A Review Dipterocarps

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
MA
L AYSI A
TI
INS TUTE
CIFOR
&(17(5)25,17(51$7,21$/)25(675<5(6($5&+
FOREST
SEARC
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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: [email protected]
Website: http://www.cgiar.org/cifor
Contents
Authors
Abbreviations
Acknowledgements
Foreword
v
vii
ix
xi
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
45
Chapter 3. Seed Physiology

P.B. Tompsett
57
Chapter 4. Seed Handling

B. Krishnapillay and P.B. Tompsett
73
Chapter 5. Seedling Ecology of Mixed-Dipterocarp Forest

M.S. Ashton
89
Chapter 6. Root Symbiosis and Nutrition

S.S. Lee
99
Chapter 7. Pests and Diseases of Dipterocarpaceae

C. Elouard
115
Chapter 8. Management of Natural Forests

S. Appanah
133
Chapter 9. Plantations
G. Weinland
151
Chapter 10. Non-Timber Forest Products from Dipterocarps

M.P. Shiva & I. Jantan
187
Scientific Index
General Index
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 Biomtrie, Gntique et Biologie
des Populations
Universit 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
52109 Kuala Lumpur
Malaysia
S. S. Lee
Kepong
52109 Kuala Lumpur
Malaysia
G. Maury-Lechon
U.M.R. C.N.R.S. 5558
Laboratoire de Biomtrie, Gntique et Biologie
des Populations
Universit 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
50660 Kuala Lumpur
Malaysia
Abbreviations
ABA
ACOM
AFTSC
ASEAN
ASTAG
BHC
BIO-REFOR
BIOTROP
CIFOR
DABATTS
DENR
DFID
DNA
EEC
FAO
FD
FORSPA
FRIM
GTZ
IBPGR
ICFRE
IIED
IPGRI
ITTO
IUCN
Abscisic acid
Asian Conference on Mycorrhizae
ASEAN Forest Tree Seed Centre
Association of Southeast Asian
Nations
Agriculture Division in the Asian
Technical Department, World Bank
(ceased January 1993)
Benzene hexachloride
Biotechnology assisted
Reforestation
See SEAMEAO-BIOTROP
Center for International Forestry
Research
Database of tropical tree seed
research
Department of Environment and
Natural Resources, Philippines
Department for International
Development (United Kingdom)
Deoxyribonucleic acid
European Economic Community
Food and Agriculture Organization of
the United Nations
Forest Department of Peninsular
Malaysia
Forestry Research Support Program
for the Asia-Pacific
Deutsche Gesellschaft fr
Technische Zusammenarbeit
International Board for Plant Genetic
Resources (now IPGRI)
Indian Council of Forestry Research
and Education
International Institute for
Environment and Development
International Plant Genetic
Resources Institute
International Tropical Timber
Organisation
The World Conservation Union
IUFRO
IWGD
JICA
JIRCAS
LN
LSMC
MC
MP
MTC
MUS
NCT
NTFPs
ODA
OLDA
OTA
PAR
PCARRD
PCT
PEG
PSLS
RAPA
RAPD
RIF
RIL
ROSTSEA
SEAMEOBIOTROP
SMS
SPDC
International Union of Forestry

Research Organizations
International Working Group on
Dipterocarps
Japan International Cooperation
Agency
Japan International Research Centre
Liquid nitrogen
Lowest-safe moisture content
Moisture content
Melting point
Malaysian Timber Council
Malayan Uniform System
Non-crop trees
Non-timber forest products
Overseas Development Authority
(United Kingdom) (now DFID)
Orthodox with limited desiccation
ability
Office of Technology Assessment
Photosynthetically active radiation
Philippine Council for Agriculture,
Forestry and Natural Resources
Research and Development
Potential final crop trees
Polyethylene glycol
Philippine Selective Logging System
Regional Office for Asia and Pacific
(FAO)
Random Amplified Polymorphic
DNAs
Regeneration Improvement Fellings
Reduced Impact Logging
Regional Office for Science and
Technology for South East Asia
(UNESCO)
South-East Asian Regional Centre for
Tropical Biology
Selective Management System
Special Programme for Developing
Countries (IUFRO)
viii
SPINs
TPI
TPTI
TROPENBOS
Species Improvement Network

Tebangan Pilih Indonesia (Indonesian
Selective Cutting System)
Tebang Pilih Tanam Indonesia
(Modified Indonesian Selective
Cutting System)
The Tropenbos Foundation,
Netherlands
TSI
UNESCO
UPM
USDA
VAM
Timber Stand Improvement

United Nations Educational,
Scientific and Cultural Organization
Univesiti Pertanian Malaysia
(Agriculture University of Malaysia)
United States Department of
Agriculture
Vesicular arbuscular mycorrhizas
Acknowledgements
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 worlds tropical rain forests. At that time it was
evident that a particular focus for CIFORs research
should be in the moist tropical forests of Asia. Trees in
the family Dipterocarpaceae, the dipterocarps are a
major component of southeast Asias 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.
2
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
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
3
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
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
4
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, 19151928. 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 worlds 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.
Chapter 1
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 (MauryLechon 1991). Excessive canopy openings provoke the
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
xx
$XVWUDOLDQ
VKHOI
x
x
x x
x x
xx
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 (Londoo et al. 1995,
:DOODFHVOLQH
XXX
S
F
D
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

Table 1. Recent content of Dipterocarpaceae family.
Families
Sub families
Genera
Maguire et al. 1977, Maguire and Ashton 1980

Dipterocarpaceae
Monotoideae
Monotes
Pakaraimoideae
Pakaraimaea
Marquesia
Dipterocarpoideae
see table 2
Maury 1978, Maury-Lechon 1979a, b*

Monotaceae
Monotoideae
Monotes
Marquesia
Pakaraimoideae
Dipterocarpaceae
see table 2
Pakaraimaea
see table 2
Kostermans 1978, 1985, 1989

Monotaceae
Monotes
Marquesia
Pakaraimaea
Dipterocarpaceae
see table 2
see table 2
Londoo 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
Table 2. Recent (1994) genera, sections and sub-sections related to Dipterocarpaceae and authors. (Londoo 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
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
not yet created
2 Neobalanocarpus
3 Shorea
s.Shorea
s.s.Shoreae
3 Balanocarpus
2 Shorea
3 Shorea
s.Shoreae
4 Shorea
Shorea including
s.g.Eushorea= Shorea
s. Barbatae
Pentacme genus
s.g.Richetia
4 Richetia
s.Richetioides
s.s.Barbata
(1992: p.60)
s.Richetioides
s.s.Richetioides
s.s.Polyandrae
s.Anthoshorea
s.Mutica
s.Maximae
s.g.Anthoshorea
5 Anthoshorea
s.g.Rubroshorea
6 Rubroshorea
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.Smithiana
subgr.Smithiana
s.Pachycarpae
2 Neobalanocarpus
2 Balanocarpus heimii
subgr.Pinanga
s.s.Brachypterae
s.s.Smithianeae
s.Pachycarpae
s.Doona
7 Doona
s.Pentacme
8 Pentacme
5 Doona
4 Parashorea
3 Parashorea
9 Parashorea
5 Dryobalanops
4 Dryobalanops
10 Dryobalanops
7 Dryobalanops
6 Dipterocarpus
5 Dipterocarpus
11 Dipterocarpus
8 Dipterocarpus
9 Anisoptera
7 Anisoptera
6 Anisoptera
12 Anisoptera
s.Anisoptera
s.Pilosae
s.Anisoptera
6 Parashorea
s.Glabrae
s.Glabrae
s.Glabrae
8 Upuna
7 Upuna
13 Upuna
9 Cotylelobium
8 Cotylelobium
14 Cotylelobium
10 Vatica
9 Vatica
15 Sunaptea
11 Sunaptea (+Coty.)
s.Sunaptea
s.g.Synaptea
12 Vatica
s.Vatica
s.g.Isauxis
16 Vatica
s.Vatica
(s.Pachynocarpus 1964)
s.g.Pachynocarpus
10 Upuna
s.Pachynocarpus
11 Stemonoporus
17 Stemonoporus
13 Stemonoporus
12 Vateria
18 Vateria
14 Vateria
13 Vateriopsis
19 Vateriopsis
15 Vateriopsis
14 Monotes
15 Marquesia
16 Pakaraimaea
s.: section;
s.s.: sub-section;
s.g.: sub-genus;
subgr.: sub-group.
inflorescence
paniculate (compound raceme)
(+)
cyme appearance
(+)
+
5-merous perianth
Sarcolaenaceae
+
+
+
imbricate
valvate
imbricate
valvate
numerous
many
subversatile anthers
connectival appendage
ovary
2-celled anthers generally dehiscing longitudinally
exine pollen
+
(-)
centrifugal stamens
pollen
+
(-)
imbricate
persistent sepals and calyx
hypogynous stamens
valvate
contorted corolla
fruit-sepals
Ochnaceae
unisexual flower
open-flower sepals
Elaeocarpaceae
bisexual flower
bud-flower sepals
Tiliaceae
Theaceae
sensu lato
Pakaraimoideae
racemi-paniculate
Guttiferae
Dipterocarpaceae*
Dipterocarpoideae
BOTANICAL CHARACTERS
Monotoideae
Table 3. Affinities between Dipterocarpaceae sensu lato and other close angiosperm families.
tricolporate
tricolpate
2-3 layers
4 layers
(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
+
+
10
leaves
opposite
alternate
leaf venation
prominent pinnate
vertically transcurrent
entirely transcurrent and
Sarcolaenaceae
Ochnaceae
Elaeocarpaceae
Tiliaceae
Theaceae
sensu lato
Pakaraimoideae
Guttiferae
Dipterocarpaceae*
Dipterocarpoideae
BOTANICAL CHARACTERS
Monotoideae
Table 3. (continued) Affinities between Dipterocarpaceae sensu lato and other close angiosperm families.
+
+
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
geniculate petiole
indumentum + complex anatomy petiole (Malvales type)
rays
mixed uni/multiseriate
+
+
+
intercellular resin canals
mucilage canals in cortex and cells in the epidermis
+
+
pith and primary cortex with indumentum
anomocytic stomata
complex petiolar vascular supply
n=7
n=11
chromosomes
: present and other possibilities;

+: present;
-: absent;
into outwardly tapering wedges
presence of resins
elongate medullary mucilage cells
multiseriate
+
-
uniseriate
biseriate
complex indumentum
arrangement of bast fibres *
+
-
+
+
(+)
-
(+)
(+)
(-) or (+): exceptions;

