the wild but some experimental plantations 

of S. macrophylla, S. stenoptera, S. mecistopteryx, S. 

aptera and other related species exist in Sarawak and 

Kalimantan (Tantra 1979). The fruiting is somewhat 

aperiodic but at about four year intervals the forests fruit 

heavily. The natives of Borneo extract oil from the nuts 

for use as cooking oil (Anderson 1975). The kernels are 

exported to Europe, Japan and West Malaysia. The illipe 

fat extracted from the kernel is used in the confectionery 

industry, especially in the manufacture of chocolate. The 

illipe fat has a high melting point, and when blended with 

cocoa butter remains solid at room temperatures. 

Likewise, illipe fat is added to cosmetics such as lipstick. 

The illipe nuts have a high value with prices from US 

$2300-2700 per tonne in the 1980s (Anon. 1985b), and 

during peak fruiting years exports from Borneo can reach 

50 000 tonnes (Wong Soon 1988). 

Shorea robusta (sal) from the Indian region is 

another important source of butter fat. The kernels, 

constituting 72% of the nut weight contain 14-20% of 

fatty oil known as sal-butter. Sal seed oil has assumed 

great importance for use as a cooking medium, industrial 

oil, illuminant, lubricant and as a substitute for cocoabutter. 

It is also suitable for soap making after blending 

with other softer oils. The sal fat is obtained by boiling 

the husk seeds in twice the volume of water and skimming 

off the oil which solidifies to a buttery consistency in 

cold weather. In India sal fruits must be collected before 

193 

the onset of the monsoon when it becomes difficult to 

dry and decorticate them. The dried fruits can be 

decorticated by hand or with mechanised decorticators 

after manually dewinging them. The fruits are spread on 

a hard surface to a thickness of about 10 cm and beaten 

with sticks to dewing them. The oil is also obtained by 

solvent extraction of seeds by flaking procedure. The 

particle size of the kernel is reduced to 7-10 mesh by 

using fluted rolls and cooked at 2.25 kg cm -2 steam 

pressure with limited open steam injection so as to adjust 

the meal moisture content in the flaking rolls to about 

15%. A steam jacketed flight screw kettle is most suitable 

for cooking the meal. The flakes are tempered to a 

thickness of 0.24 - 0.3 mm with a moisture content of 

8%. They do not show any sign of disintegration on 

solvent impact due to the kernels' high starch content. 

Studies show that, even with proper conditioning of the 

kernels, it is not possible to obtain a good yield of fat by 

expeller. The fat is refined by a conventional method of 

alkali refining. However, the small recoverable fat 

content of 14% is disadvantageous because the fat 

contains various kinds of pigments even after refining. 

The glycerides of the kernel fat are a rich source of stearic 

and oleic acid (44.2 and 44.9%) in addition to palmatic 

(4.6%) and arachidic acid (6.3%). 

The kernels of Vateria indica from India yield about 

22% fat by solvent extraction. This is known as piney 

tallow, malabar tallow or dhupa tallow. It is extracted by 

boiling the powdered kernels in water, then allowing the 

extract to cool and skimming off the floating fat. The fat 

has a slight, pleasant odour and is greenish yellow at first 

but rapidly lightens in the air. It consists of glycerides 

of solid acids (53%) and liquid acids. (Puntembaker and 

Krishna 1932). The tallow is edible after refining, but is 

not in common use. It is used in confectionery and as an 

adulterant of ghee, in candle and soap manufacture, and 

for sizing cotton yarn instead of animal tallow. It is also 

used as a local application for rheumatism. 

Tannin 

The leaves and bark of several dipterocarps are a source 

of tannin. The bark of Hopea parviflora from India is a 

good tanning material for heavy leather, particularly when 

used with other tanning materials, for example myrobalan 

bark in a 2:1 ratio which gives a good quality, reddish 

brown leather resistant to mould. The bark contains 14- 

28% tannins and the solid extract, an astringent with slow 

diffusion speed, 70% tannins (Anon 1985a). The tannin


content in the bark of Dipterocarpus tuberculatus is 

24%, while young leaves have 10-12% and may be used 

in direct light leather tanning. Sal bark, together with the 

leaves and twigs, is also a promising tanning material. 

The tannin content is: bark 7%, young leaves 20%, twigs 

and leaves 22% and powder dust 12%. The aqueous 

extract of bark is a pale reddish colour and the tannins 

are of pyrogallol type. The extract is used locally for 

cheap tanning or in a blend with other tanning materials. 

The dry leaves of H. odorata contain 10% tannin and 

are used in crude tannery. The tannin extract is rich and 

produces strong leather (Anon. 1985a, Agarwal 1986). 

The fruit of Vateria indica contains 25% tannin. 

Lac Host 

A few dipterocarps are known to host the lac insect 

(Lacifer lacca), a source of lac. Shorea roxburghii, a 

species found in Burma and India, is a valuable host in 

South India, and yields a good crop when inoculated with 

‘Deverbettakusum’ variety in Karnataka. Shorea talura 

is another important lac host plant of Karnataka in India 

(Krishnamurthy 1993). Shorea obtusa, a species found 

in Burma, is an occasional host and sal is the source of 

the ‘Kusumi’ strains of lac insect. 

Other Products 

In addition to the important products described above 

there are other dipterocarp NTFPs. The sal tree yields 

many of these products. Its leaves are a good source of 

income to the tribals in India who make them into plates 

and cups or use them as wrappers for home-made cigars. 

They are also used for thatching huts in the villages and 

as a medium to poor grade fodder containing 0.94% 

nitrogen and 2.97% ash. Sal leaves are one of the primary 

hosts of tassar silk-worm (Antheraea mylitta). Roasted 

sal seeds, although not very palatable, are sometimes 

eaten, and decorticated seeds are used as poultry feed. 

Dried seed meal contains: moisture 5.23%; protein 

6.16%; ether extractive 16.77%, crude fibre 4.81%, N. 

free extractive 63.25%, calcium 0.18%, total ash 3.78% 

and acid insoluble ash 0.95%. 

A light grey, somewhat granular cellulose gum is 

prepared from the bleached, bright cellulose obtained 

from the spent bark. This compares favourably with 

commercial grade technical gums. The cellulose from 

the spent bark is also suitable for making wrapping paper. 

Lignins from wood waste are used as wood-adhesive. The 

bark is oily, bitter, acrid and anthelmintic and can cure 

194 

ulcers, wounds and itches. It is also a useful raw material 

for fibreboards (WOI 1988). Sal oil cake, used as cattle 

and poultry feed, contains 10-12% protein and about 50% 

starch. It can also be used as a fertiliser. Sal flowers are 

produced in abundance and are the source of honey. Santal 

tribals use the bark for preparing red and black dyes and 

wood ash in dyeing. 

A number of minor products derived from the wood 

also need mention. Wood of Shorea robusta and Vatica 

lanceaefolia is extensively used as firewood and for 

making charcoal. However, fuelwood should only be 

harvested at the time of clear felling at fixed rotations 

when unsuitable wood for timber can be utilised for 

firewood and charcoal making. The branches and thick 

twigs can be converted into charcoal in a specially 

designed kiln for supplementing the energy requirements 

after converting charcoal into briquettes. Briquetted 

charcoal and sawdust are good fuels for domestic and 

industrial purposes. Briquettes made with suitable 

binders from inferior grade gum, gum resin or pulp and 

juice from Agave/Furcraea species (Verma et al. 1979, 

Gulati et al. 1983) without the traditional use of clay 

and molasses ignite easily, do not emit smoke and 

provide sustained heat. 

The sal tree is considered to be the home of spirits 

and many gods, and tribals build their shrines under its 

shade and worship the tree as a whole. The Bagdis and 

Bauris tribes of Bengal are married under an arbour made 

of its branches. The sal tree in full bloom is worshipped 

in some villages by childless couples. Buddhists also 

worship the tree as it is believed that Buddha’s mother 

held a branch in her hands when Buddha was born, and it 

was under the shade of this tree that Buddha passed the 

last night of his life on earth (Bennet et al. 1992). 

Other valuable dipterocarps in the South Asian region 

include: 

• Hopea odorata of which the bark is an astringent and 

masticatory for gums. 

• Vateria copallifera of which the cotyledons are ground 

into an edible flour and the bark is used for arresting 

toddy fermentation (Anon. 1985a). 

• V. indica of which the fruit is ground into flour. The 

seed cake, unpalatable to livestock, is used as a manure, 

especially in coffee plantations. However, the cake, 

when mixed with other concentrates such as bran or 

groundnut cake, can be utilised for cattle feeding. The 

bark is an antidote (alexipharmic) in Ayurvedic 

preparations. The juice of the leaves is applied to burns


and is orally administered to prevent vomiting (WOI 

1989b). 

Socio-economic Perspectives 

In general dipterocarp NTFPs have been mainly used as 

subsidiary products by village people. However, some 

products from a few species have assumed much greater 

importance due to their demand. These few products have 

gained commercial importance in industry and trade due 

to their properties and chemical constituents. At present 

in the southern Asian countries, forest management 

systems have banned or restricted timber harvesting and 

so there is a need to generate more revenue from the 

NTFPs, especially during the prescribed long rotation 

for tree felling. 

Amongst the dipterocarp genera, Dipterocarpus, 

Dryobalanops, Hopea, Shorea, Vateria and Vatica are 

the important sources for NTFPs. The oleoresin and 

seeds are the most important for various uses while the 

leaves, bark, and twigs are useful for medicinal or tanning 

purposes. Shorea robusta is the only tree considered 

sacred and associated with different beliefs and religions. 

Critical analysis reveals that commercial extraction/ 

harvesting of different NTFPs aids socioeconomic 

development. Local harvesting of NTFPs by village and 

forest dwellers for traditional uses will persist. The 

present system of exploitation by local people is 

generally detrimental and therefore, improved collection 

methods are needed to provide sustained production and 

income. Value adding by local processing is desirable to 

increase returns. 

Strategies for NTFP Development in 

Forest Management 

Development strategies will differ for the extraction of 

oleoresin, seeds, bark and leaves from any species of 

NTFP importance. Owing to the erratic and unregulated 

extraction of NTFPs from different species, the 

economic returns do not properly accrue to the 

collectors. Therefore, it is essential that there are 

scientific measures to ensure better gains for improving 

the socioeconomic condition of not only the village and 

forest dwellers, including tribals, but also other industrial 

entrepreneurs associated with the utilisation, marketing 

and trade of the various products. Specific mention is 

made below of the development of NTFPs from 

dipterocarps: 

195 

1. Resins (Oleoresins and Dammars): These are the 

most important commercial products obtainable 

from Dipterocarpus alatus, D. grandiflorus, D. 

indicus, D. tuberculatus (gurjan/In oil), D. 

turbinatus (gurjan/kanyin oil), Hopea odorata (rock 

dammar), Shorea robusta (sal dammar), Vateria 

indica (white dammar) and Vatica lanceaefolia. 