* : adapted from Ashton 1982, Maury-Lechon 1979, and other
works (see in text: Classification).
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), 300750 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,
Wallaces 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).
13

Table 4. Phytogeographical regions and distribution of numbers of genera and species.
Area Country
Number of genera
area
Malesia
country
10
Number of species
area
Authors
country
465*
*Ashton 1982
Malaya
14
155* 168
Symington 1943
Borneo
13
267* 276
"
Sumatra
106*
*Ashton 1982
Philippines**
11
50* 52
"
Sulawesi
*Ashton 1982
Moluccas
*Ashton 1982
New Guinea
15
"
W. Wallaces line
E. Wallaces line
Mainland
Southeast Asia
76
Smitinand 1980
Smitinand et al.1990
Burma
33
"
Thailand
66
"
Laos
20
"
Cambodia
28
"
Vietnam
36
"
China
24
Huang 1987
11
Xu and Yu 1982
Sri Lanka
44-45
South Asia:
58
Kostermans1992
5 (6)
31
Tewary 1984
India+Andamans
Ashton 1977
North India
10
South India
14
Andamans
Seychelles
2
1
Africa
plus Madagascar
Africa
Madagascar
South America
8
1
Jacobs 1981
"
"
Parkinson 1932
37*
* Ashton 1982
49
Shaw 1973
48
Shaw 1973
36
Ashton 1982
30
Verdcourt 1989
Maguire et al.1977
Maguire and Ashton 1980
*: numbers in Ashtons 1982 publication;

: Shaws numbers in Jacobs 1981;
Philippines**: east of Wallaces line only 3 genera and 13 species.
NB: - Number of genera in China assumes Chinas 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.
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 (Londoo 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 Ashtons 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 Wallaces
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 Wallaces 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 Wallaces 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
Wallaces line. This is an important fact.
15

Table 5. Distribution of living and fossil dipterocarp genera or section.
Taxa
S.Am Afri
Pakaraimaea
Marquesia
Monotes
Vateriopsis
Mada Seyc India Sri-L
Burm InCh
Thai
Mal
Born Indo
Phil
N.Gu
O*
O*
O*
O*
O
O
Vateria
O*
Stemonoporus
Doona
Balanocarpus K.
Vatica Kosterm.
Dipterocarpus
Chin
* (?)
O*
O*
Anisoptera
Anthoshorea
O?
s.Shorea (*)
O*
O?
O*
O*
O*
s.Hopea
O*
(O)
s.Dryobalanoides
Parashorea
Pentacme
*(?)
Sunaptea
Cotylelobium
(O)
O
O
O*
O*
s.Richetioides
(O)
s.Rubroshoreae
(O)
O
O
Neobalanocarpus
Dryobalanops
s.Brachypterae
s.Pachycarpae
s.Rubellae
s.Neohopea
Upuna
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
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). MauryLechons previous conclusions (Maury 1978, MauryLechon 1979a, b: Fig. 16, p.100) based on cotyledonary
shapes and structures have been vindicated by
Wickneswaris 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 Wickneswaris 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, Krasel 1922, Schweitzer 1958, all in
Aubrville 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 Aubrville 1976). Doubts
were cast on these identifications (Bancroft 1933, Harris
1956 and Hughes 1961 in Aubrville 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 AsiaticMalesian 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 cest avec des bois de la famille des
Dipterocarpaceae, notamment ceux du genre Monotes
que notre echantillon parait avoir le plus daffinites...
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

Dipterocarpaceae
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 PermoTriassic 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 Aubrville 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. Aubrville (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).
18

Figure 2. Continental drifts concerning the area of Dipterocarps from Primary to Tertiary (adapted from
Renous 1989).

0<
Miocene

0<

0<
Late
Cretaceous

0<

0<
(XUDVLD
1$P
Middle & late

Jurassic
$I
6$P
,
$X
0
$Q
I: India
M: Madagascar
S.Am: South America
N.Am: North America
Af: Africa
Au: Australia
An: Antarctica
SrL: Sri Lanka
6U/

0<
LAURASIA
Triassic
GONDWANA

0<
Permian

0<
MY - Million of years
19

Table 6. Dipterocarp fossil distribution in time and space.
Fossil genera
Number of species
total
Geological
country
II
Cret.
Monotes oeningensis flower?

Monotoid fruit remants?
Monotoids?? :
Calicites alatus, C.obovatus
Monotoid fossil?
Dipterocarpoxylon? Woburnia
Dipterocarpophyllum?
Dipterocarpophyllum
Dipterocarpus-type pollen
"
Dipterocarpoxylon? Monotoid
(Lemoigne 1978)
Dipterocarpoxylon? Monotoid?
Dipterocarpoxylon
"
"
"
"
Anisopteroxylon
"
Vaticoxylon
"
Vaterioxylon
Shoreoxylon
"
"
"
"
"
"
"
Pentacmoxylon??
Hopenium
3
2
1
1
1
1
1
1
30
Dryobalanoxylon
13
"
"
Dryobalanops pollen
Dryobalanoxylon
"
# : early period;
* : mid period;
& : upper period;

+ : early+mid-period;
7
2
2
23
1
4
1
1
Hungary
Germany, Austria
Switzerland
2
1
1
1
1
1
1
1
New-York USA
Alaska
Great Britain
Egypt
Nepal, N. India
Nepal, N. India
Vietnam
Ethiopia
2
11
2
2
3
7
5
2
1
1
2
13 3
1
1
3
1
1
1
7
1
2
2
1
1
1
1
6
5
Somalia
North India
Burma
Vietnam
Sumatra
Java
N. and NW India
India
Sumatra
Java
North India
India-Assam
Northwest India
North India
South India
Assam-Cambodia
Burma
Thailand
Sumatra
India
North India
South India
Cambodia
South Vietnam
Borneo
Borneo
Sumatra
Java
periods
III
Eoc.
IV
Olig.
Mio.
*
*
Plio.
Plei.
&
#
*
*
&
&
&
&
&
III
#
+
III
III
III
III
IV
**
**
#
#
&
**
**
*
#
#
#
**
IV
III
*
III
*
#
*
#
IV
III?
*
*
IV
#
*
*
*
**
*
#
** : mid+upper period.
The two successive hypotheses, of Aubrvilles

double-cradle areas, and of the Vozenin-Serra and SalardCheboldaeffs archipelago connection anterior to Indias
contact with Laurasia, could support a possible very
remote common ancestor of American, African and Asian
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.
20

Figure 3. Paleogeographical reconstitution of southeast Asian Permo-Triassic in conformity with paleobotanic
data (from Vozenin-Serra and Salard-Cheboldaeff 1994).
1
9
2
8
3
4
7
6
1.
2.
3.
4.
5.
6.
7.
8.
9.
Sinkiang block (Tarim) + Sino-Korean block (North China).

Yangtse block (North China).
North Tibet block (North Xizang + North-West Xunnan + Shan Plateau).
Khorat-Kontum block (East Thailand, most of Laos, Cambodia and Vietnam, South of Song Ba suture).
South Tibet block (Lhassa plate + lands located South of Yalu Tsangpo).
Kashmir.
Iran and Afghanistan (Helmand block).
Turkey.
Spain.
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
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)). Wallaces 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-SeychellesSri 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) IndiaSri Lanka-Madagascar-Africa-America-Eurasia (Fig. 2
(C)); b) the putative eastern archipelago northeast of
Gondwana to Eurasia (Fig. 3); c) later, India-Sri LankaEurasia (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 Wallaces 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
22

Table 7. Distribution and importance of endemicity in Dipterocarpaceae family.
Geographical areas
Ashton 1982
number of
species
Seychelles
Sri Lanka
South India
North India
Andamans
Burma
China mainland
Hainan
Vietnam
Laos
Cambodia
Thailand
Peninsular Malaysia
Sumatra
Java
Lesser Sundas
Lombok
Borneo
Philippines
Celebes
Moluccas
New Guinea
North Peninsular Malaysia
(Malayan-Burmese floristics)
Jacobs 1981
% of
endemicity
155
106
10
18
10
20
3
267
50
7
6
15
0
58
47
29
16
73
total number
of species
number of
endemics
% of
endemicity
1
44
13
13
8
32
5*
1
35
19
27
63
156
95
10
3
1
43
11
4
1
0
3
0
3
1
0
0
26
10
2
0
267
45
7
6
15
47
158
21
2
1
11
23
100
98
85
40
12
0
60 ?
0
9
5
0
0
17
10
20
0
0
55
47
29
16
73
49
* 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
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 (Ashtons 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,
Ashtons 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 Ourissons 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 Ashtons and MauryLechons 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 Kengs (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, MauryLechon 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
30
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).
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 MauryLechon 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 Ashtons 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):
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.
Tribes (2):
31
the herbarium collections of Asia (Forest Research

Institute Malaysia, Kepong including Symingtons
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 Ashtons, Meijers, and Maurys,
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-Lechons 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
Taxonomical levels
Supra-family
level (1) (order
or sub-order):
Family level
(2):
Sub-family
level (2):
Tribe level (4):
a)
b)
c)
d)
Dipterocarpales (2 families)
Monotaceae (3 genera:
Pakaraimaea, Marquesia,
Monotes)
Dipterocarpaceae (2 infra-family
groups: sub-families, 4 sub-groups:
tribes, 9 sub-subgroups: sub-tribes,
19 genera)
Imbricate [2 tribes (a & b), 3 subtribes, 9 genera, 19 sections]
Valvate [2 tribes (c & d), 7 subtribes, 10 genera, 8 sections]
Hopeae (2 genera, 4 sections):
Hopea (4 sections),
Neobalanocarpus
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)
Dipterocarpae (2 sub-tribes, 3
genera, 2 sections):
Dryobalanoinae (1 genus):
Dryobalanops
Dipterocarpinae (2 genera):
Dipterocarpus, Anisoptera (2
sections)
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
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.
32