Methods of obtaining oleoresin/dammar from these 

species have been discussed, but as yet there is no 

foolproof scientific method of tapping. Research 

should be done, according to the species, on: the 

optimum size, shape and depth of the blaze to avoid 

damage to a tree on a harvest rotation of several decades; 

the appropriate collection season and duration 

with adequate freshenings; and obtaining sustained, 

optimum oleoresin yields. 

Further, depending upon the constituents of each oleoresin, 

they can be put to specific industrial uses. 

Therefore, in order to make maximum gains from 

the value-added products, the raw material must be 

graded and processed prior to manufacturing the essential 

oil and various derivatives. These exercises 

will go a long way in improving the socio-economic 

conditions of all those involved in the oleoresin trade. 

2. Camphor: Dryobalanops aromatica is the only important 

dipterocarp producing camphoraceous oleoresin. 

This is extracted when the trees are felled 

for wood. Marco Polo, in 1299, mentioned that its 

camphor was traded by Arabs in the sixth century. The 

camphor was obtained, from concentrated occurrences 

of this species in North and East Sumatra and 

Johore. Now, camphor from this species is not commonly 

used, owing to the convenient availability of 

alternative sources. 

3. Seeds: Collection of seeds either for edible and medicinal 

purposes or recovery of fatty oil is made on 

large scale from Shorea robusta in South Asia and 

S. macrophylla in Malaysia, followed by S. aptera, 

S. obtusa, S. stenoptera, Vateria indica, V. 

copallifera, and Vatica lanceaefolia. To maintain the 

product quality and achieve maximum returns it is 

essential to collect the seeds in the appropriate season 

and stage of development and to properly grade 

and process them. 

4. Leaves: Shorea robusta leaves are important for local 

and commercial manufacture of cups, platters and 

cigar wrappers. The leaves of other species, such as 

Dipterocarpus tuberculatus, Hopea odorata and


Vateria indica, are mostly used for tanning and medicinal 

purposes. To increase the collector’s income, 

the leaves must be collected at the appropriate season 

and stage of growth and be properly dried and 

graded. 

5. Bark: This is used for tanning and medicinal purposes, 

for example: Dipterocarpus alatus (medicine), 

D. tuberculatus (tannin), Hopea odorata (tannin), 

H. parviflora (tannin), Shorea robusta (tannin, 

medicine, gum), Vateria copallifera (toddy fermentation) 

and V. indica (medicine). 

Only the bark of S. robusta is utilised on a large scale. 

Collection of bark from standing trees is detrimental 

to tree growth, so improved methods of bark 

extraction are needed or bark utilised only from felled 

or dead trees. 

There is good scope for greater utilisation of various 

dipterocarp NTFPs for socioeconomic development. 

This will be enhanced by further research and training in 

a range of technologies. These include: oleoresin and 

dammar tapping techniques; seed, leaf and bark 

harvesting; grading and processing standardisation; 

chemical evaluation of derivatives; and marketing and 

pricing analysis. 


We would like to thank Manuel Ruiz-Perez (Center for 

International Forestry Research) for comments and suggestions 

for the improvement of this chapter. 

References 

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Prakashan, Calcutta. 

Anderson, J.A.R. 1975. The potential of illipe nuts 

(Shorea spp.) as an agricultural crop. In: Williams et 

al. (eds.) Proceedings of Symposium on South East 

Asian Plant Genetic Resources, Bogor, Indonesia, 20- 

22 March, 1975, 217-230. BIOTROP, Bogor. 

Anonymous. 1985a. Dipterocarps of South-Asia. RAPA 

Monograph 4/85, FAO Regional Office for the Asia 

and the Pacific, Bangkok. 

Anonymous. 1985b. Statistics of external trade in 

Sarawak (1976-1986). Dept. of Statistics (Sarawak 

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Balfour, E. 1985. Cyclopedia of India and of Eastern 

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A 

Acacia arabica, 154 

Acacia auriculiformis, 161 

Acacia mangium, 154 

Adenanthera spp., 161 

Aegle marmelos, 165 

Agave, 194 

Agrobacterium tumefaciens, 115, 121 

Alati section, 32 

Albizia, 91 

Alstonia, 91 

Amanita, 99, 104 

ANCISTROCLADACEAE, 25 

ANGIOSPERMS, 21, 31, 49 

Angulati section, 32 

Anisoptera, 8, 14, 15, 16, 17, 23, 24, 28, 30, 31, 32, 

33, 34, 45, 46, 59, 75, 100, 159 

Anisoptera costata, 49, 58, 60, 64, 78, 152, 161 

Anisoptera costata Korth., 100 

Anisoptera curtisii, 49 

Anisoptera glabra, 118, 152 

Anisoptera laevis, 75, 152, 153, 155, 156, 158 

Anisoptera laevis Ridl.Pen., 100 

Anisoptera marginata, 58, 64, 79 

Anisoptera marginata Korth., 100 

Anisoptera megistocarpa, 78 

Anisoptera oblonga Dyer, 100 

Anisoptera polyandra, 119 

Anisoptera scaphula, 75, 152, 153, 155, 156 

Anisoptera scaphula (Roxb.) Pierre, 100 

Anisoptera section, 8, 24, 32 

Anisoptera sp., 157 

Anisoptera thurifera , 160, 134, 135 

Anisoptera thurifera (Blco) Bl., 100 

Anisopteroxylon, 17, 19 

ANNONACEAE, 45 


Antheraea mylitta, 194 

Antherotriche section, 32 

Anthoshorea Heim, 14, 33 

Anthoshorea section, 8, 32 

Anthoshorea sub-genus, 8 

Anthoshorea, 8, 15, 16, 22, 24, 28, 29, 31, 32, 33 

ANTHOSHORINAE sub-tribe, 31, 32 

Apis spp, 48, 50 

Araucaria columnaris, 61 

Araucaria hunsteinii, 61, 81, 83 

ARAUCARIACEAE, 61, 81 

ASCOMYCETES, 122, 103 

Aspergillus niger, 120 

Aspergillus, 120 

Asterina, 121 

Astraeus hygrometricus, 104 

Auriculatae section, 32 

Auriculatae sub-section, 8, 32 

Aurificaria shoreae, 121 

B 

Bactronophorus, 119 

Baillonodendron , 32 

Balanocarpus Bedd., 33 

Balanocarpus heimii, 8, 32 

Balanocarpus Kosterm., 15, 15 

Balanocarpus, 8, 32, 33, 59, 101 

‘Balau’ group of Shorea , 59 

Barbata section, 32 

Barbata sub-section, 8, 32 

Barbatae section, 8 

Barringtonia sp., 167 

BASIDIOMYCETEAE, 120, 121, 122 

BASIDIOMYCETES, 103 

Baukia, 119 

‘Beraliya’ group of Shorea , 52


BIXALES, 25 

BOLETACEAE, 104 

Boletus sp., 106 

Boletus, 99, 104, 158 

BOMBACACEAE, 26 

Bombax, 26 

BOSTRICHIDAE, 119, 123 

Botriodiplodia theobromae , 123 

Brachypterae section, 8, 15, 24, 32 

Brachypterae sub-section, 8, 32 

Bracteata section, 32 

Buchanania lanzan, 165 

BUPRESTIDAE, 123 

C 

Calicites alatus, 19 

Calicites obovatus, 19 

Callosciurus notatus, 116 

Callosciurus prevostii, 116 

Calophyllum, 91 

Camphora officinalis, 193 

Capnodium, 121 

Cenococcum, 104 

Cephaleuros virescens, 115, 121 

CERAMBYCIDAE, 115, 117, 119, 123 

Ceratocystis spp., 123 

Cercospora, 121 

Chaetomium globosum, 122 

Chukrassia tabularis, 151 

CICALIDAE, 118 

Ciliata section, 32 

Cinnamomum camphora, 192, 193 

Cladosporium chlorocephalum, 120 

Cladosporium cladiosporioides, 120 

COCCIDAE, 115, 118 

Coleoptera, 48, 117 

Colletotrichum, 120, 121 

COLYTIDAE, 116 

Coniophora cerebella, 122 

Coptotermes curvignathus, 119 

Coriolus versicolor, 122 

Corticium, 121 

Cotylelobium , 6, 8, 14, 16, 24, 26, 28, 29, 30, 32, 33, 

100 

Cotylelobium burckii, 58, 62, 64, 79 

Cotylelobium malayanum Sloot.Pen. , 100 

Cotylelobium melanoxylon, 58, 62, 64, 79 

Cotylelobium Pierre, 14, 28 

Cotylelobium scabriusculum (Thw.) Brandis, 16 

Cotylelobium scabriusculum Brandis, 100 

Cryptotermes cynocephalus, 119 

CURCULIONIDAE, 116 

Curvularia harveyi, 122 

Curvularia, 120 

Cylindrocladium, 120 

200 

D 

Daedalea quercina, 122 

DEUTEROMYCETES, 120, 121 

Dialeges pauper, 119 

Dicyathifer, 119 

DILLENIALES, 25 

DILLENIFLORAE, 25 

Diplodia spp., 123 

Diptera, 48 

DIPTEROCARPACEAE sensu lato, 5, 9, 10, 14, 26, 27, 35 

DIPTEROCARPACEAE sensu stricto, 5, 26 

DIPTEROCARPACEAE, 5, 6, 7, 8, 16, 17, 22, 23, 25, 26, 

27, 28, 29, 30, 31, 33, 35, 45, 46, 49, 83, 89, 99, 

103, 115, 117, 133 

DIPTEROCARPAE tribe, 31, 32 

DIPTEROCARPALES, 31 

DIPTEROCARPI tribe, 27, 28, 30, 32, 34 

DIPTEROCARPINAE sub-tribe, 31, 32, 33 

DIPTEROCARPOIDEAE, 5, 7, 9, 10, 11, 15, 26, 30, 133 

Dipterocarpophyllum, 19 

Dipterocarpoxylon, 17, 19 

Dipterocarpus alatus Roxb., 100 

Dipterocarpus alatus, 46, 58, 60, 61, 63, 64, 67, 78, 

83, 104, 151, 152, 160, 161, 170, 188, 189, 195, 

196 

Dipterocarpus Baudii Korth., 100 

Dipterocarpus baudii, 49, 64, 75, 152, 153, 156, 

158, 160 

Dipterocarpus bourdilloni, 151, 191 

Dipterocarpus caudatus ssp. penangianus, 78 

Dipterocarpus caudatus, 63 

Dipterocarpus caudiferus, 159 

Dipterocarpus chartaceus Sym., 100 

Dipterocarpus chartaceus, 60, 78 

Dipterocarpus confertus Sloot., 100 

Dipterocarpus confertus, 106 

Dipterocarpus cornutus Dyer, 100 

Dipterocarpus cornutus, 49, 78, 82, 116, 188, 190 

Dipterocarpus costatus Gaertn. f., 100 

Dipterocarpus costatus, 58, 60, 62, 78, 151, 152, 

188, 191


Dipterocarpus costulatus Sloot., 100 

Dipterocarpus costulatus, 49, 75, 153, 156 

Dipterocarpus crinitus, 188, 190 

Dipterocarpus dyeri, 152 

Dipterocarpus elongatus Korth., 100 

Dipterocarpus Gaertn.f., 5 

Dipterocarpus gracilis Bl., 100 

Dipterocarpus gracilis, 78, 157, 191 

Dipterocarpus grandiflorus (Blco) Blco, 100 

Dipterocarpus grandiflorus, 60, 64, 78, 151, 153, 

154, 156, 159, 160, 163, 169, 188, 190, 191, 195 

Dipterocarpus hasseltii Bl., 100 

Dipterocarpus hasseltii, 188, 190 

Dipterocarpus hispidus Thw., 100 

Dipterocarpus hispidus, 191 

Dipterocarpus humeratus Sloot., 100 

Dipterocarpus humeratus, 64, 78 

Dipterocarpus indicus Bedd., 100 

Dipterocarpus indicus, 119, 151, 170, 191, 195 

Dipterocarpus intricatus Dyer., 100 

Dipterocarpus intricatus x tuberculatus, 78 

Dipterocarpus intricatus, 60, 61, 63, 64, 67, 68, 79, 

80, 83, 134, 152, 160 

Dipterocarpus kerrii, 75, 151, 153, 156, 188, 190 

Dipterocarpus kunstleri King, 100 

Dipterocarpus kunstleri, 78, 105 

Dipterocarpus macrocarpus, 119, 151, 152, 158, 

188, 191 

Dipterocarpus oblongifolius Bl., 100 

Dipterocarpus oblongifolius, 47, 81 

Dipterocarpus obtusifolius Teysm. ex Miq., 100 

Dipterocarpus obtusifolius, 58, 60, 61, 62, 64, 68, 

78, 134, 152, 191 

Dipterocarpus pilosus, 152 

Dipterocarpus retusus, 153, 159 

Dipterocarpus rotundifolius, 11 

Dipterocarpus sp., 118, 119 

Dipterocarpus spp., 118 

Dipterocarpus spp., 153, 160, 170 

Dipterocarpus sublamellatus Foxw., 100 

Dipterocarpus tempehes Sloot., 100 

Dipterocarpus tempehes, 153 

Dipterocarpus tuberculatus Roxb., 100 

Dipterocarpus tuberculatus var. grandifolius, 78 

Dipterocarpus tuberculatus var. turbinatus, 46 

Dipterocarpus tuberculatus, 30, 46, 58, 59, 60, 63, 

64, 68, 78, 118, 134, 152, 188, 189, 194, 195, 196 

Dipterocarpus turbinatus, 58, 62, 64, 68, 78, 81, 

119, 151, 152, 160, 170, 172, 188, 189, 195 

201 

Dipterocarpus verrucosus Foxw., 100 

Dipterocarpus verrucosus, 156 

Dipterocarpus warburgii, 154 

Dipterocarpus zeylanicus Thw., 100 

Dipterocarpus zeylanicus, 58, 62, 64, 78 

Dipterocarpus, 8, 14, 15, 16, 19, 22, 23, 24, 26, 27, 

28, 29, 30, 31, 32, 33, 34, 45, 46, 49, 52, 57, 59, 

61, 67, 68, 75, 79, 80, 82, 91, 100, 135, 159, 188, 

195 

Dirochloa spp., 165 

Doona section, 8, 32 

Doona zeylanica, 26 

Doona, 8 , 14, 15, 16, 21, 24, 31, 32, 33, 34 

Drosicha stebbingi, 115 

Dryobalanoides section, 8, 15, 32 

Dryobalanoides sub-section, 8, 32 

DRYOBALANOINAE sub-tribe, 31, 32 

Dryobalanops aromatica Gaertn. f., 100 

Dryobalanops aromatica, 48, 49, 58, 63, 64, 75, 82, 

83, 105, 116, 117, 146, 152, 153, 154, 155, 156, 

157, 158, 159, 161, 163, 164, 166, 169, 187, 191, 

192, 193, 195 

Dryobalanops beccarii, 192 

Dryobalanops keithii Sym., 100 

Dryobalanops keithii, 60, 62, 64, 78 

Dryobalanops lanceolata Burck., 100 

Dryobalanops lanceolata, 46, 48, 47, 62, 64, 78, 

105, 153, 156, 157, 159, 160, 163 

Dryobalanops oblongifolia Dyer, 100 

Dryobalanops oblongifolia, 75, 82, 105, 152, 155, 

156, 158, 159, 162, 164, 169, 170 

Dryobalanops oocarpa Sloot., 100 

Dryobalanops rappa, 79 

Dryobalanops spp., 171 

Dryobalanops, 8, 15, 16, 17, 19, 24, 26, 27, 28, 29, 

30, 31, 32, 33, 34, 45, 47, 59, 75, 80, 82, 91, 100, 

159, 188, 195 

Dryobalanoxylon, 17, 19 

Durio, 91 

Duvallelia, 32 

Dyerella, 32 

E 

Ehretia laevis, 165 

ELAEOCARPACEAE, 9, 10 

Elaeocarpus stipularis, 167 

Endogone, 103 

Endospermum malaccense, 167


EPHEMEROPTERA, 119 

EUCOSMIDAE, 117 

Eugeissona triste, 161, 165, 169 

Eugenia, 91 

Euhopea section, 32 

EUPHORBIACEAE, 45 

Eushorea section, 32 

Eushorea sub-genus, 8 

Eustemonoporus section, 32 

Euvatica section, 32 