Table 8. Comparative classifications of Asian Dipterocarpaceae.
MAURY 1978
HEIM 1892
Hopea
Euhopea
Pierrea
Dryobalanoides
Bracteata
Pachynocarpus
Parvifolia
subgr.
SHOREA
Dryobalanops
Dryobalanops
Dryobalanops
Dipterocarpus
Dipterocarpus
Dipterocarpus
Anisoptera
Pilosae
Glabrae
Anisoptera
Anisoptera
Glabrae
Upuna
Upuna
Cotylelobium
Vatica
Synaptea
Isauxis
Pachynocarpus
DIPTEROCARPI
IMBRICATE
SUB-VALVATE
--------------------------------------
Damar hitam
Parashorea
Vaticae
Vaticinae Stemonoporinae Vaterinae
Upuna
(Upuninae)
-----------------------Cotylelobium
Sunaptea
Sunapteae
Vaticoides
-----------------------Vatica
Vaticae
Balau
Pauciflora
subgr.
Anisoptera
Pilosae
Glabrae
Cotylelobium
Cotylelobium
Vatica
Sunaptea
Vatica
(Pachynocarpus*)
Vatica
Synaptea
Isauxis
Pachynocarpus
Stemonoporus
Eustemonoporus
Monoporandra
Vesquella
Vateriopsis
Stemonoporus
Sphaerae
Ovoides
-----------------------Vateriopsis
Stemonoporus
Vateria
Paenoe
Hemiphractum
Vateria
Vateria
*: Ashton 1964, 1968, spelling changes in and after 1982;
Ovalis subgr.
Red Meranti
Dipterocarpus
Shorea
Shorea, Barbata
Neohopea
Richetioides
Richetioides
Polyandrae
Pentacme
Shorea
Meranti Paang
Smithiana subgr.
Pinanga subgr.
Parashorea
Sunaptinae
VALVATE
Balanocarpus
hemii
Mutica
Mutica
Auriculatae
Ovalis
Rubellae
Brachypterae
Brachypterae
Smithiana
Pachycarpae
Parashorea
Pauciflora subgr.
Dryobalanops
(Dryobalanoinae)
Anisoptera
Anisoptera,
Glabrae
Rubroshorea
Parvifolia subgr.
Ovalis subgr.
Dipterocarpinae
Vatica
Euvatica, Isauxis
Retinodendron
Pachynocarpus
Shorea
Isoptera
Barbata, Ciliata
Richetia
Dipterocarpae
Cotylelobium
Sunaptea
Dyerella
SYMINGTON 1943
Hopea
Hopea
Hopea, Pierrea
Dryobalanoides
Dryobalanoides
Sphaerocarpae
Shorea
Pentacme
Doona
Anthoshorea
Pentacme
Shorea
Anthoshorea
Parashorinae
Brachypterae
Brachypterae
Smithianeae
Pachycarpae
Parashorea
Pentacme
Antoshorea
Anthoshoreae
Bracteolatae
Shorea
Shoreae
Barbatae
Richetia
Richetioides
Maximae
Rubroshorea
Muticae
Muticae
Auriculatae
Ovalis
Rubellae
Brachypterae
Brachypterae
Smithianeae
Pachycarpae
Parashorea
Tomentellae
Smithianae
Shorinae
Rugosae
Dipterocarpus
Sphaerales,
Angulati, Plicati,
Alati, Tuberculati
Anisoptera
Pilosae, Glabrae,
Antherotriche
ASHTON 1964-68-82
Doona
Doona
Shoreae
Isoptera
Richetia
Shorea
Eushorea
Anthoshorea
Hopeoides
Richetioides
Dryobalanops
Baillonodendron
Hopea
Group I
Group II
Neobalanocarpus
hemii
Anthoshorinae
Pentacme
MEIJER 1964
Hopea
Hopeae
Pierreae
Dryobalanoides
Sphaerocarpae
Balanocarpus hemii
Hopeae
Hopea
Euhopea, Hancea,
Dryobalanoides,
Petalandra
Pierrea
Duvallelia
Balanocarpus
Eubalanocarpus
Pachynocarpoides
Microcarpae
Sphaerocarpae
Parahopea
Doona
Vateriopsis*
subgr.: sub-group.
33
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 Meijers work, only the Asian genera
represented in this geographical area were analysed in
detail. Contrary to Maury-Lechons 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).
Taxonomical levels
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
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 (Londoo 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, Hall
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 (MauryLechon 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.0717 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
Unit 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 IndoChina. Likewise, there is a need for contact with African
colleagues
Acknowledgements
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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Chapter 2
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
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

Table 1. Infraspecific variation in chromosome number
(from Ashton 1982).
Species
Dipterocarpus alatus
Chromosome
Number
20, 22
D. tuberculatus
20
D. tuberculatus var. turbinatus
30
Hopea beccariana
H. odorata
H. subalata
20, 21, 22
14, 20, 21, 22
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
Tixiers 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
46
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.
47
Table 2. Breeding systems of Dipterocarps.

Species/Section
Shorea cordifolia
Section Doona
S. disticha
Section Doona
S. trapezifolia
Section Doona
S. trapezifolia
Section Doona
S. hemsleyana
Section Muticae
S. macroptera
Section Muticae
S. lepidota
Section Muticae
S. acuminata
Section Muticae
S. leprosula
Section Muticae
S. splendida
Section Pachycarpae
S. stenoptera
Section Pachycarpae
S. ovalis
Section Ovales
Dipterocarpus oblongifolius
Dryobalanops lanceolata
Hopea glabra
Section Richetioides
S. maxima
Section Richetioides
S. multiflora
1: Dayanandan et al. (1990);
Percent Fruit Set

Selfed
Crossed
0.5
21.0
Inferred Breeding
System
References
Self-incompatible
0.0
10.0
Self-incompatible
0.8
3.8
Self-incompatible
0.5
11.3
Self-incompatible
0.0
15.2
Self-incompatible
2.5
19.9
Self-incompatible
1.7
24.4
Self-incompatible
1.1
34.1
Self-incompatible
1.6
17.0
Self-incompatible
0.0
37.5
Self-incompatible
n/a
n/a
Self-incompatible
16.2
17.6
Self-compatible
69.3
n/a
n/a
64.0
n/a
n/a
Self-compatible
Self-incompatible
Self-compatible or
apomictic
2
3
2
n/a
n/a
Self-incompatible
n/a
n/a
Self-incompatible
2: Chan (1981);
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
3: Mamose et al. (1994).
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
48
ranges and therefore disperse pollen

over varying distances. Appanah and
Species
Outcrossing Rate tm References
Chan (1981) implicated thrips as
( standard error)
pollen vectors for several species of
Malaysian species of Shorea, section
Dryobalanops
Kitamura et al. (1994)
0.794 (0.059)
aromatica
0.856 (0.063)
Muticae. The thrips breed in flower
Shorea congestiflora
Murawski et al. (1994)
0.874 (0.021)
buds of the species they pollinate and,
S. megistophylla
0.860 (0.058)
as flowering progresses, they multiply
S. trapezifolia
0.617 (0.033)
in number. The adult thrips feed on
Stemonoporus
Murawski & Bawa
0.898 (0.022)
stamens and petals. As the petals of the
oblongifolius
(1993)
flowers are shed from the tree the
thrips fall on the ground and then move
to a new cohort of subsequently opened flowers. The
tropical tree species (Murawski and Hamrick 1990,
distances over which thrips move are not known but,
1991). Apparently, the low density of trees in logged
because of their relatively small body size, they
stands reduces the inter-tree movement of pollinators
apparently do not fly over long distances. It is presumed
and promotes self-pollination. Selfing, in part, may be
that their restricted movement is not a drawback in their
aided by a weak self-incompatibility system.
effectiveness as pollinators because the species they
Kitamura et al. (1994) compared outcrossing rates
pollinate are relatively abundant.
of Dryobalanops aromatica in primary and secondary
Dayanandan et al. (1990) present evidence for
forests but found no significant differences.
pollination of Shorea megistophylla (section Doona) and
Self-incompatibility as well as mating system studies
Vateria copallifera by bees (Apis spp.). They also
suggest that dipterocarps are predominantly outcrossed.
observed a wide variety of other insect floral visitors
Outcrossing in large populations can allow populations
including thrips. However, the thrips acted as flower
to harbour considerable genetic variation. In dipterocarps,
predators rather than pollinators, particularly in V.
mass flowering is also likely to enhance outcrossing by
copallifera.
allowing exchange of gametes among a very large number
More recently, Momose et al. (1994) have presented
of individuals. Not surprisingly, therefore, populations
evidence for pollination of Dryobalanops lanceolata, a
of dipterocarps show considerable genetic variation (see
large canopy tree species in Sarawak, by medium sized,
below).
stingless bees (Trigona spp.). They also noted the presence
Although the analysis of mating systems shows that
of many other types of flower visitors (Coleoptera and
the rates of outcrossing are high, it is also clear that there
Diptera). Momose et al. suggest that medium sized,
is a considerable potential for selfing in almost all species
stingless bees constitute an important group of pollen
examined so far. Moreover, apomixis has been reported
vectors for canopy and subcanopy trees in Sarawak (see
in several species (see below). While outcrossing
also Chan and Appanah 1980).
continuously generates new genetic variation, potential
Clearly, the dipterocarps are pollinated by a wide
for self-pollination and apomixis allows occasional new
variety of insects. The three detailed studies, respectively
variants to spread in the population or colonise new sites,
by Appanah and Chan (1981), Dayanandan et al. (1990),
and thereby promote differentiation of taxa.
and Momose et al. (1994) have revealed three different
classes of pollinators. Ashton (1982) in his extensive
Pollen and Seed Dispersal
review also lists beetles and moths as flower visitors,
Pollen dispersal influences the mating system, and both
although their role in pollination has not yet been
pollen and seed dispersal affect population genetic
demonstrated. Among the pollinators implicated so far,
structure. Limited dispersal results in inbreeding, small
all, except thrips, are capable of moving pollen over long
effective population sizes, and a high level of
distances. The extent of gene flow via pollen (and seeds)
differentiation among populations. Extensive dispersal
is further discussed in subsequent sections.
has the opposite effect.
Seed dispersal in most dipterocarps is by wind
Outcrossing in dipterocarps is achieved through a
(Ashton 1982). In most species, sepals are modified into
wide variety of pollinators that differ in their foraging
Table 3. Outcrossing rates of Dipterocarps.
49
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.
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.
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
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.
50
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
51
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.
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.
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
52
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
53
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.
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.
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. Ashtons earlier review. Sri Lanka particularly
deserves serious consideration because of the high
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.
Acknowledgements
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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Momose, K., Nagamitsu, T. and Inoue, T. 1994.
Reproductive ecology of an emergent tree,
Dryobalanops lanceolata, Dipterocarpaceae, in a
non-general flowering period in Sarawak. In: Inoue,T.
and Hamid, A.A. (eds.) Plant reproductive systems
and animal seosanol dynamics: Long term study of
dipterocarp forests in Sarawak. Canopy Biology
Programme in Sarawak, Series I. Kyoto University,
Japan.
Murawski, D.A. and Bawa, K.S. 1994. Genetic structure
and mating system of Stemonoporus oblongifolius
(Dipterocarpaceae) in Sri Lanka. American Journal
of Botany 81: 155-160.
Murawski, D.A., Dayanandan, B. and Bawa, K.S. 1994a.
Outcrossing rates of two endemic Shorea sp. from
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Murawski, D.A., Gunatilleke, I.A.U.N. and Bawa, K.S.
1994b. The effect of selective logging on mating
patterns of Shorea megistophylla (Dipteracarpacea)
from Sri Lanka. Conservation Biology 8: 997-1002.
Murawski, D.L. and Hamrick, J.L. 1990. Mating systems
of two Bombacaceous trees of a neotropical moist
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Murawski, D.L. and Hamrick, J.L. 1991. The effect of
the density of flowering individuals on the mating
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67:167-174.
Seavey, S.R. and Bawa, K.S. 1983. Late-acting selfincompatability in Angiosperms. The Botanical
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Slatkin, M. and Maruyama, T. 1975. The influence of

gene flow on genetic distance. American Naturalist
109: 597-601.
Stebbins, G. L., Jr. 1950. Variation and evolution in
plants. Columbia University Press, New York.
Symington, C.F. (1943). Foresters manual of
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Tixier, P. 1960. Donnes cytologiques sur quelques
Guttifrales rcoltes au Laos. Revue Cytolologique
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Tsai, L. M. and Yuan, Ct. T. 1995. A practical approach to
conservation of genetic diversity in Malyasia: genetic
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Jakarta.
Wickneswari, R. and Norwati, M. 1994. Genetic
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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
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REFOR, Kangar, Malaysia.
Wickneswari, R., Norwati, M., Tsumura, Y., Kawahara,
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Wright, S. 1931. Evolution in Mendelian populations.
Genetics 16: 97-159.
Chapter 3
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),
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 26-31C 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 30-35C was deemed optimal because
germination rates were faster. More recently, a standard
58
Seed Physiology
Table 1. Optimum germination temperatures, germination rates and base temperature values (Tompsett and Kemp
1996a, b).
Species
Shorea almon
Shorea siamensis
Shorea smithiana
Shorea pinanga
Hopea parviflora
Shorea roxburghii
Shorea robusta
Anisoptera marginata
Hopea foxworthyi
Hopea odorata
Shorea leprosula
Shorea parvifolia
Shorea contorta
Shorea affinis
Parashorea smythiesii
Dipterocarpus costatus
Dryobalanops aromatica
Parashorea tomentella
Shorea guiso
Anisoptera costata
Shorea ferruginea
Dipterocarpus obtusifolius
Cotylelobium burckii
Vatica mangachapoi
Dipterocarpus turbinatus
Cotylelobium melanoxylon
Dipterocarpus tuberculatus
Shorea amplexicaulis
Dipterocarpus zeylanicus
Shorea argentifolia
Base
temperature
(C)
Time to 50% final

germination at the
optimum
temperature (d)
Optimum
temperature
(C)
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
6.4
7.1
7.5
8.3
8.8
9.4
9.7
10.2
10.5
10.5
11.0
11.4
12.2
12.4
13.0
13.0
15.2
15.4
15.8
15.8
16.2
16.4
n/a
n/a
n/a
n/a
11
11
20
4
6
4
2
4
4
7
5
3
30
3
9
4
5
6
3
9
9
5
6
13
7
11
4
26
26
26
31
31
31
26
31
26
31
31
31
26
26
26
31
31
26
31
26
26
26
26
26
31
26
31
31
26
26
26
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 26C and 31C 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
Radicle length
defining
germination
(mm)
5
5
3
10
5
5
5
5
3
3
5
3
3
3
3
3
3
3
10
3
3
3
5
3
3
10
3
5
3
10
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 16C 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
Seed Physiology
(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 4C. By contrast, he believed that seeds of Shorea
species in the red meranti and balau groups were
intolerant of temperatures below 15C. 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 14C; secondly, seed of Shorea species in the
sections Mutica, Pachycarpae and Brachypterae were
considered intolerant of temperatures below 22-28C;
and, finally, seed of Shorea species in the Anthoshorea
section and seed of Hopea and Parashorea could be
cooled to 4C (but were recommended to be stored at
14C). 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 2C -5C relative to seed kept at
warmer temperatures (Purohit et al. 1982, Tompsett
1985). Another example of chilling damage which
59
occurred over a lengthy period of time is that to H.
hainanensis. For this species, seed at 5C almost all
died after 6 months; by contrast, at 15C -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
60
Seed Physiology
Table 2. Percentage moisture content and oil values for processed, dewinged whole seed and excised
seed parts (Tompsett and Kemp 1996a, b).
Species
Dipterocarpus intricatus ***

Dipterocarpus alatus ***
Dipterocarpus tuberculatus ***
Shorea ferruginea
Shorea argentifolia
Hopea ferrea
Shorea parvifolia
Hopea foxworthyi
Hopea odorata
Shorea gibbosa
Shorea macrophylla
Dipterocarpus grandiflorus
Anisoptera costata
Shorea fallax
Shorea affinis
Dipterocarpus chartaceus
Shorea leptoderma
Parashorea malaanonan
Dryobalanops keithii
Stemonoporus canaliculatus
Shorea macroptera
Whole-seed moisture
content* (percentage)
Axis moisture
content*
(percentage)
Embryo oil content

** (percentage)
n/a
16
11
11
29
29
32
32
34
36
37
38
38
40
40
40
42
42
44
47
47
48
50
53
55
56
n/a
13
n/a
51
n/a
62
52
54
64
n/a
66
63
69
70
n/a
70
63
n/a
61
66
56
64
n/a
74
7
19
61
n/a
9
n/a
n/a
20
n/a
10
n/a
n/a
57
n/a
33
n/a
n/a
8
n/a
n/a
n/a
n/a
n/a
n/a
*: 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 2C, 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).
***: 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:
Seed Physiology
orthodox; and
recalcitrant.
The orthodox type is capable of desiccation to a low
moisture content (approximately 5%) and storage for
several years at -20oC 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
61
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.
62
Seed Physiology
Table 3. Relationship between germination and moisture content during desiccation (Tompsett and Kemp 1996a, b)*.
Species
Shorea leprosula
Shorea argentifolia
Shorea ferruginea
Hopea ferrea
Hopea mengerawan
Hopea foxworthyi
Hopea odorata
Shorea parvifolia
Shorea roxburghii
Shorea obtusa
Shorea ovalis
Vatica mangachapoi
Shorea macrophylla
Shorea trapezifolia
Shorea fallax
Shorea macroptera
Shorea robusta
Vatica odorata ssp. odorata
Shorea affinis
Shorea almon
Shorea leptoderma
Lowest-safe moisture
content values
(percentage)**
Slope of probit line (probits

per unit of moisture content
percentage)
Intercept of probit line

(probit percentage
germination)
26
28
29
30
30
31
32
32
32
33
33
34
34
35
35
35
37
38
38
38
38
38
40
40
40
40
41
42
42
42
43
43
50
50
n/a
0.1814
0.1912
0.1050
0.1400
0.0994
0.1303
n/a
0.1000
0.0660
0.1816
0.1303
0.0919
0.1400
0.1743
0.0978
n/a
0.0789
0.0584
0.1427
0.0869
0.0867
0.2000
0.1307
0.0904
0.1300
0.0961
0.0475
0.1400
0.0460
0.1300
0.1200
0.1233
0.0965
n/a
-3.5780
-4.8300
-3.5660
-3.6700
-2.5555
-3.6450
n/a
-2.7700
-2.4420
-5.0550
-3.1790
-3.4460
-3.2200
-4.9920
-2.6190
n/a
-1.9360
-2.2470
-3.8090
-2.2470
-4.2300
-7.1400
-5.5600
-3.6510
-4.2200
-3.6290
-1.6120
-5.4700
-2.5280
-5.5300
-6.0400
-5.0770
-3.9830
*: 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
**: LSMC for seeds dried at 10-15% relative humidity and 1520C.
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
63
Seed Physiology
Table 4. Lowest-safe moisture content values (wet weight basis) for mature seeds*.
Species
Source
Dipterocarpus alatus**
Dipterocarpus caudatus
Dipterocarpus intricatus**
Dipterocarpus tuberculatus**
Hopea hainanensis
Hopea helferi
Shorea sumatrana
Shorea acuminata
Shorea acuminata
Shorea argentifolia
Shorea gibbosa
Shorea lepidota
Shorea pachyphylla
Shorea selanica
Shorea siamensis
Shorea singkawang
Shorea xanthophylla
Vatica umbonata
Tompsett (unpub.)
Tompsett (unpub.)
Tompsett (1987)
Tompsett (1987)
Tang and Tamari (1973)
Song et al. (1984)
Tang and Tamari (1973)
Yap (1986)
Chin et al. (1984)
Yap (1986)
Yap (1986)
Tompsett (unpub.)
Yap (1986)
Tompsett (unpub.)
Tompsett (1986)
Tompsett (unpub.)
Yap (1986)
Tompsett (unpub.)
Tompsett (unpub.)
Mahdi (1987)
*: 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
LSMC (%)
<11***
<47***
<12
<12
37
c.30
30
>40
<42
>40
35
>32***
27
>56***
>25
51***
55
>41***
43***
74****
***: 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
64
Seed Physiology
Table 5. Temperatures, moisture contents and germination of mature seeds for the optimum reported storage conditions.
Optimum storage achieved
Species
MC
(%)
Other conditions
Source
ventilated incubator, 99%

rh, rib-channel PB,
ventilated weekly
ventilated incubator, 99%
rh, PB, ventilated weekly
gas box, over water,
ventilated weekly
gas box, over water,
ventilated weekly
hermetic, laminated
aluminium foil bag
n/a
No information.
Tompsett
(unpub.)*
Germination
(%)
Days
Temp.
(C)
Anisoptera costata
44
30
18
44
45
84
21
48
52
28
21
29
Cotylelobium
melanoxylon
Dipterocarpus alatus**
46
67
21
36
44
2829
-13
11
Dipterocarpus baudii
Dipterocarpus
grandiflorus
Dipterocarpus
humeratus
Dipterocarpus
intricatus**
Dipterocarpus
obtusifolius
25
87
30
4
14
n/a
n/a
40
30
28
15
26
30
2373
-20
10
20
60
18
59
Dipterocarpus
tuberculatus**
Dipterocarpus
turbinatus
Dipterocarpus
zeylanicus
Dryobalanops
aromatica
77
1369
-20
12
20
177
16
42
53
100
21
39
50
16
14
38-40
54
23
16
45
Dryobalanops
lanceolata
Hopea ferrea
92
62
21
56
40
300
16
30-50
Hopea foxworthyi
68
365
18
35
Hopea hainanensis
80
365
18
35-38
Hopea helferi
85
40
15
48
n/a
Hopea mengerawan
40
67
21
44
Hopea nervosa
Hopea odorata
19
48
330
93
25
16
n/a
38
ventilated incubator at
99% r.h., loose
n/a
polythene rib-channel
bag within polythene box,
ventilated weekly
PB, sealed, inflated with

nitrogen
hermetic, laminated
aluminium foil bag
ventilated incubator 99%
rh, rib-channel PB,
ventilated weekly
hermetic, laminated
aluminium foil bag
closed box, over water,
ventilated weekly
rh, loose
n/a
PB tied, sawd. (18%
MC), ventilated weekly
PB sealed and inflated,
ventilated weekly
PB then perl.
ventilated incubator at
99% rh, rib-channel PB,
ventilated weekly
n/a
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Yap (1981)
Tompsett
(unpub.)*
Maury-Lechon
et al. (1981)
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Yap (1981)
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Song et al.
(1984, 1986)
Tang and
Tamari (1973)
Tompsett
(unpub.)*
Sasaki (1980)
Tompsett
(unpub.)*
65
Seed Physiology
Table 5. (continued) Temperatures, moisture contents and germination of mature seeds for the optimum reported storage
conditions.
Species