F 

FAGACEAE, 95 

Ficus, 91 

Fragraea fragrans, 157, 161 

FUNGI IMPERFECTI, 120, 122 

Furcraea, 194 

Fusarium, 120 

G 

Gaertnera, 91 

Garcinia indica, 190 

Garcinia, 91 

GEOMETRIDAE, 117 

Girroniera nervosa, 167 

Glabrae section, 8, 32 

Gleichenia spp., 161 

Gliocladium, 120, 124 

Gossypium, 26 

Greenia jackii, 168 

GUTTIFERAE, 9, 10, 25, 26 

GUTTIFERALES, 25 

H 

Hancea section, 32 

Hemiphractum section, 32 

HEMIPTERA, 92 

Herpes simplex, 192 

Heterocerambyx spinicornis, 123 

Hopea , 5, 8, 14, 16, 23, 24, 26, 28, 29, 30, 31, 32, 

33, 34, 45, 46, 47, 50, 52, 59, 67, 80, 82, 101, 

151, 159, 191, 195 

Hopea bancana (Boerl.) Sloot., 101 

Hopea bancana, 153 

Hopea beccariana, 46, 105, 160, 192 

Hopea celebica, 192 

Hopea dryobalanoides Miq., 101 

202 

Hopea dryobalanoides, 79, 192 

Hopea ferrea Laness., 101 

Hopea ferrea, 60, 62, 64, 79, 80, 152 

Hopea ferruginea Parijs, 101 

Hopea foxworthyi, 58, 60, 62, 64, 79, 117, 160 

Hopea glabra, 47, 151 

Hopea hainanensis, 59, 63, 64, 66, 13, 81, 118 

Hopea helferi, 60, 63, 64, 82, 104, 106, 160 

Hopea iriana Sloot., 101 

Hopea jucunda Thw., 101 

Hopea latifolia, 49 

Hopea mengerawan Miq., 101 

Hopea mengerawan, 62, 64, 79, 104, 153 

Hopea micrantha, 192 

Hopea montana Sym., 101 

Hopea nervosa King, 101 

Hopea nervosa, 64, 103, 105, 116, 153, 160 

Hopea nigra, 79 

Hopea nudiformis Thw., 101 

Hopea nutans, 170 

Hopea odorata Roxb., 101 

Hopea odorata, 30, 46, 49, 50, 57, 58, 60, 62, 64, 79, 

82, 83, 104, 105, 106, 118, 120, 152, 153, 155, 

156, 157, 158, 161, 192, 194, 195, 196 

Hopea parviflora, 58, 65, 79, 119, 122, 151, 170, 

172, 193, 196 

Hopea parvifolia (Warb.) Sloot., 101 

Hopea pierrei, 157, 159 

Hopea plagata (Blco) Vidal, 101 

Hopea plagata, 160 

Hopea sangal Korth., 101 

Hopea sangal, 153 

Dryobalanoides section, 15 

Hopea section, 8, 15, 22, 32 

Hopeae section, 14, 15 

Hopea spp., 151 

Hopea sub-section, 8, 32 

Hopea subalata, 46, 49, 65 

Hopea utilis, 151 

Hopea wightiana, 65, 151, 170, 172 

HOPEAE tribe, 31, 32 

Hopenium, 17, 19 

Hopeoides section, 32 

Hoplocerambyx spinicornis, 115, 117, 118, 119 

HYMENOMYCETES, 121 

HYMENOPTERA, 92 

Hypoxylon mediterraneum, 122


I 

IMBRICATE group, 27, 28, 31, 32, 34 

Imperata cylindrica , 161, 165 

Indigofera teysmanii, 161 

Isauxis section, 32 

Isauxis sub-genus, 8 

Isoptera section, 32 

L 

Lacifer lacca, 118, 194 

Lactarius, 104 

Lantana camara, 165 

Lasiodiplodia theobromae, 123 

Lasiodiplodia, 120 

LAURACEAE, 17, 45 

LEGUMINOSAE, 99 

LEPIDOPTERA, 116, 117 

Lepiota, 104 

Limnoria, 119 

LOPHIRACEAE, 25 

LORANTHACEAE, 121 

Loranthus, 122 

LYCTIDAE, 119 

Lyctus brunneus, 119 

Lymantria, 117 

Lymantria mathura, 117 

Lymantriidae, 117 

M 

Macaranga spp., 167 

Macaranga, 91 

MAGNOLIALES, 25 

Mallotus philippinensis, 165 

MALVACEAE, 26 

MALVALES, 25, 26 

Mangifera, 91 

Marasmius, 121 

Marquesia , 6, 7, 8, 11, 12, 14, 15, 17, 23, 24, 26, 27, 

29, 30, 31, 33, 101 

Marquesia acuminata Gilg., 101 

Marquesia excelsa, 11, 17, 23, 24 

Marquesia macroura Gilg., 101 

Martesia sp., 119 

Martesia, 119 

Massicus venustus, 118 

Maximae section, 8, 32 

Meliaceae, 67 

Meliococcus, 81 

Meliola sp., 121 

Mesua ferrea, 168 

Michelia champaca, 151 

Microcarpae section, 32 

Microcerotermes cameroni, 119 

Microcydus ulei, 115 

Mikania scandens, 165 

Mikania spp., 165 

Miliusa velutina, 165 

Monoporandra section, 32 

MONOTACEAE, 25, 31, 33 

Monotes , 6, 7, 8, 11, 12, 14, 15, 16, 17, 20, 24, 25, 

27, 29, 30, 31, 33, 101 

Monotes africanus (Welw.) A.D.C., 101 

Monotes elegans Gilg., 101 

Monotes kerstingii, 14, 20, 21, 65, 79 

Monotes madagascariensis, 14 

Monotes oeningensis (Heer) Weyland, 16 

Monotes oeningensis, 19 

Monotoideae, 6, 7, 8, 9, 10, 14, 16, 25, 26, 27, 30, 

133, 

Mutica section, 8, 32 

Mutica sub-section, 8, 32 

Mutica, 28 

Muticae section, 8, 32 

Muticae sub-section, 8 

Muticae, 29 

MYRTACEAE, 45, 81 

203 

N 

Nanophyes shoreae, 116 

Nanophyes, 116 

Nausitora, 119 

Neobalanocarpus heimii (King) Ashton, 101 

Neobalanocarpus heimii, 31, 32, 49, 65, 116, 119, 

137, 152, 157, 158, 160, 169, 192 

Neobalanocarpus, 24, 26, 28, 30, 31, 33, 45, 59, 191 

Neobalanocarpus, 101 

Neohopea section, 8, 15, 32 

Neohopeae section, 8 

NOCTUIDAE, 117 

Nototeredo, 119 

O 

OCHNACEAE, 9, 10, 26 

OCHNALES, 25 

Oecophylla smaragdina, 117 

Ougeineia oojeinensis, 165


Ovalis section, 8, 32 

Ovalis sub-group, 8, 32 

Ovalis, 24 

Ovoides section, 32 

P 

Pachycarpae section, 8, 15, 32 

Pachynocarpoides section, 32 

Pachynocarpus section, 8, 32 

Pachynocarpus sub-genus, 8 

Pachynocarpus, 16, 24, 29, 32, 33 

Paenoe section, 32 

Pakaraimaea , 6, 7, 8, 11, 14, 15, 17, 23, 24, 25, 26, 

31, 33 

Pakaraimaea dipterocarpacea ssp. dipterocarpacea, 

14 

Pakaraimaea dipterocarpacea ssp. nitida, 14 

Pakaraimaea dipterocarpacea, 11, 14, 25 

Pakaraimoideae, 5, 7, 9, 10, 14, 25, 27, 30, 133 

PALMAE, 69 

Pammene theristhis, 117 

Parahopea, 32 

Paraserianthes falcataria, 153, 154, 161, 164 

Parashorea chinensis, 170 

Parashorea densiflora Sloot. & Sym., 101 

Parashorea densiflora, 49, 65 

Parashorea lucida (Miq.) Kurz., 101 

Parashorea malaanonan (Blco) Merr., 101 

Parashorea malaanonan, 60, 62, 65, 78, 118, 119, 

153, 156, 157, 159 

Parashorea plicata, 154, 158, 159 

Parashorea robusta, 118 

Parashorea smythiesii, 58, 62, 65, 78, 80 

Parashorea stellata, 118, 191 

Parashorea tomentella, 58, 60, 62, 65, 78, 119, 159 

Parashorea, 8, 15, 23, 24, 27, 28, 29, 30, 31, 32, 33, 

34, 45, 59, 80, 101, 159, 188 

PARASHORINAE sub-tribe, 31, 32 

Paropsia varediformis, 167 

Parvifolia sub-group, 8, 32 

Pasania sp., 167 

Pauciflora sub-group, 8, 32 

Peltophorum spp., 161 

Penicillium albicans, 120 

Penicillium canadense, 120 

Penicillium, 120 

Pentacme contorta (Vidal) Merr. & Rolfe, 101 

Pentacme section, 8, 32 

Pentacme siamensis (Miq.) Kurz., 101 

Pentacme suavis, 118 

Pentacme, 5, 8, 14, 15, 24, 31, 32, 33, 34, 101 

Pentacmoxylon, 19 

Pestaliopsis, 120 

Pestalotia, 120 

Petalandra section, 32 

Petalandra, 5 

Phellinus caryophylli, 123 

Phellinus fastuosus, 123 

Pierrea , 32 

Pierrea section, 32 

Pierrea sub-section, 8 

Pilosae section, 8, 32 

Pinanga sub-group, 8, 32 

Piper longum, 190 

Pisolithus , 103, 104 

Pisolithus tinctorius, 104, 106 

PLATYPODIDAE, 119 

Plicati section, 32 

Polyandrae sub-section, 8, 32 

Polyporus shoreae, 121 

Polystictus versicolor, 122 

Presbytis rubicunda, 116 

Pseudomonotes , 7, 8, 11, 14, 23, 24, 26 

Pseudomonotes tropenbosii, 6, 11, 14, 33 

Psittacula sp., 116 

Psychotria, 91 

PYRALIDAE, 117 

Q 

Quercus lucida, 167 

R 

Randia anisophylla, 168 

Randia scortechenii, 167, 168 

‘Red Meranti’ group of Shorea , 15, 32, 59, 

Retinodendron genus, 32 

Rhizopus oryzae, 120 

Rhodophyllus sp., 105 

Richetia Heim, 33 

Richetia sub-genus, 8, 32 

Richetia, 8, 28, 31, 32, 33 

Richetiodes sub-section, 8, 32 

Richetioides section, 8, 15, 24, 29, 32, 34, 47 

RUBIACEAE, 45 

Rubroshorea sub-genus, 8, 32 

204


Rubroshorea, 8, 31, 32, 33 

Rubroshoreae section, 15 

Rugosae section, 32 

Russula amatic, 155 

Russula sp., 106 

Russula, 104, 158 

RUSSULACEAE, 104 

S 

SAPINDACEAE, 81 

SAPOTACEAE, 69 

SARCOLAENACEAE, 9, 10, 25 

Schyzophyllum commune, 120, 121, 122 

Scleroderma columnare, 104, 106, 155 

Scleroderma dicstyosporum, 106 

Scleroderma sp., 106 

Scleroderma spp., 158 

Scleroderma, 104 

SCLERODERMATACEAE, 104 

Sclerotium, 120 

SCOLYTIDAE, 119, 123 

Scorodocarpus borneensis, 168 

171 

Shorea , 6, 8, 11, 13, 14, 16, 23, 24, 26, 28, 29, 30, 

31, 32, 33, 34, 45, 46, 47, 50, 59, 61, 67, 68, 75, 

80, 82, 91, 92, 93, 94, 101, 117, 122, 140, 159, 

170, 188, 191, 193, 195 

Shorea academia , 101 

Shorea acuminata Dyer, 101 

Shorea acuminata, 47, 63, 65, 75, 82, 117, 152, 153, 

155, 156 

Shorea affinis (Thw.) Ashton, 101 

Shorea affinis, 58, 60, 62, 65, 79 

Shorea agamii ssp agamii, 49 

Shorea albida, 49 

Shorea almon, 57, 58, 62, 65, 68, 78, 154, 156 

Shorea amplexicaulis, 58, 60, 62, 65, 78 

Shorea aptera, 193, 195 

Shorea argentifolia, 49, 58, 60, 62, 63, 65, 78, 153 

Shorea assamica Dyer Pen. , 101 

Shorea assamica ssp. globifera, 192 

Shorea assamica, 65, 118, 151, 160 

Shorea balangeran (Korth.) Burck, 101 

Shorea beccariana, 78 

Shorea bracteolata Dyer Pen. , 102 

Shorea bracteolata, 16, 65, 75, 82, 105, 156 

Shorea compressa Burck, 102 