Germination
(%)
Days
Temp.
(C)
MC
(%)
Hopea parviflora
84
104
18
41
Hopea subalata
Hopea wightiana
Monotes kerstingii
40
5
16
51
60
90
4
4
2
32-43
n/a
7

ventilated weekly
n/a
n/a
PB, ventilated weekly
Neobalanocarpus
heimii
Parashorea densiflora
Parashorea
malaanonan
80
50
14
28-47
n/a
90
67
60
141
25
18
54
45
50
317
18
44
40
91
16
40
Shorea acuminata
Shorea affinis
70
56
30
253
21
21
38-43
35
Shorea almon
18
32
16
45
30
168
21
45
Shorea argentifolia
Shorea assamica
Shorea bracteolata
60
50
4
52
45
98
60
49
21
4
4
21
43
n/a
n/a
39
Shorea contorta
35
32
21
67
Shorea curtisii
Shorea dasyphylla
Shorea fallax
20
24
50
30
14
50
25
21
21
n/a
40-46
40
Shorea ferruginea
36
77
21
34
Shorea hypochra
Shorea javanica
10
87
60
30
4
20
n/a
13-15
Shorea leprosula
Shorea macrophylla
45
40
30
22
21
21
32
31
Shorea obtusa
44
11
16
36
Shorea ovalis
87
92
21
37
Shorea pachyphylla
28
16
18
66
Source
Other conditions
n/a
rib-channel PB,
ventilated weekly
rh, PB, perl. (0% MC),
ventilated weekly
PB, 4% moisture content
perl., ventilated weekly
n/a
rh, loose
stored on agar, some
seed germinated and are
included
PB, sealed, perl. 0-8%
MC, ventilated weekly
No information.
n/a
n/a
PB, top folded over,
within gas box, ventilated
weekly
rh, PB, ventilated weekly
n/a
n/a
PB tied, with sawd.,
ventilated weekly
PB, sealed and inflated,
ventilated weekly
n/a
moisture low
Tompsett
(unpub.)*
Sasaki (1980)
Sasaki (1980)
Tompsett
(unpub.)*
Yap (1981)
Yap (1981)
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Sasaki (1980)
Tompsett
(unpub.)*
Tompsett
(1985)
Tompsett
(unpub.)*
Sasaki (1980)
Sasaki (1980)
Sasaki (1980)
Tompsett
(unpub.)*
No information.
PB tied, with sawd.,
ventilated weekly
PB, sealed and inflated,
ventilated weekly
No information.
Tompsett
(unpub.)*
Yap (1981)
Sasaki (1980)
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Sasaki (1980)
Umboh
(1987)
Sasaki (1980)
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Sasaki (1980)

rh, over water
Tompsett
(unpub.)*
66
Seed Physiology
Table 5. (continued) Temperatures, moisture contents and germination of mature seeds for the optimum reported
storage conditions.
Species

Days
Temp.
MC
Other conditions
(C)
(%)
57
18
35
ventilated incubator
99% rh, PB, with perl.,
ventilated weekly
45
25
38-51 n/a
112
21
46
99% rh, loose
58
25
n/a
n/a
49
15
0
partial vacuum, MC
unknown
307
16
36
PB, tied and inflated,
ventilated weekly
56
15
40-48 No information.
Shorea parvifolia
Germination
(%)
40
Shorea pauciflora
Shorea pinanga
67
50
Shorea platyclados
Shorea robusta
80
54
Shorea roxburghii
52
Shorea siamensis
83
Shorea smithiana
28
46
26
44
Shorea sumatrana
Shorea trapezifolia
60
100
15
63
25
21
n/a
n/a
Stemonoporus
canaliculatus
Vatica mangachapoi
20
77
18-21
47
24
85
21
40-45
Vatica odorata ssp.

odorata
Vatica umbonata
48
148
18
40
10-50
60
10-15
n/a

ventilated weekly
n/a
97% rh, PB, ventilated
weekly, seed was pregerminated.
99% rh, loose
PB, perl. 0% MC,
ventilated weekly
99% rh, loose
n/a
Source
Tompsett
(unpub.)*
Sasaki (1980)
Tompsett
(unpub.)*
Yap (1981)
Khare et al.
(1987)
Tompsett
(1985)
Panochit et
al. (1984)
Tompsett
(unpub.)*
Yap (1986)
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
Tompsett
(unpub.)*
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;
perl.: stored in perlite;

sawd.: stored in sawdust;
PB: stored in ventilated polythene bag.
Note: not all reports specified the seed maturity.
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.
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:
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,
V = Ki - P/10KE - CWlog10m - CHt - CQt2............. (Eqn 1).
In this equation, V is the predicted viability, Ki 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 (oC). Viability is expressed as probit
germination. The constants KE, CW, CH and CQ are
67
Seed Physiology
Table 6. Viability constants and standard errors for two OLDA species of dipterocarps (Tompsett and Kemp
1996a, b).
Species
KE (se)
CW (se)
Dipterocarpus
alatus
6.44 (0.72)
3.09 (0.61)
0.0329 (0.0017)
0.000478 (0.000000)
Dipterocarpus
intricatus
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 = KE - CWlog10m - CHt - CQt2............. (Eqn 2).
In this equation, s represents the rate of loss of viability

in days per probit.
The viability constants KE, CW, CH and CQ 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 -13C 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
CH (se)
CQ (se)
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
68
Seed Physiology
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.
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 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 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.
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
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.
69
Seed Physiology
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 KE and, CW 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.
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 Departments 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.
Relevant Institutions
Berjak, P. and Pammenter, N.W. 1996. Recalcitrant

(desiccation-sensitive) seeds. In: Olesen, K. (ed.)
Innovations in tropical tree seed technology, 14-29.
Danida Forest Seed Centre, Humlebaek.
Chin, H.F., Hor, Y.L. and Mohd Lassim, M.B. 1984.
Identification of recalcitrant seeds. Seed Science and
Technology 12: 429-436.
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
Acknowledgements
I thank the Royal Botanic Gardens Kew and the Center
for International Forestry Research for facilities and
financial support
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Seed Physiology
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Maury-Lechon, G., Hassan, A.M. and Bravo, D.R. 1981.
Seed storage of Shorea parvifolia and Dipterocarpus
humeratus. Malaysian Forester 44: 267-280.
Mori, T. 1979. Physiological studies on some
dipterocarp species of peninsular Malaysia as a basis
for artificial regeneration. Research Pamphlet no. 78,
Forest Research Institute, Kepong, Kuala Lumpur, 76p.
Nautiyal, A.R. and Purohit, A.N. 1985a. Seed viability in
sal. I. Physiological and biochemical aspects of seed
development in Shorea robusta. Seed Science and
Nautiyal, A.R. and Purohit, A.N. 1985b. Seed viability
in sal. II. Physiological and biochemical aspects of
ageing in seeds of Shorea robusta. Seed Science and
Nautiyal, A. R. and Purohit, A.N. 1985c. Seed viability
in sal. III. Membrane disruption in ageing seeds of
Shorea robusta. Seed Science and Technology 13:
77-82.
Nautiyal, A.R. Thapliyal. A.P. and Purohit, A.N. 1985.
Seed viability in sal. IV. Protein changes accompanying
loss of viability in Shorea robusta. Seed Science and
Ng, F.S.P. 1980. Germination ecology of Malaysian
woody plants. Malaysian Forester 43: 406-437.
Olesen, K. (ed.) 1996. Innovations in tropical tree seed
technology. Danida Forest Tree Seed Centre,
Humlebaek. 302p.
Panochit, J., Wasuwanich, P. and Hellum, A.K. 1984.
Collection, germination and storage of Shorea
siamensis Miq. seeds. Embryon 1: 1-13.
Collection and storage of seeds of Shorea roxburghii
G. Don. Embryon 2: 62-67.
Purohit, A.N., Sharma, M.M. and Thapliyal, R.C. 1982.
Effect of storage temperatures on the viability of Sal
(Shorea robusta) and talura (Shorea talura) seed.
Seed Physiology
Roberts, E.H. 1973. Predicting the storage life of seeds.

Seed Science and Technology 1: 499-514.
Roberts, E.H. and Ellis, R.H. 1989. Water and seed
survival. Annals of Botany 63: 39-52
Sasaki, S. 1979. Physiological study on Malaysian
tropical tree species. Studies on storage and
germination of Leguminosae and Dipterocarpaceae
seeds. Tropical Agriculture Research Series 12: 7587.
Sasaki, S. 1980. Storage and germination of dipterocarp
seeds. Malaysian Forester 43: 290-308.
Scott, E.S., Rao, A.N. and Loh, C.S. 1988. Production
of plantlets of Shorea roxburghii G. Don from
embryonic axes cultured in vitro. Annals of Botany 61:
233-236.
Smits, W.T.M. and Struycken, B. 1983. Some preliminary
results of experiments with in vitro culture of
dipterocarps. Netherlands Journal of Agricultural
Science 31: 233-238.
Song, X., Chen, Q., Wang, D. and Yang, J. 1983. A study
of ultrastructural changes in radicle-tip cells and seed
vigour of Hopea and Vatica in losing water process.
Scientia Silvae Sinicae 19: 121-125.
on the principal storage conditions of Hopea
hainanensis seeds. Scientia Silvae Sinicae 20: 225236.
Song, X., Chen, Q., Wang, D. and Yang, J. 1986. A further
study on ultrastructural changes in radicle-tip cells of
Hopea hainanensis during deterioration resulted from
losing water. Tropical Forestry 4: 1-6.
Tamari, C. 1976. Phenology and seed storage trials of
dipterocarps. Research Pamphlet no.69. Forest
Department, Kuala Lumpur. 73p.
Tang, H.T. and Tamari, C. 1973. Seed description and
storage tests of some dipterocarps. Malaysian
Forester 36: 38-53.
Tompsett, P.B. 1982. The effect of desiccation on the
longevity of Araucaria hunsteinii seed. Annals of
Botany 50: 693-704.
71
Tompsett, P.B. 1984. The effect of moisture content on
the seed storage life of Araucaria columnaris. Seed
Science and Technology 12: 801-316.
Tompsett, P.B. 1985. The influence of moisture content
and temperature on the viability of Shorea almon, S.
robusta and S. roxburghii seed. Canadian Journal of
Forest Research 15: 1074-1079.
Tompsett, P.B. 1986. The effect of desiccation on the
viability of dipterocarp seed. In: Nather, J. (ed.) Seed
problems under stressful conditions. Proceeding of
the IUFRO Symposium, 181-202. Report, no. 12.
Federal Research Institute, Vienna.
Tompsett, P.B. 1987. Desiccation and storage studies
on dipterocarp seeds. Annals of Applied Biology 110:
371-379.
Tompsett, P.B. 1992. A review of the literature on storage
of dipterocarp seeds. Seed Science and Technology
20: 251-267.
Tompsett, P.B. 1994. Capture of genetic resources by
collection and storage of seed: a physiological
approach. In: Leakey, R.R.B. and Newton, A.C. (eds)
Proceedings of the IUFRO Conference Tropical
Trees; the Potential for Domestication and the
Rebuilding of Forest Resources, August 1992, 6171. Her Majestys Stationery Office, London.
Tompsett, P.B. and Kemp, R. 1996a. Database of tropical
tree seed research (DABATTS). Database Contents.
Royal Botanic Gardens Kew, Richmond, Surrey. 263p.
Tompsett, P.B. and Kemp, R. 1996b. Database of tropical
tree seed research (DABATTS). User Manual. Royal
Botanic Gardens Kew, Richmond, Surrey. Includes two
3.5 computer disks. 28p.
Umboh, M.I.J. 1987. Storage and germination tests on
Shorea javanica seeds. Biotropica 1: 58-66
Yap, S.K. 1981. Collection, germination and storage of
dipterocarp seeds. Malaysian Forester 44: 281-300.
Yap, S.K. 1986. Effect of dehydration on the germination
of dipterocarp fruits. In: Nather, J. (ed.) Seed problems
under stressful conditions. Proceeding of the IUFRO
Symposium, 168-181. Report no.12. Federal Forest
Research Institute, Vienna.
Chapter 4
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.
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
74
Seed Handling
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 (< 16o 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.
75
Seed Handling
Table 1. Likely periods for flowering and seed production of important Dryobalanops, Dipterocarpus, Shorea, and Anisoptera
species (Krishnapillay, unpublished).
Species
Months
J
Frequency
A
xxxx
xxxxxxx
biennial
Dryobalanops oblongifolia
xxxx
xxxxxx
biennial
Shorea leprosula
xxxxxxx
3-4 years
Shorea parvifolia
xxxxxxx
3-4 years
Dipterocarpus baudii
xxxxxxx
annual
Dipterocarpus costulatus
xxxx
Anisoptera scaphula
xxxx
Dipterocarpus kerrii
Shorea ovalis
4-5 years
xxxx
Shorea macrophylla
4-5 years
xxxxxx
xxx
xxxxxx
xx
xxxxxxx
Shorea platyclados
4-5 years
4-5 years
Anisoptera laevis
Shorea macroptera
xxx
xxxx
annual
3-4 years
3-4 years
xxxxx
3-4 years
Shorea acuminata
xxxxxxx
2-3 years
Shorea bracteolata
xxxxxxx
2-3 years
Shorea curtisii
xxxxx
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
3-4 years
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
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
Seed Handling
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
76
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
77
Seed Handling
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 16oC or above 32oC 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
78
Seed Handling
Table 2. Seed (fruit) weight and size indicators at harvest (Tompsett and Kemp 1996a, b).
Species
Shorea pinanga
Shorea macrophylla
Dipterocarpus grandiflorus
Dipterocarpus kunstleri
Dipterocarpus humeratus
Dipterocarpus cornutus
Dipterocarpus caudatus ssp. penangianus
Shorea palembanica
Shorea beccariana
Shorea fallax
Dipterocarpus chartaceus
Shorea smithiana
Anisoptera megistocarpa
Dipterocarpus tuberculatus
Shorea almon
Shorea ferruginea
Shorea robusta
Shorea trapezifolia
Shorea siamensis
Dipterocarpus tuberculatus var. grandifolius
Dipterocarpus gracilis
Shorea ovalis
Shorea gibbosa
Shorea argentifolia
Shorea macroptera
Shorea roxburghii
Anisoptera costata
Dipterocarpus intricatus x tuberculatus
Shorea parvifolia
Shorea selanica
Mean seeds per kilo
Mean length (mm)
Mean width (mm)
30
33
50
64
80
90
90
100
110
120
120
120
140
160
160
160
170
180
200
200
220
230
270
360
440
540
588
670
680
690
760
790
790
930
940
1100
1100
1195
1200
1200
1300
1300
59
n/a
58
46
55
35
20
n/a
29
26
36
26
n/a
36
n/a
n/a
30
30
28
29
27
27
n/a
38
26
15
n/a
16
26
n/a
n/a
15
17
n/a
17
n/a
19
16
11
24
19
17
32
n/a
38
26
43
29
19
n/a
28
23
23
23
n/a
24
n/a
n/a
20
20
22
17
20
23
n/a
30
13
14
n/a
9
16
n/a
n/a
13
11
n/a
11
n/a
10
8
11
17
8
10
1300
n/a
n/a
79
Seed Handling
Table 2. (continued) Seed (fruit) weight and size indicators at harvest.
Species
Mean seeds per kilo
Mean length (mm)
Mean width (mm)
1400
1400
1600
1800
1800
1900
1900
1900
2800
2900
2900
2900
4000
4000
4100
4200
5300
5500
5800
6300
9000
17000
45000
17
n/a
14
10
16
n/a
n/a
n/a
20
10
9
n/a
10
8
7
11
8
8
n/a
10
8
5
n/a
9
n/a
9
10
10
n/a
n/a
n/a
17
10
8
n/a
7
7
6
6
6
5
n/a
4
5
5
n/a
Dryobalanops rappa
Shorea faguetiana
Shorea laevis
Shorea leprosula
Shorea affinis
Shorea leptoderma
Shorea ovata
Dipterocarpus intricatus
Shorea obtusa
Hopea dryobalanoides
Vatica odorata ssp. odorata
Hopea parviflora
Shorea guiso
Hopea odorata
Hopea foxworthyi
Hopea ferrea
Hopea mengerawan
Hopea nigra
Vatica mangachapoi
Monotes kerstingii*
*: 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,
80
Seed Handling
Table 3. Mean insect infestation statistics for species received at Kew (Tompsett and Kemp 1996a, b).
Genus
Number of Number of
Mean
species
species
percent
infested
infestation examined
Dipterocarpus
35
10
10
Shorea
16
18
12
Hopea
Parashorea
Dryobalanops
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.
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.
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
15C (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.
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
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.
Species
Source
Optimum storage recorded

Days
Temp.
o
( C)
Germination
(%)
Medium
MC
(%)
Shorea platyclados
Tang (1971)
20
16
64
27
Hopea ferrea
Tompsett (1992)
300
16
40
30-50
Tompsett (1992)
317
18
46
45
Perlite
Shorea fallax
Tompsett (1992)
50
21
50
40
Sawdust
MC: moisture content.
Vermiculite
Mainly perlite
81
Seed Handling
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
15C 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 21-24C. 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 16C 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.
82
Seed Handling
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
16C, 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
Hopea odorata
Period of storage
(months)
9-12
Hopea helferi
Dipterocarpus cornutus
Shorea macrophylla
Shorea leprosula
6-9
Shorea acuminata
Shorea longisperma
Shorea parvifolia
8-9
Shorea ovalis
8-9
Shorea curtisii
8-9
Shorea platyclados
8-9
Shorea bracteolata
Shorea macroptera
Shorea maxwelliana
Shorea pauciflora
Dryobalanops oblongifolia
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 -196C, 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
Seed Handling
(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
-20C 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 -30C at the rate of 1C per minute, finally being
reduced to -196C by plunging into LN. After one week
the embryo axes were removed, thawed at 40C 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 -20C (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).
83
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 18C. 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 2C; for use in the long term, transport material at
as low a temperature as possible, but not below -20C.
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 20C 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
84
Seed Handling
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 18oC 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
18oC 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 18oC. 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 -20oC. 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
2C 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 oC -31oC. 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.
Seed Handling
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
85
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.
86
Seed Handling
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 governments
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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Pritchard, H.W., Tompsett, P.B., Manger, K. and Smidt,
W.J. 1995. The effects of moisture content on the
low temperature responses of Araucaria hunsteinii
seed and embryos. Annals of Botany 76: 79-88.
seeds. Malayan Forester 43: 290-308.
Schaffalitzky de Muckadell, J. and Malim, P. 1983.
Preliminary observations on harvesting, handling and
storage of seeds of some dipterocarps. Working paper
no. 18. FAO/UNDP-MAL/78/009. 42p.
Sen Gupta, J.N. 1939. Dipterocarpus (gurjan) forests
in India and their regeneration. Indian Forest Record
3, No. 4. 164p.
on the principal storage conditions of Hopea
hainanensis seeds. Scientia Silvae Sinicae 20: 225236.
Ng. F.S.P. 1981. Vegetative and reproductive phenology

of dipterocarps. Malaysian Forester 44: 197-216.
Sukwong, S., Dhamanitayakul, P. and Pongumphai, S.

1975. Phenology and seasonal growth of dry
dipterocarp forest tree species. Kasetsart Journal 9:
105-114.
Ng. F.S.P. 1984. Plant phenology in the humid tropics.