Shorea compressa, 106, 158 

205 

Shorea congestiflora, 47, 62, 65, 78 

Shorea contorta, 58, 65, 156, 158, 160, 161, 164 

Shorea cordifolia, 47 

Shorea crassifolia, 192 

Shorea curtisii Dyer ex King, 102 

Shorea curtisii, 29, 49, 65, 75, 82, 105, 106, 123, 

152, 156, 158, 160, 165, 169 

Shorea dasyphylla Foxw., 102 

Shorea dasyphylla, 65 

Shorea disticha, 47 

Shorea elliptica, 122 

Shorea faguetiana Heim, 102 

Shorea faguetiana, 79 

Shorea fallax, 60, 62, 65, 78, 80, 156 

Shorea ferruginea, 58, 60, 62, 65, 78 

Shorea foxworthyi Sym., 102 

Shorea foxworthyi, 160 

Shorea Gaertn., 33 

Shorea gibbosa, 60, 63, 78 

Shorea glauca King, 102 

Shorea glauca, 120 

Shorea gratissima, 49 

Shorea guiso (Blco) Bl., 102 

Shorea guiso, 58, 79, 118, 122, 153, 154, 156 

Shorea hemsleyana, 47 

Shorea henryana Pierre, 102 

Shorea henryana, 152 

Shorea hopeifolia, 163 

Shorea hypochra Hance, 102 

Shorea hypochra, 65 

Shorea hypoleuca, 122 

Shorea javanica K & V., 102 

Shorea javanica, 65, 117, 121, 122, 170, 173, 192 

Shorea johorensis Foxw., 102 

Shorea johorensis, 153, 157, 162, 163 

Shorea laevifolia, 119 

Shorea laevis Ridl., 102 

Shorea laevis, 79, 116, 122, 156 

Shorea lamellata Foxw., 102 

Shorea lamellata, 192 

Shorea lepidota (Korth.) Bl., 102 

Shorea lepidota, 47, 63, 153 

Shorea leprosula Miq., 102 

Shorea leprosula, 47, 49, 50, 58, 62, 65, 75, 79, 82, 

103, 104, 105, 106, 117, 119, 120, 152, 153, 154, 

155, 156, 157, 158, 160, 161, 162, 163, 167, 168, 

169, 171, 172


Shorea leptoderma, 60, 62, 79 

Shorea longisperma, 82 

Shorea macrophylla (de Vriese) Ashton, 102 

Shorea macrophylla, 49, 60, 62, 65, 75, 78, 82, 116, 

140, 152, 153, 155, 156, 158, 169, 170, 171, 173, 

193, 195 

Shorea macroptera Sloot., 102 

Shorea macroptera ssp. baillonii, 29 

Shorea macroptera ssp. macropterifolia, 29 

Shorea macroptera, 47, 49, 60, 62, 75, 78, 82, 105, 

116, 152, 153, 154, 155, 156, 158, 160, 169 

Shorea maxima, 47 

Shorea maxwelliana King, 102 

Shorea maxwelliana, 82, 119 

Shorea mecistopteryx Ridl., 102 

Shorea mecistopteryx, 106, 153, 193 

Shorea megistophylla, 47, 48, 50, 51, 191 

Shorea multiflora, 47, 153, 158, 169 

Shorea negrosensis, 154 

Shorea obstusa Wall., 102 

Shorea obtusa, 62, 65, 79, 118, 122, 134, 152, 160, 

191, 194, 195 

Shorea ovalis (Korth.) Bl., 102 

Shorea ovalis ssp sericea, 49 

Shorea ovalis ssp. sericea, 30 

Shorea ovalis, 30, 47, 62, 65, 75, 78, 82, 105, 116, 

117, 119, 120, 152, 153, 155, 156, 157, 158, 160, 

162 

Shorea ovata Dyer ex Brandis, 102 

Shorea ovata, 79 

Shorea pachyphylla, 63, 65 

Shorea palembanica Miq., 102 

Shorea palembanica, 78, 104, 153, 160 

Shorea parvifolia Dyer Pen., 102 

Shorea parvifolia, 49, 58, 60, 61, 62, 66, 75, 78, 82, 

105, 106, 117, 152, 153, 154, 155, 156, 157, 158, 

159, 160, 161, 162, 163, 167, 169, 172 

Shorea parvistipulata, 163 

Shorea pauciflora King, 102 

Shorea pauciflora, 49, 66, 82, 116, 153, 156, 157, 

160 

Pinanga group of Shorea, 153, 170, 171, 172 

Shorea pinanga Scheff., 102 

Shorea pinanga, 58, 66, 78, 106, 122, 153, 155, 156, 

158, 160, 163, 168 

Shorea platyclados Sloot. ex Foxw., 102 

Shorea platyclados, 66, 75, 80, 82, 152, 153, 155, 

156, 159, 172 

206 

Shorea polyandra Ashton, 102 

Shorea polyandra, 118 

Shorea polysperma, 154, 156, 162 

Shorea quadrinervis, 106 

Shorea resinosa, 16, 49 

Shorea retinodes, 192 

Shorea robusta Gaertn. f., 102 

Shorea robusta, 6, 57, 58, 59, 62, 66, 68, 78, 80, 106, 

115, 117, 118, 119, 120, 121, 122, 123, 133, 151, 

152, 154, 155, 156, 157, 158, 160, 161, 162, 164, 

165, 166, 169, 170, 171, 172, 187, 189, 190, 192, 

193, 194, 195, 196 

Shorea roxburghii G. Don, 102 

Shorea roxburghii, 15, 16, 57, 58, 59, 60, 62, 66, 68, 

78, 80, 81, 105, 151, 152, 160, 191, 194 

Shorea scabrida Sym., 102 

Anthoshorea section, 14, 15, 26, 33, 34, 59 

Brachyptera section, 14 

Doona section, 47, 48, 52, 93 

Muticae section, 47 

Ovales section, 47 

Ovalis section, 34 

Pachycarpae section, 47, 59 

Richetioides section, 34, 47 

Rubella section, 8, 32 

Shoreae section, 8, 34 

Brachypterae section, 59 

Mutica section, 48, 59 

Rubellae section, 8, 15, 24, 32, 34 

Shorea section, 8, 14, 15, 23, 24, 32 

Shorea selanica (Lamk.) Bl., 102 

Shorea selanica, 63, 78, 104, 153, 156, 158, 159, 

160 

Shorea seminis (de Vriese) Sloot., 102 

Shorea seminis, 153 

Shorea sericeiflora Fisher & Hance, 102 

Shorea siamensis Miq., 103 

Shorea siamensis, 58, 59, 60, 63, 66, 78, 118, 122, 

134, 152, 189, 191 

Shorea singkawang, 63 

Shorea smithiana Sym., 103 

Shorea smithiana, 49, 58, 66, 78, 116, 153, 156 

Shorea sp., 119 

Shorea splendida, 47, 172 

Shorea spp., 118, 121, 139, 146, 160, 167, 169, 170, 

171 

Shorea squamata, 154, 156 

Shorea stenoptera Burck, 103


Shorea stenoptera, 47, 104, 106, 118, 156, 158, 164, 

168, 171, 193, 195 

Shorea stipularis, 26 

Shorea sub-genus, 8, 32 

Shorea sub-section, 8, 23, 32 

Shorea sumatrana (Sloot. ex Thor.), 103 

Shorea sumatrana, 63, 66, 140, 152, 160, 164 

Shorea talura Roxb., 103 

Shorea talura, 15, 16, 80, 118, 151, 157, 160, 163, 

194 

Shorea teysmanniana Dyer ex Brandis, 103 

Shorea teysmanniana, 117, 159, 163 

Shorea thorelii, 152 

Shorea trapezifolia, 47, 49, 50, 51, 62, 66, 78, 146, 

169 

Shorea tumbuggaia, 151, 191 

Shorea virescens, 192 

Shorea Wiesneri, 192 

Shorea xanthophylla, 63 

Shorea zeylanica, 26 

SHOREAE tribe, 27, 28, 30, 31, 32, 34 

Shoreoxylon, 17, 19 

SHORINAE sub-tribe, 31, 32 

Smithiana sub-group, 8, 32 

Smithiana sub-section, 32 

Smithianeae sub section, 8, 32 

Sphaerae section, 32 

Sphaerales section, 32 

Sphaerocarpae section, 32 

Sphaerocarpae sub-section, 8, 32 

Stemonoporinae sub-tribe, 31, 32 

Stemonoporus affinis, 29 

Stemonoporus canaliculatus, 60, 63, 66, 78 

Stemonoporus , 8, 15, 32 

Stemonoporus oblongifolius, 47, 50, 51, 52 

Stemonoporus reticulatus, 29 

Stemonoporus Thw., 28 

Stemonoporus, 11, 14, 20, 21, 23, 24, 27, 28, 30, 31, 

33, 34, 47, 49, 50 

STERCULIACEAE, 26 

STYRACACEAE, 192 

Styrax benzoin, 192 

Sunaptea , 8, 14, 15, 16, 23, 24, 26, 29, 31, 32, 33, 

34 

Sunaptea scabriuscula (Thw.) Brandis, 16 

Sunaptea section, 8, 32 

Sunaptea type, 16 

Sunapteae section, 32 

SUNAPTINAE sub-tribe, 31, 32 

Sus scrofa, 116, 117 

Swietenia humilis, 67 

Synaptea Griff., 28, 33 

Synaptea section, 32 

Synaptea sub-genus, 8 

T 

Tectona grandis, 152, 160 

Teredo, 119 

TERNSTROEMIACEAE, 25, 26 

THEACEAE, 9, 10, 25, 26 

THEALES, 25 

Theobroma, 25 

Thespesia, 25 

Tilia, 25 

TILIACEAE, 9, 10, 25, 26 

TILIALES, 26 

Tomentellae section, 32 

TORTRICIDAE, 117 

Trametes versicolor, 122 

Trema ambionensis, 167 

Trema, 91 

TRENTEPHOLIACEAE, 121 

Trichoderma, 124 

Tricholoma, 104 

Trigona spp, 48 

Tuberculati section, 32 

Tyromyces palustris, 122 

U 

Upuna borneensis, 121 

Upuna section, 32 

Upuna, 8, 14, 15, 16, 19, 20, 21, 24, 26, 28, 29, 30, 

31, 33, 34, 45 

UPUNINAE sub-tribe, 31, 32 

V 

VALVATE group, 27, 28, 31, 32, 33 

Vateria copallifera, 29, 48, 194, 195, 196 

Vateria indica L., 103 

Vateria indica, 119, 122, 151, 152, 162, 170, 190, 

192, 193, 194, 195, 196 

Vateria L., 33 

Vateria macrocarpa, 151 

Vateria, 5, 8, 14, 15, 20, 21, 23, 24, 27, 28, 29, 30, 

31, 32, 33, 34, 103, 195 

207


VATERINAE sub-tribe, 31, 32 

Vateriopsis seychellarum Heim, 103 

Vateriopsis seychellarum, 12 

Vateriopsis, 8, 15, 20, 21, 23, 24, 28, 30, 31, 32, 33, 

103 

Vaterioxylon, 17, 19 

Vatica , 6, 8, 14, 15, 16, 20, 23, 24, 26, 27, 28, 30, 33, 

34, 45, 59, 103, 188, 195 

Vatica astrotricha, 118 

Vatica chartacea Ashton, 103 

Vatica chinensis, 191 

Vatica heteroptera, 16 

Vatica Kosterm., 15 

Vatica lanceaefolia, 191, 194, 195, 152 

Vatica mangachapoi, 58, 62, 66, 79 

Vatica nitens, 169 

Vatica oblongifolia ssp. crassibolata, 29 

Vatica oblongifolia ssp. multinervosa, 29 

Vatica oblongifolia ssp. oblongifolia, 29 

Vatica obscura, 191 

Vatica odorata ssp. odorata, 62, 66 

Vatica odorata ssp.odorata, 79 

Vatica odorata, 26, 152 

Vatica pallida, 49 

Vatica papuana Dyer ex Hemsl., 103 

Vatica pauciflora, 15, 16, 49, 160 

Vatica pro parte, 31, 33, 34 

Vatica rassak (Korth.) Bl., 103 

Vatica rassak, 49, 192 

Vatica roxburghiana, 152 

Pachynocarpus section, 29 

Vaticae section, 29 

Vatica section, 8, 32 

Vatica senlu lato, 33 

Vatica sensu Kostermans, 14 

Vatica sp.1, 103 

Vatica sp., 119 

Vatica sumatrana Sloot., 103 