Research Pamphlet No. 96. Forest Research Institute
Malaysia, Kepong. 60p
Tang, H.T. 1971. Preliminary tests on the storage and

collection of some Shorea seeds. Malaysian Forester
34: 84-98.
Owens, J.N., Sornsathapornkul, P. and Tangmitcharoen,

S. 1991. A manual on studying flowering and seed
ontogeny in tropical forest trees. ASEAN-Canada
Forest Tree Seed Centre Project, Muak Lek, Saraburi,
Thailand. 134p.
Tompsett, P.B. 1983. The influence of gaseous

environment on the storage life of Araucaria
hunsteinii seed. Annals of Botany 52: 229-237.
Pence, V.C. 1992. Desiccation and survival of Aesculus,

Castanea and Quercus embryo axes through
cryopreservation. Cryobiology 29: 391-399.
Prasad, R. and Jalil, P. 1987. Observations on the seed
viability of sal (Shorea robusta). Indian Forester 113:
89-94.
Press, M.C., Brown, N.D., Barker, M.G. and Zipperlen,
S.W. 1996. Photosynthetic responses to light in
tropical rain forest tree seedlings. In: Swaine, M.D.
and Alexander, I. (eds.) The ecology of tropical rain
forest seedlings. Parthenon Press, London.
Pritchard, H.W. 1995. Cryopreservation of seeds. In:
Day, J.G. and McLellan, M.R. (eds.) Methods in modern
biology, Volume 38: Cryopreservation and freezedrying protocols, 133-144. Humana Press, Tolowa,
New Jersey.
Tompsett, P.B. 1985. The influence of moisture content

and temperature on the viability of Shorea almon, S.
robusta and S. roxburghii seed. Canadian Journal of
Forest Research 15: 1074-1079.
Tompsett, P.B. 1987. Desiccation and storage studies
on Dipterocarpus seeds. Annals of Applied Biology
110: 371-379.
of dipterocarp seeds. Seed Science and Technology
20: 251-267.
Tompsett, P.B. 1994. Capture of genetic resources by
collection and storage of seed: a physiological
approach. In: Leakey, R.R.B. and Newton, A.C. (eds.)
Proceedings of the IUFRO Conference Tropical
Trees: the Potential for Domestication and the
Rebuilding of Forest Resources, August 1992, 6171. HMSO, London.
Seed Handling
Tompsett, P.B. and Kemp, R. 1996a. Database of tropical

tree seed research (DABATTS). Database Contents.
Royal Botanic Gardens Kew, Richmond, Surrey. 263p.
Tompsett, P.B. and Kemp, R. 1996b. Database of tropical
tree seed research (DABATTS). User Manual. Royal
Botanic Gardens Kew, Richmond, Surrey. Includes two
3.5 computer disks. 28p.
Toy, R.J. and Toy, S.J. 1992. Oviposition preferences
and egg survival in Nanophes shoreae (Coleoptera,
Apionidae), a weevil fruit predator in South-east Asian
rain forest. Journal of Tropical Ecology 8: 195-203
Toy, R.J, Marshall, A.G. and Tho, Y.P. 1992. Fruiting
phenology and the survival of insect fruit predators: a
case study from the South-east Asian
Dipterocarpaceae. Philosophical Transactions of the
Royal Society. Biological Science 335: 417-423.
88
Troup, R.S. 1921. The silviculture of Indian trees, Vol. I.
Clarendon Press, Oxford. 336p.
Willan, R.L. 1985. A guide to forest seed handling with
special reference to the tropics. FAO Forestry Paper.
No. 20/2. Food and Agriculture Organisation, Rome.
Wood, G.H.S. 1956. The dipterocarp flowering season
in North Borneo, 1955. Malayan Forester 19: 193201.
Yap, S.K. 1986. Effect of dehydration on the germination
of dipterocarp fruits. In: Seed Problems Under
Stressful Conditions. Proceedings of the IUFRO
Symposium, 168-181. Report no. 12. Federal Forest
Research Institute, Vienna.
Chapter 5
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 forest1 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-
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.
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.
1
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 VazquezYanes 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
LATE-SUCCESSIONAL UNDERSTORY
(Gaertnera, Psychotria)
LATE-SUCCESSIONAL SUBCANOPY
(Calophyllum, Garcinia)
LATE-SUCCESSIONAL NON-DOMINANTS
(Durio, Ficus, Mangifera)
LATE-SUCCESSIONAL DOMINANTS
(Dipterocarpus, Dryobalanops, Shorea)
PIONEERS OF BUILDING PHASE

(Albizia, Alstonia)
PIONEERS OF GAP PHASE

(Macaranga, Trema)
MATURE
LATE SUCCESSIONAL
Canopy break-up
Stand canopy dominance. The breadth of the bar represents the degree of dominance in relation 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).
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.
MID-SUCCESSIONAL
Approximate point of canopy closure

Approximate peak in canopy exclusion
Initiation of groundstory
BUILDING
EARLY SUCCESSIONAL
Disturbance
GAP
PHASES OF STAND DEVELOPMENT
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).
91
92
(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,
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
(hymenoptera, hemiptera)
small
insects
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

germination and early survival
protection
for
require an increase in light

compared to understorey conditions)
satisfactory establishment and growth
(as
for
Seedlings
Seedling survival and establishment is usually site
according
to
particular
specific,
microclimatic and edaphic characteristics
biotic,
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. Beckers (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 (PN), 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 PN rate was a good measure of the
light tolerance of Shorea. The shade tolerant species had
maximum PN rates at relatively lower light intensity
compared to that of more light demanding species. Ratios
between rates of PN and E of species at their maximum
PN 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 PN 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!
Acknowledgements
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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Chapter 6
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
Hgberg 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
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
100
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
Thailand
Dry deciduous forest
Aniwat (1987)
Pen. Malaysia
Lowland rainforest
Singh (1966)
Kalimantan
Lowland rainforest
Smits (1987)
Pen. Malaysia
Rainforest
Mohd. Noor (1981)
Anisoptera
A. costata Korth. *(VAM also)
A. laevis Ridl.
A. marginata Korth.
A. oblonga Dyer
A. scaphula (Roxb.) Pierre
A. thurifera (Blco) Bl.
"
Luzon
"
"
Rainforest
Zarate et al. (1993)
Cotylelobium
C. malayanum Sloot.
C. scabriusculum Brandis
Pen. Malaysia
Dipterocarp arboretum
Hong (1979)
Sri Lanka
Lowland rainforest
de Alwis & Abeynayake (1980)
Aniwat (1987)
Dipterocarpus
D. alatus Roxb.
D. baudii Korth.
D. chartaceus Sym.
D. confertus Sloot.
D. cornutus Dyer
D. costatus Gaertn. f.
D. costulatus Sloot.
D. elongatus Korth.
D. gracilis Bl.
D. grandiflorus (Blco) Blco
D. hasseltii Bl.
D. hispidus Thw.
D. humeratus Sloot.
D. indicus Bedd.
D. intricatus Dyer.
D. kunstleri King
D. oblongifolius Bl.
D. obtusifolius Teysm. ex Miq.
D. sublamellatus Foxw.
D. tempehes Sloot.
D. tuberculatus Roxb.
D. verrucosus Foxw.
D. zeylanicus Thw.
Thailand
Semi-evergreen forest
Pen. Malaysia
Rainforest
Mohd. Noor (1981)
Pen. Malaysia
Hong (1979)
Kalimantan
Lowland rainforest
Smits (1992)
"
"
Bimaatmadja in Hadi et al. (1991)
Thailand
Aniwat (1987)
Pen. Malaysia
Rainforest
Mohd. Noor (1981)
Kalimantan
Lowland rainforest
Smits (1992)
"
"
"
"
"
Sri Lanka
Kalimantan
"
Lowland rainforest
"
"
Smits (1983)
Smits (1992)
Smits (1992)
India
Wet evergreen forest
Thailand
Alexander & Hogberg (1986)

Aniwat (1987)
Sarawak
Kerangas
Pen. Malaysia
Lowland rainforest
Singh (1966)
Thailand
Aniwat (1987)
"
"
Kalimantan
"
Lowland rainforest
Smits (1992)
Thailand
Aniwat (1987)
Pen. Malaysia
Rainforest
Sri Lanka
"
Mohd. Noor (1981)

Dryobalanops
D. aromatica Gaertn. f.
"
D. keithii Sym.
D. lanceolata Burck
Pen. Malaysia
Lowland rainforest
Singh (1966)
Kalimantan
Lowland rainforest
Smits (1992)
"
Java
"
"
Nuhamara et al. in Hadi et al.
(1991)
"
Sabah
Lowland rainforest
Unpublished data
"
Kalimantan
Lowland rainforest
Smits (1992)
D. oblongifolia Dyer
D. oocarpa Sloot.
Pen. Malaysia
Hong (1979)
Kalimantan
Lowland rainforest
101

Table 1. (continued) Dipterocarp species reported to be ectomycorrhizal based on root examination.
Genera Species
Location
Vegetation
Reference/Source
H. bancana (Boerl.) Sloot.
Java
H. dryobalanoides Miq.
H. ferrea Laness.
"
H. ferruginea Parijs
H. iriana Sloot.
H. jucunda Thw.
H. mengerawan Miq.
H. montana Sym.
H. nervosa King
"
"
H. nudiformis Thw.
H. odorata Roxb.
"
"
Kalimantan
Pen. Malaysia
Thailand
Pen. Malaysia
?
Sri Lanka
Kalimantan
Pen. Malaysia
"
Sabah
Kalimantan
Java
Pen. Malaysia
Thailand
Java
Lowland rainforest
Rainforest
Lowland rainforest
?
Lowland rainforest
Lowland rainforest
Rainforest
"
Lowland rainforest
Lowland rainforest
Rainforest

(1991)
Smits (1992)
Mohd. Noor (1981)
Aniwat (1987)
Singh (1966)
Ashton (1982)
Smits (1992)
Mohd. Noor (1981)
"
Unpublished data
Smits (1992)
Setiabudi in Hadi et al. (1991)
Mohd. Noor (1981)
Aniwat (1987)
H. parvifolia (Warb.) Sloot.