Vatica sumatrana, 106, 158, 159 

Vatica tumbuggaia, 191 

Vatica umbonata (Hook. f.) Burck, 103 

Vatica umbonata, 16, 49, 63, 66 

Vatica wallichii, 15, 16 

Vaticae tribe, 31, 32 

Vaticae type, 16, 31 

Vaticae, 16 

Vaticinae sub-tribe, 31, 33, 32 

Vaticoxylon, 16, 17, 19 

Vesquella, 32 

W 

‘White Meranti’ group of Shorea, 59 

Woburnia porosa, 16 

Woburnia, 19 

Y 

‘Yellow Meranti’ group of Shorea , 59 

X 

Xyleborus declivigranulatus, 119 

Xyleborus pseudopilifer, 119 

Z 

Zygomycetes, 103 

208

A 

abscisic acid (ABA), 81 

acids, 63, 99, 158, 191 

adaptation trials, 155 

affinities, 14, 25-6 

afforestation, on degraded land, 151, 170 

Africa, 12, 18, 23, 24 

affinities, 25 

distribution, 6, 7, 13, 14, 15, 17, 21 

mycorrhirzas, 101 

agamospermy, 30 

age, 63 

felling, and heart-rot incidence, 123 

flowering and seeding, 168-9 

growth rates, 171-2 

longevity of seeds, 66-7, 68, 73, 81 

nurse crop, 161 

planting stock, 157 

aged planting stock, rejuvenation of, 160 

agriculture, 92 

see also plantations 

agroforestry, 170, 192 

Aided Natural Regeneration, 151 

airlayering, 160 

airtight containers, seed storage in, 80-1 

algae, 115, 121 

aldrex, 119 

alkaloides, in resin, 116 

altitudinal zonation, 11, 12 

ambrosia beetles, 119 

America, see South America; United States of America 

anatomy, 29, 33-4, 94-5 

ancestral forms, 17-20 

Andaman Canopy Lifting System, 151 

Andaman Islands, 13, 22, 134, 151 

oleoresins, 188, 189, 191 

aneuploid series, 46 

Angola, 14 

animals, destruction by, 116, 117, 157 

anthers, 7, 9, 11, 24 

General Index 

ants, 117 

apomixis, 30, 34-5, 49, 51 

arboricides, 165 

arbuscular mycorrhizas (VAM), 99, 100, 102 

artificial induction of flowering and seeding, 68 

aseasonal areas/forests, 23, 24, 134 

flowering and fruiting, 73, 74-5, 135 

Asia 

affinities, 25-6 

biological characteristics, 23, 26-7 

conservation status, 52 

distribution, 5-6, 7, 12-13, 14-16, 17, 19, 21-2 

ecological presentation, 11, 12 

morphological trends, 24-5 

natural forest management, 133-49 

non-timber forest products, 187-97 

plantation management, 151-85 

supraspecific taxa, 27-8 

see also South Asia; Southeast Asia 

Austria, 16 

autecology, 92 

axis moisture content, 59, 60 

B 

bacterial disease, 115, 121 

bags, storage in, 77, 80-1, 159, 162 

Bali, 134 

ball-rooted transplants, 160 

balsam, 191 

bamboo (climbing), control of, 165 

Bangladesh, 134, 152, 157 

dammars, 192 

diseases, 122 

oleoresins, 188, 189, 190, 191 

pests, 117 

bare-root planting stock, 157-9, 160, 162-3 

wildlings, 158-9159 

bark, 28, 121 

products, 193-4, 196 

barkchipped tapping, 190

General Index 

barus kapur, 192-3 

basket plants (potted seedlings), 157, 160, 161, 162 

Batavian dammar, 192 

Bavistin, 120 

beaches, 170 

bees, 24, 48, 50 

beetles, 119 

Bengal, 160, 170 

benlate, 81, 82, 120 

BHC, 119 

bhimsaini-kapur, 192-3 

Bhutan, 117 

biogeography, 5-44 

biological characteristics, 7, 9-11, 23-33, 99-103 

biotic interactions, 93-4 

mycorrhizas, 93, 99-114, 154-5, 158 

birch, 106 

birds, destruction by, 116 

black stain, 123 

blue-stain, 123 

boat wood, 119 

borers, 115, 117-18, 119 

Borneo, 53, 134 

agroforestry, 170 

butter fat, 193 

distribution, 12, 13, 15, 19, 22 

endemic species, 14, 22 

habitats, 12 

Borneo camphor, 192-3, 195 

bostrichid, 119 

botany, 5, 7-11, 23-33 

Brassical, 120 

Brazil, 104 

breeding systems, 46-51 

broadcast sowing, 157, 163-4 

brown ants, 117 

brown rot, 122 

Brunei, 12, 106 

buds, 28, 121, 160 

Burma, 12 

dammars, 192 

distribution, 13, 15, 19, 22 

lac host plants, 194 

natural forests, 133-4 

oleoresins, 188, 189, 191 

pests, 117 

silviculture, 151-2 

burning, see fire 


C 

cacao seeds, 81 

calcium, see mineral nutrition 

calyx, 7, 9, 11, 24, 25, 28 

OLDA seeds, 68 

removal of lobes, 79 

Cambodia, 13, 19, 22, 152 

camphor, 192-3, 195 

cankers, 120-1 

canopies, 90, 91, 92 

crowns and, 23-4 

regeneration and, 169-70 

secondary, planting under, 161 

size of opening, 92 

canvas, collection using, 76 

carbon dioxide and storage life, 81 

‘Carbon Offset’ Project, 141 

carpophores, 120, 122 

caterpillars, 117 

CCA, 119, 122 

Celebes, 22 

cells, 63 

cellulose, 122 

cellulose gums, 194 

chamber storage, 82 

charcoal, storage in, 80 

chemicals, 171 

arboricides, 165 

fungicides, 81, 82, 120 

herbicides, 165 

insecticides, 119 

preservatives, 119, 122-3, 124 

see also fertilisers and fertilisation 

chemotaxonomy, 16, 29 

chilling damage, 58-9, 60 

China, 134 


pests, 118, 119 

reforestation on degraded land, 170 

seed research, 69 

Chittagong, 134 

chlordane, 119 

chromosome variation, 10, 27-8, 29-30, 45-6 

chua oil, 190, 191 

classifications, 5-7, 15-16, 25-33 

forests, 133-4 

stems, for thinning, 166 

clayey sediments, 11 

210

General Index 

cleanings, 164-5, 167 

clearfelling, 137, 161 

Clearfelling System, 137 

climatic conditions, 121, 122, 123 

before seed harvest, 73-4 

climbers (plants), 165 

climbing collection methods, 76-7 

clonal propagation, 159-60 

collar cankers, 120-1 

collar rot, 120 

collection of resins, 188, 189-90, 192 

collection of seeds, 75-7, 83 

Colombia, 11, 12, 14 

compacted soil, 155, 163 

conservation of genetic resources, 45-55, 107 

container plants (potted seedlings), 157, 160, 161, 

162 

containers, seed storage in, 80 

continental drifts, 17-21 

copper-chrome-arsenic, 119, 122 

Coppice System, 136, 143 

coppicing, 135 

cotyledons, 26, 27-8, 29, 31, 194 

diseases, 120 

pests, 116, 117 

crawl tractors, 155 

creosote, 119 

cross-pollination, 46-7 

crowns, 23-4, 168 

architecture, 155 

disengagement regimes, 166, 172 

fertilisation and, 163 

cryopreservation, 82-3 

cultivation, 163, 170 

cuttings, 159 

cylcones, 92 

cytoplasm, 63 

D 

DABATTS, 69 

dammars, 191-2, 195 

damping-off, 120 

death, see mortality 

decay fungi, 122 

deciduousness, 11, 23 

deer browsing, 117 

defoliation, 92, 117, 121 

deforestation, 1, 51, 53 

degraded forests, 141, 151 

see also regeneration 

degraded land/soils, 105, 151, 157, 161, 170 


dessication, 60-3, 68 

partial, 81 

destructive logging, rehabilitation of degraded forest 

sites following, 170 

dhupa, 190 

dhupa tallow, 193 

diameter, see growth 

die-back, 117, 121, 123 

diedrex, 119 

differentiation, 15-16 

dimethyl sulphoxide, 83 

direct sowing, 157, 163-4 

directional felling, 142 

diseases, 115, 119-25, 170-1 

dispersals, 20, 21, 48-9, 89 

distribution, 5-6, 7, 12-25 

disturbance regimes, 92, 171 

Dithane-45, 120 

diversification, 15-16 

diversity, 45-55, 107 

drainage, 123 

drought, 123 

tolerance, 94, 99, 106 

dry deciduous forests, 11 

dry evergreen forests, 11, 133-5, 136 

see also sal forests 

dry habitats, storage in relation to, 68 

drying, see dessication 

E 

East Kalimantan, see Kalimantan 

ecology, 11-12, 89-98 

economic assessments, 168, 172 

economic losses, 115, 122 

see also yield 

ecophysiology, 33-5 

ectomycorrhizas, 93, 99-114 

Egypt, 17 

embryogenesis, 29, 30 

embryos, 27, 30, 31 

diseases, 120 

moisture content, 59, 60 

multiple, 30, 49 

embryo oil content, 60, 67 

211

General Index 

endangered species, identification of, 52 

endemicity, 14, 21-2 

engkabang, 193 

enrichment planting, 141, 161, 164 

establishment of canopy tree species, 92 

establishment of seedlings, 90-4, 135 

establishment of stands, 155-64 

on degraded land, 170 

Ethiopia, 17 

ethylene, and storage life, 81 

eucalyptus, 104 

Europe, 16, 17, 19, 23 

evergreen associations, 11 

evergreen trees/forests, 11-12, 23, 24, 133-6 

flowering and fruiting, 73, 74-5, 135 

everwet areas, 12 

evolutionary systematics, 5-44 

exploitation damage, 141-2 

F 

falls, 92 

families, 5-7, 30-3 

felling, 141-2 

age, with incidence of heart-rot, 123 

thinning, 165-8, 172 

treatment before, 169 

fenpropathrim, 119 

fenvalerate, 119 

feral pigs, destruction by, 116, 117 

fertilisers and fertilisation, 93, 105-6, 107, 163 

effect on growth and mycorrhizal infection, 154-5 

mycorrhizas, 103 

nursery planting stock, 158 

fertility of soil, 93, 170 

fire, 92, 121, 123 

before reforestation/afforestation on degraded 

land, 170 

before seed-fall, 135 

before sowing, 161 

fire-climax woodlands, 23 

fire wood, 194 

‘fishing line’ collection method, 76 

flooding, 92 

flowering, 23, 73, 74-5, 135 

age, 168 

artificial induction, 68 

growth during, 172 

flowers, 7, 9, 11, 23-4, 28 

destruction, 122 

fossils, 16 

sal trees, 194 

forest floor seedling storage, 82 

forest fragmentation, 51, 52-3 

forest management, 1-4 



pests and diseases, 123-4 

plantations, 171-2 

regeneration, 90 

forest products, 118-19, 122-3, 187-97 

see also timber 

fossils, 6, 15, 16-17 

France, 23 

free climbing collection method, 76-7 

freshwater swamps, 12 

frost, 121, 122, 123 

fruiting, 74-5, 135 

age, 168 

canopy tree species, 92 

dry evergreen forests, 134 

fruits, 7, 11, 24, 27 


dessication rates, 61 

dispersal and colonisation by, 20 

fossils, 16 

life span, 73 

pests, 116 

supraspecific taxa, 27, 28 

winged, 20, 21, 24-5, 29 

see also seeds; sepals 

fuelwood production, 166, 194 

funding, 110 

fungi, 93, 99-114, 154-5, 158 

causing diseases, 119-20, 121-3 

fungicides, 81, 82, 120 

G 

Gabon, 11 

galls, 116, 121 

gammexane, 119 

gaps in forests, 92, 93, 171 

gaseous environments for storage, 80-1 

gene dispersal, 50 

gene flow, 52-3 

212

General Index 

genera, 6-7, 8, 13 

chromosome numbers, 45-6 

dessication rates among species of same, 61 

genetic resources, conservation of, 45-55, 107 

geographical distribution, 5-6, 7, 12-25 

geographical origin, 17-22 

Germany, 16 

germination, 57-9, 60, 84, 90, 152 


canopy conditions, 169 

moisture content percentage and, 61 

site preparation, 161-2 

storage conditions, 64-6, 80 

while storing, 60 

while transporting on long journeys, 79 

see also seedlings 

germination inhibitors, storage using, 81 

gibelleric acid, 158 

girdling, 138, 141, 165, 166, 169 

glabrousness, 7, 10, 23 

grafting, 160 

Great Britain, 16, 69 

green moulds, 123 

ground collection, 76 

groundstorey, 90-3 

growth, 139-41 

fertilisation, 105-6, 107, 154-5 

hormone treatment, 159, 160 

mixed forests, 94-5 

plantations, 171-2: on degraded forest land, 170 


site preparation, 161-2 

stand density regimes, 155 

thinning regimes, 166 

see also survival 

gums, 191, 194 

gurjan oil, 188-91 

Guyana, 12, 14 

H 

habitats, 11-12, 51 

mycorrhizas, 101-3 

storage physiology and, 68 

see also evergreen trees/forests 

Hainan, 22, 134 

hairiness, 7, 10, 23 

hairy caterpillars, 117 

hardwoods, 155, 171 

heart-rot fungi, 122 

pests, 119 

harvest, 59-63, 73-4 

harvesting systems, 141-2 

heart-rot, 122, 123, 171 

heartwood borers, 115, 117 

heat stress, 93 

heavy hardwoods, 119 

height, see growth 

helicopter logging, 142 

herbicides, 165 

herbivory, 93-4 

hills, see slopes 

hormone treatment, 159, 160 

humidity, for storage, 82, 159 

Hungary, 16 

hybrydisation, 30, 49 

I 

illipe nuts, 193 

imbibed storage, 80 

imbricate calyx, 25 

‘In oil’, 189 

in vitro experiments, 160 

inbreeding, 51, 53 

India, 18, 133-4, 151-2, 187 


butter (sal) fat, 193 

dammars, 192 

diseases, 119, 120, 121, 122 

distribution, 13, 14, 15, 19, 22 

endangered species, 52 


habitats, 12 


mycorrhizas, 100, 101, 102, 103, 109 

natural regeneration, 134-5 

oleoresins, 188, 189, 190, 191 

pests, 115, 116, 117, 118, 119 

physiological disorders, 123 

reforestation on degraded land, 170 

silviculture, 136-7, 143, 151, 157, 171 

sowing, 163 

taxa for differentiation, 15 

thinning model, 166, 172 

underplanting, 162 

Indian copal, 190 

Indian Irregular Shelterwood System, 136 

Indo-Burma, see Burma; India 

Indochina, 133-4 

213

General Index 

distribution, 15 


oleoresins, 191 

pests, 117 

Indonesia, 15, 153 


diseases, 119, 121, 122 

mycorrhizas, 102, 104, 106, 108 

pests, 117, 118 

resins, 190, 192 

silviculture, 139, 171-2 

vegetative propagation, 159 

see also Borneo; Java; Kalimantan; Sumatra 

Indonesian Selective Cutting System, 139 

induced flowering and seeding, 68 

inflated bags, storage in, 80-1 

inoculation studies, 104-5, 154-5 

insect-borne diseases, 121 

insect pests, 79, 80, 92, 115-19, 170-1 

insecticides, 119 

intermediate (OLDA) seeds, 61, 63-7, 68-9 

Irian Jaya, 134 

Irregular Shelterwood System, 136 

isozyme surveys, 50 

J 

Java, 192 

diseases, 121 

distribution, 19, 22 

fruiting age, 168 

mycorrhizas, 100, 101, 102, 103 


Johore, 192 

K 

Kalimantan, 4, 12 


enrichment planting, 141 

mycorrhizas, 100-3, 104 

pests, 116, 118 

stump plants, 160 

wildings, 159 

kanyin oil, 188, 189 

Katanga, 14 

L 


land clearing, rehabilitation of degraded forest sites 

following, 170 

214 

land use, 1, 53 

landslides, 92 

Laos, 13, 22 

leaching, 99 

leaf damage, 117, 118 

seedlings and saplings, 116-17 

see also defoliation 

leaf gall formation, 115 

leaf venation, 7, 10, 28 

leaves, 7, 10, 11, 28 

dimensions, 94 

diseases, 115, 121 

mineral nutrients in, 105 

products, 194-5 

sal trees, 194 

stripped seedlings, 163 

Lepidoptera, 116, 117 

Lesser Sundas, 22 

liberation thinning, 138, 140, 141 

light, 164 

mycorrhizal inoculum, 105 

re-establishment by natural regeneration, 168 

seedling storage, 82 

seedling survival and growth, 92-3, 94, 161, 162 

light hardwoods, 155, 171 

light-demanding/shade-tolerant seeds, 169 

light-demanding/shade-tolerant species, 90-4, 154 

dry mass allocation, 94 

mycorrhizas, 105, 106 

natural regeneration, 169 

lightning, 92, 171 

lignins, 122, 194 

limestone, 12 

line opening, 164 

line plantings, 161, 162, 165 

litter, 103 

logged over forests, 47, 103, 106 

logging, 51, 52, 141-2 

destructive, rehabilitation of degraded forest site 

following, 170 

residual growth rates after, 140, 141 

logs, 118-19, 122-3 

Lombok, 22, 134 

Long Range Cable Crane System, 142 

longevity of seeds, 66-7, 68, 73, 81 

low light storage conditions, 82 

lowest-safe moisture content (LSMC), 61, 62, 63, 68 

lowland forests, 12, 116, 137 

Luzon, 101

General Index 

M 

Madagascar, 6, 13, 15, 18, 21 

magnesium, see mineral nutrition 

malabar tallow, 193 

Malayan Uniform System (MUS), 2, 137, 138, 143 

Malaysia (Malaya), 1, 134 


dammars, 192 

Departmental Improvement Fellings, 143-4 


distribution, 13, 15, 22 

drought, 123 

economic assessment of plantations, 172 

flowering and fruiting, 75, 168-9 

genetic diversification, 50 

growth and yield, 140-1 

habitats, 12 

light requirement experiments, 154 

mineral nutrients, 106 

mycorrhizas, 100-3, 104, 108 

oleoresins, 189, 190, 191 

pests, 116, 117, 118 

plantation species choice, 156 

planting techniques, 162 

pollen vectors, 48 

seed production, 157 

silviculture, 2, 137-8, 142, 143-4, 171 

sowing, 163 

thinning regimes, 166, 167 


weed vegetation, 165 

wildings, 158-9 

Malesia, 1, 134 

distribution, 13 




see also Borneo; Malaysia; Moluccas; New 

Guinea; Philippines; Sulawesi; Sumatra 

mammals, destruction by, 116, 117, 157 

marine borers, 119 

marine toredo worm, 122 

mating systems, 46-8 

maturation of seeds, 59-63 

mealybugs, 115 

mechanisation, 141-2, 155 

membranes, 63 

microclimates, 92 

215 

migration, 17, 21 

between populations, 50 

mineral nutrition, 93, 99, 105-6, 107 

effect on growth and mycorrhizal infection, 154-5 

nursery planting stock, 158 


mining, rehabilitation of degraded forest sites 

following, 157, 170 

minyak keruing, 188 

mistlotoes, 124 


Modified Malayan Uniform System, 137, 141 

moisture content, 59-63 

storage conditions, 64-6, 80, 81 

transportion conditions, 77-9 

Moluccas, 13, 22, 134 

monkeys, destruction by, 116 

monocyclic (shelterwood) systems, 135, 136-7, 143- 

4, 167-8, 169-70 

monospecific plantations, 115 

morphology, 23-5, 29, 33-5, 104 

mortality 

caused by diseases, 120-1 

caused by pests, 116, 117, 118, 119 

outplants, 103 



see also survival 

moulds, 123 

Mozambique, 14 

mud-puddled wildings, 163 

mulching, 163 

Myammar, see Burma 

Mycorrhiza Network Asia, 109-10 

mycorrhizas, 93, 99-114, 154-5, 158 

N 

natural disturbance regimes, 92, 171 

natural forests, management of, 133-49 

natural regeneration, 134-5, 168-70 

nematodes, 117 

Nepal, 19, 117, 152, 190 

nets, collection using, 76 

New Guinea, 13, 14, 15, 22 

nitrogen environment, storage in, 81 

see also mineral nutrition 


notch-planting, 162-3

General Index 

nurse crops, 161, 165 

nursery planting stock, 156-63 

‘nursing’ phenomenon, 107 

nutrition, see mineral nutrition 

nuts, see fruits 

O 

oak, 106 

oil content of seeds, 60, 67, 

oleoresins, 188-91, 195 

ontogenesis, 29, 34, 35, 74 

open sites, 161 

organotins, 122 

origin, 17-22 

orthodox seeds, 61 

orthodox with limited desiccation ability (OLDA) 

seeds, 61, 63-7, 68-9 

outcrossing, 46-8 


ovaries, 7, 9, 11, 24, 34 

overtopping vegetation, removal of, 164 

oxygen levels in storage, 81 

P 

paclobutrazol, 68 

Pakistan, 115, 117, 152, 163 

paleobotany, 16-22 

fossil sites, 6, 15 

Papua New Guinea, 117, 134, 135 

parasites, 121, 122, 124 

see also fungi 

parrots, destruction by, 116 

partial dessication, 81 

pathogens, 115, 119-25, 170-1 

Peninsula Malaysia, see Malaysia 

pericarps, 24, 61 

perlite, 80 

permethrine, 119 

peroxidation, 63 

pests, 115-19, 123-5, 157, 164-5, 170-1 

petals, 5 

petioles, 29 

phenology, 74-5, 89-90, 152 

phenols, in litter, 103 

Philippine Selective Logging System (PSLS), 137, 

139 

Philippines, 134, 153-4, 157 

diseases, 122 

216 



growth and yield, 140 

mycorrhizas, 101, 104, 109 


pests, 117, 118, 119 


silviculture, 138-9, 142, 144, 162 

phosphorus, see mineral nutrition 

photosynthesis, 94, 117, 121 

photosynthetically active radiation (PAR), 90-1, 93 

phylogeny, 5-44 


physiology, 57-71 

changes, following defoliation, 117 

growth in relation to, 94-5 

state of mother trees, 73-4 

phytogeography, 5-6, 7, 12-22 

pigs (wild), destruction by, 116, 117 

pin-hole borers, 119 

piney resin, 190 

piney tallow, 193 

plantations, 151-85, 170 

monospecific, 115 

resinous trees, 192 

planting site, 160-2 

planting stock production, 156-63 

planting techniques, 162-3 

plastic bags, storage in, 77, 80-1, 159, 162 

poison-girdling, 138, 165, 166 

pollen, 7, 9, 11, 26, 27, 31 

dispersal, 48 

grain size, 24 

see also stamens 

pollen exine, 9, 25, 27 

pollen tubes, 30 

pollination, 24, 30, 34, 46-51, 50 

polycyclic (selection) systems, 51, 135-6, 138, 139, 

142, 144-5 

polyembryony, 30, 49 

polyethylene bags, storage in, 81 

polyploids and polyploidy, 29-30, 45-6 

population densities, 51 

population diversity, 52 

post-harvest-maturation phenomenon, 60 

potassium, see mineral nutrition 

potted seedlings, 157, 160, 161, 162 

pre-felling treatment, 169

General Index 

preservatives, 119, 122-3, 124 

processing seeds, 79, 83-4 

procurement of seeds, 75-7 

propagation of seedlings, 157 

R 

rain-weeding, 165 

re-establishment by natural regeneration, 134-5, 168- 

70 

recalcitrant seeds, 61, 68 

handling, 73, 83-4 

storage, 79-83 

Reduced Impact Logging (RIL), 142 

reforestation, 115, 151, 170 

regeneration, 3, 134-5, 168-70 

enrichment planting, 141, 161, 164 


‘nursing’ phenomenon, 107 


stands, 155-64 

Regeneration Fellings, 137 

Regeneration Improvement Fellings (RIF), 137, 138 

Regeneration Improvement Systems, 154, 167 

relative humidity, for storage, 82, 159 

removal fellings, 168, 169 

reproduction, 46-51, 89-98, 92 

see also germination; pollen; fruiting; seeds 

research priorities, 1-4 

biogeography and evolutionary systematics, 35 

conservation of genetic resources, 51-3 

mycorrhizas, 106-10 

pests and diseases, 124-5 


seed handling, 84-5 

seed physiology, 68-9 

resin canals, 10, 27, 28, 29 

resins, 10, 26-7 

alkaloides, 116 

after defoliation, 117 

forest products, 187-92, 195 

riverine fringes, 12 

rock dammar, 192 

rodents, destruction by, 116, 117, 157 

root borers, 117 

root cankers, 120-1 

root pruning, 157, 159, 160, 163 

root rot, 120, 121 

roots, 99-114, 117 

in compacted soil, 155 

diseases, 120-1, 122 

mass, 93, 94 

pests, 117, 118 

‘roping up’ collection method, 77 

rotations, 168-9, 171, 172 

rots, 121-2, 171 

S 

Sabah, 12, 153 

climbing bamboo, 165 

growth and yield, 140 


nutrient availability, 105-6 

silviculture, 138, 142 

vegetative propagation, 159 

sal forests, 134-5, 151, 152, 187 

non-timber forest products, 193, 194 

resins, 190, 192 

silviculture, 136-7 

sandstones, 11 

sandy sediments, 11 

sap-staining fungi, 123 

sap suckers, 118 

saplings, see seedlings 

sapwood decay, 121 

Sarawak, 12, 153 

assignment of species to site, 155 

growth and yield, 139-40 

mineral nutrition, 106 


pests, 118 

pollen vectors, 48 

silviculture, 138, 142, 172 

wilding planting stock, 159 

savanna woodlands, 11 

sawdust, storage in, 80, 157, 159 

sea beaches, 170 

scented balsam (chua), 191 

seasonal areas/forests, 11, 12, 23, 24, 134, 136 

flowering and fruiting, 73, 135 

secondary forest, 167, 168 

sections, 8 

seed abortion, 122 

seed-borne fungi, 120 

seed cakes, 194 

seed fungi, 119-20 

seed handling, 73-88 

217

General Index 

seed material, cryopreservation of, 82-3 

seed physiology, 57-71 

seed trees, treatment of, 169 

seed water potential, 67 

seeding 

artificial induction, 68 

age, 168 

seedling felling, 168 

seedlings, 27, 31, 34, 89-98, 135 

diseases, 120-1 

fertilisation, 105-6, 107 

in vitro, 160 

light requirements, 105 

multiple, 30, 49 

mycorrhizal infection, 103, 104-5, 154-5, 108 

pests, 116-17 

planting stock, 157-8 

storage, 82 


seeds, 23, 34, 135 

canopy tree species, 92 


dispersal, 48-9, 89 

pests, 116 


polyembryonic, 30 

products, 193, 194 

see also fruits 

Selection Improvement Fellings, 138 

selection systems, 51, 135-6, 138, 139, 142, 144-5 

Selective Management System (SMS), 137-8, 144 

self-incompatible breeding systems, 46-7 

sepals, 7, 9, 11, 26, 27, 28, 48-9 

base, 24, 27 

Seychelles, 13, 15 

shelter, 160-1, 162, 164 

see also light 

shelterwood systems, 135, 136-7, 143-4, 167-8, 169- 

70 

shoot borers, 117 

shoot-pruning, 157-8, 159 

silvics, 92, 152, 153, 154-5 

silviculture, 2, 3, 151-2 


plantations, 161, 171-3 

regeneration methods, 93 

stand species, 155 


Singapore, 116, 118, 121, 189 

site quality, 172 

site requirements, 155 

sites, 160-2 

size, 7-11, 29 

anthers, 24 

flowers, 23 

leaves, 94 

pollen grains, 24 

pots/containers, 162 


seeds, 61, 68, 78-9 

wildlings, 159 

skyline yarding systems, 142 

slash, disposal of, 123-4 

slopes, 11, 12, 92 

logging on, 142 

soft rot, 122 

soil, 12, 93, 161-2 

compacted, 155, 163 

degraded, 105, 151, 157, 161, 170 

fertility, 170 

mycorrhizas and, 99, 108 

see also fertilisers and fertilisations 

soil water, 93 

Somalia, 17 

South America, 5-6, 11, 12, 13, 14, 15, 18 

affinities, 25 

mycorrhizas, 104 

South Asia, 152 

distribution, 13, 14 


oleoresin trees, 188-90, 191 

seed handling recommendations, 83-4 

see also India; Sri Lanka 

Southeast Asia, 13, 14, 83-4, 19-2 

see also Burma; Cambodia; Thailand; Vietnam 

sowing, 157, 163-4 

speciation, 103 

species, 13, 52, 53 

criteria for definition of, 29 

see also light-demanding/shade-tolerant species 

Species Improvement Network, 157 

Species Improvement Program (Bangladesh), 152 

splice grafting, 160 

spurs, using to climb, 77 

squirrels, 116 

Sri Lanka, 18, 24, 134 

218

General Index 




habitats, 12 

mycorrhizas, 100, 101, 104, 109 


regeneration conditions, 169 


stamens, 5, 7, 9, 11, 24, 26, 28 

stand density, 51, 155, 172 

stands, 151-85 

starch, 160 

stem canker, 120 

stomatal conductivity, 94 

storage, 63-8, 79-83, 84 

category designations, 60-1 

chilling damage, 58-9, 60 

fungi, 120 

planting stock, 157: wildings, 159 


Stratified Uniform System, 138 



subfamilies, 5-6, 7, 8, 30-3 

subspecies, criteria for definition of, 29 

suckers, 118 

sucrose, 83 

Sulawesi, 13, 134 

Sumatra, 134 


dammars, 192 


distribution, 13, 19, 22 

oleoresin production, 190 

pests, 117 

wildings, 159 

Sumatra camphor, 192-3, 195 

Sundaland, 14 

superoxide, 63 


survival, 90-4, 105 

bare-root/ball-root transplants, 160 


container plants, 160 

insect pests, 116 


planting site, 161-2 

planting stock, 157-8 

planting techniques, 162-3 

seeds, in storage, 80-1 

wildings, 159, 163 

see also mortality 

swamps, 12 

swidden agriculture, 92 

Switzerland, 16 

T 

tannin, 193-4, 196 

Tanzania, 101 

tap roots, 157 

tapping resins, 188, 189-90, 192 

taungya plantations, 152, 155, 168, 170 

taxonomic diversity, 52 

taxonomy, 5-7, 15-16, 25-33 

telodrine, 119 

temperature 

flowering and, 75 

germination and, 57-9 

heat stress, 93 

storage conditions, 64-6, 80, 81, 82 

transportation of seeds, 77 

tending stands, 164-8 

tengkawang, 193 

termites, 118-19 

tetraploids, 30 

Thailand, 4, 133-4, 152, 157 


diseases, 119, 121, 122 


mycorrhizas, 100, 101, 102, 103, 104, 108-9 

oleoresins, 190, 191 

pests, 117, 118, 119 

phenological studies, 75 


thinnings, 165-8, 172 

thiram, 81, 82, 120 

thread blights, 121 

threatened species, identification of, 52 

thrips, 24, 48 

timber, 1, 166 


pests, 118-19 

tissue chemistry, 94-5 

tissue culture, 67, 83, 160 

topography, 92, 93 

see also slopes 

219

General Index 

TPI, 139 

tractors, 155 

transpiration, 94, 163 

transplants, 160, 163 

transportation of seeds, 77-9, 83 

tree bicycles, 77 

tree climbers, collection using, 76-7 

tree diseases, 121-2 

tree falls, 92 

tree pests, 117-18 

tribes, 30-3 

triploids, 30, 49 

U 

underplanting, 3, 161, 162 

understorey, 23, 94, 163 

Uniform System, 136 

United Kingdom, 16, 69 

United States of America, 16, 19, 23 

V 

vacuolar membrane rupture, 63 

vegetation, 160-2, 164, 168 

vegetative propagation, 103, 159-60 

Venezuela, 12, 14 

ventilation, for storage, 80-1 

ventilation, for transportation, 77 

vermiculite, storage in, 80 

vesicles, 63 

vesicular-arbuscular mycorrhizas (VAM), 99, 100, 102 

vicarious species, 22 

Vietnam, 13, 19, 22, 152, 170 

W 

Wallace’s line, 12, 14, 21 

waste land, 170 

water, 93, 94 

disperal by, 49 

drainage, 123 

water status of seeds, 67 

see also moisture content 

weeding, 157, 164-5 

weevils, 116 

weight of seeds, 78-9 

West Bengal, 160, 170 

West Kalimantan, see Kalimantan 

West Malaysia, see Malaysia 

West Malesia, see Malesia 

wet regions (everwet regions), 11, 12, 169 

storage physiology in relation to, 68 

white dammar, 190, 192 

white rot, 122 

whole-seed moisture content, 59, 60 

wild pigs, destruction by, 116, 117 

wilding planting stock, 158-9, 163 

wilting, 120-1 

wind, 60, 48-9 

winged fruits, 20, 21, 24-5, 29 

wood anatomy, 28 

wood decay, 122 

wood rays, 7, 10, 11, 26 

wood resins, see resins 

wood products, 118-19, 122-3, 124, 194 

wood staining fungi, 122-3 

woody vegetation, 164 

wrenching, 157 

Y 

yield, 139-41 

camphor, 192 

dammar gardens, 192 

seed production stands, 156-7 

Yunnan, 134 

Z 

Zaire, 14 

Zambia, 14, 101 

zonation, 11, 12 

zygotic embryos, 30 

220

A review of dipterocarps - Center for International Forestry Research

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