H. plagata (Blco) Vidal
H. sangal Korth.
S. India
Luzon
Kalimantan

Rainforest
Lowland rainforest
Zambia
"
Miombo
Zambia
Tanzania
Miombo
Miombo
Hogberg & Piearce (1986)

Hogberg (1982)
Pen. Malaysia
Lowland rainforest
Singh (1966)
Pen. Malaysia
"
Sabah
Lowland rainforest
"
Lowland rainforest
Mohd. Noor (1981)

"
Unbubl. data
?
Rainforest
Tupas & Sajise (1976)

Mohd. Noor (1981)
Nursery
Rainforest
Lowland rainforest
Rainforest
Lowland rainforest
Ogawa (1992a)
Mohd. Noor (1981)
Mohd. Noor (1981)
Smits (1987)
Hopea

(1991)
Zarate et al. (1993)
Julich (1985)
Marquesia
M. acuminata Gilg.
M. macroura Gilg.
"
Hogberg & Piearce (1986)

"
Monotes
M. africanus (Welw.) A.D.C.
M. elegans Gilg.
Neobalanocarpus
(Balanocarpus)
N. heimii (King) Ashton
Parashorea
P. densiflora Sloot. & Sym.
P. lucida (Miq.) Kurz.
P. malaanonan (Blco) Merr.
Pentacme
P. contorta (Vidal) Merr. & Rolfe Philippines
P. siamensis (Miq.) Kurz.
Pen. Malaysia
Shorea
S. academia (?)
S. acuminata Dyer
S. affinis (Thw.) Ashton
S. assamica Dyer
S. balangeran (Korth.) Burck
Kalimantan
Pen. Malaysia
Sri Lanka
Pen. Malaysia
Kalimantan
102

Table 1. (continued) Dipterocarp species reported to be ectomycorrhizal based on root examination.
Genera Species
Location
S. bracteolata Dyer
"
"
*(VAM also)
Vegetation
Reference/Source
Pen. Malaysia
Rainforest
Mohd. Noor (1981)
Kalimantan
Logged over forest
Suhardi et al. (1992)

Norani (pers. comm.)
Pen. Malaysia
Lowland rainforest
S. compressa Burck
Java
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
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)
"
"
Smits (1992)
S. lepidota (Korth.) Bl.
Pen. Malaysia
Lowland rainforest
Berriman (1986)
S. leprosula Miq.
Pen. Malaysia
Lowland rainforest
Singh (1966)
Kalimantan
Lowland rainforest
"
S. lamellata Foxw.
"
S. macrophylla (de Vriese) Ashton Sarawak
Chong (1986)
S. macroptera Sloot.
Lowland rainforest
Singh (1966)
"
Becker (1983)
Pen. Malaysia
S. maxwelliana King
S. mecistopteryx Ridl.
"
Hadi et al. (1991)
Indonesia
S. obtusa Wall.
Thailand
Aniwat (1987)
S. ovalis (Korth.) Bl.
Pen. Malaysia
Lowland rainforest
Singh (1966)
Kalimantan
Lowland rainforest
S. ovata Dyer ex Brandis
Pen. Malaysia
Rainforest
Mohd. Noor (1981)
S. palembanica Miq.
Java
Hadi et al. (1991)
"
S. parvifolia Dyer
"
S. pauciflora King
"
S. pinanga Scheff.
"
Pen. Malaysia
Rainforest
Mohd. Noor (1981)
Kalimantan
Lowland rainforest
Pen. Malaysia
Lowland rainforest
Singh (1966)
Kalimantan
Lowland rainforest
Smits (1992)
Java

Smits (1992)
Kalimantan
Lowland rainforest
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
Aniwat (1987)
S. scabrida Sym.
Sarawak
Kerangas
S. selanica (Lamk.) Bl.
Java
S. seminis (de Vriese) Sloot.

S. sericeiflora Fisher & Hance
"
Pen. Malaysia
"
Rainforest
Hibau in Hadi et al. (1991)

Mohd. Noor (1981)
103

Table 1.
(continued) Dipterocarp species reported to be ectomycorrhizal based on root examination.
Genera Species
Location
Vegetation
Reference/Source
S. siamensis Miq.
Thailand
Aniwat (1987)
S. smithiana Sym.
Kalimantan
Lowland rainforest
S. stenoptera Burck
Java
Setiabudi in Hadi et al. (1991)
Kalimantan
Lowland rainforest
Smits (1992)
Pen. Malaysia
Rainforest
Mohd. Noor (1981)
"
S. sumatrana (Sloot. ex Thor.) Sym.

S. talura Roxb.
"
"
"
S. teysmanniana Dyer ex Brandis
"
"
"
Vatica
Vatica sp. 1
Kalimantan
V. chartacea Ashton
Lowland rainforest
Smits (1992)
"
"
V. papuana Dyer ex Hemsl.
Pen. Malaysia
Lowland rainforest
Singh (1966)
"
V. rassak (Korth.) Bl.
Kalimantan
Lowland rainforest
Smits (1992)
V. sumatrana Sloot.
Java
Hadi & Santoso (1988)
V. umbonata (Hook. f.) Burck
Kalimantan
Lowland rainforest
Smits (1992)
S. India
Aberdeen greenhouse
Potted plant
Unpublished data
Vateria
V. indica L.
Vateriopsis
V. seychellarum Heim
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
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
Yasmans (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
106
in Sabah showed that increasing concentrations of

fertiliser resulted in increased growth rates but that
growth was reduced when 2000 mg of NH4NO3 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
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.
Research Priorities
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 BIOREFOR 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 isolates 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 FRIMCommission 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
Baos, 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.
Acknowledgements
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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Chapter 7
Pests and Diseases of

Dipterocarpaceae
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.
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
117
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).
118
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.
119
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
120
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
hosts organs except by the byssus, but develop an aerial
network which, when too excessive, can hinder the hosts
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
121
(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
122
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
123
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, RamPrasad 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
124
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).
2.
3.
Research Priorities
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.
4.
5.
6.
7.
Chemicals as well as biological controls should be

tested.
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.
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.
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.
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.
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).
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.
Acknowledgements
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. MauryLechon (Centre National de la Recherche Scientifique/
Universit 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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Chapter 8
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 worlds 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
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)
134
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 evergreen1a. Evergreen dipterocarpMalaysia, 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-evergreenNorth Thailand (part), Chittagong, Laos,
Cambodia, Vietnam (?) (part)
3. Tropical moist deciduousMaluku (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
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
135
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
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
136
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.
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.
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.
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.
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
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 7m3/
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 (WyattSmith 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 (2559 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
Sarawaks 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
139
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
(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,
140
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:
dbh class (cm)
10
20
30
40
50
60
70
80
90
Mean annual diameter

increment (cm/yr)
10-16
16-22
>22
0.44
0.45
0.55
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 70100%. 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
Liberation
Felling
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:
cm/yr
0.44
0.58
0.69
0.78
0.83
0.86
0.86
0.85
0.79
Location
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
Residual basal area

2
(m /ha)
Year 1 Year 2 Year 3 Year 4 Year 5
Peninsular
MalaysiaTekam F.R.
0.72
0.57
0.68
0.67
0.57
Peninsular
MalaysiaLabis F.R.
0.63
0.79
0.71
0.63
n/a
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):
Sarawak:
Mersing
Bako
Philippines:
Mindanao
Visayas
Luzon
Mean annual diameter

increment (cm)
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
141
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
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. >15o), 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 regions 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 societys
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
145
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
15o. 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.
2.
3.
4.
5.
6.
7.
8.
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
9.
that are more specific to the site (site categories,

floristic groups, etc.).
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.
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.
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.
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?
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.
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?
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?
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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regenerating dipterocarp forests in Malaysia. In:
Mergen, E. and Vincent, J.R. (eds.) Natural management
of tropical moist forests, 23-46. Yale University,
School of Forestry and Environment Studies, New
Haven, CT.
Tang, H.T. and Wadley, H. 1976. Report on the survival
and development survey of areas re-forested by line
planting in Selangor. Research Pamphlet no. 67. Forest
Tang, H.T. and Wan Razali, W.M. 1981. Report on growth
and yield studies in inland mixed indigenous forests
in Peninsular Malaysia. Forest Department Peninsular
Malaysia, Kuala Lumpur. 73p.
Thang, H.C. 1987. Forest management system of tropical
high forests, with special reference to Peninsular
Malaysia. Forest Ecology and Management 21: 3-20.
Tiwari, S.D.N. 1968. The concept of coppice with
reserve system. Indian Forester 94: 303-314.
Uebelhoer, K. and Hernandez, A.Z. 1988. Timber stand

improvement: a chance for the Philippine dipterocarp
forest. RP-German TSI program. FMB. Diliman,
Quezon City. 24p.
Vidal, J. 1979. Outline of ecology and vegetation of the
Indochinese Peninsula. In: Larsen, K. and HolmNielsen, L.B. (eds.) Tropical botany, 109-123.
Academic Press, New York.
Weidelt, H.J. 1996. Sustainable management of
dipterocarp forests - opportunities and constraints. In:
Schulte, A. and Schone, D. (eds.) Dipterocarp forest
ecosystems: towards sustainable management, 249273. World Scientific, Singapore.
149
Weidelt, H.J. and Banaag, V.S. 1982. Aspects of
management and silviculture of Philippine dipterocarp
forests. GTZ Publication no. 132. Eschborn, West
Germany.
for inland forests. Malayan Forest Record no. 23.
Forest Department, Kuala Lumpur.
Wyatt-Smith, J. 1987. The management of tropical
moist forest for the sustained production of timber:
some issues. IUCN/IIED Tropical Forest Policy Paper
no. 4. 20p.
Wyatt-Smith, J. and Foenander, E.C. 1962. Damage to
regeneration as a result of logging Malayan Forester
25: 40-44.
Chapter 9
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
(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).
Troups 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
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
152
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
Plantations
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 InnopriseFace 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
153
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
Plantations
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
154
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, P2O5 and K2O 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
Plantations
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
155
terms of crown structure, branching habit, growth
dynamics etc. Hall 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,
156
Plantations
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
Plantations
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.
157
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
Plantations
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 P2 O5 (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.
158
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, P2O5 and
K2O) 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.
Plantations
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
159
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). MouraCosta (1995) discusses a procedure of selecting best
genotypic material at the seedling stage, the so-called
Plantations
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
(Hall 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
160
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
Plantations
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 1-2 year old secondary forest while
sowing in cut lines proved a failure. The ecological role
161
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.
Plantations
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. djers
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
162
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:
Plantations
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
163
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.
Plantations
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-
164
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
Plantations
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 planters
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,
165
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,
Plantations
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.
166
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
Plantations
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
167
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 Durants 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
Plantations
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
168
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
Plantations
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
169
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
170
Plantations
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 IndianSoutheast 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
Plantations
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 (Brnig 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
171
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
172
Plantations
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.
Research Priorities
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.
173
Plantations
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.
Acknowledgements
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. HahnSchilling (Malaysian-German Project Forest
Mananagement Information System Sarawak), Dr. R.
Nussbaum (Socit Gnrale 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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Chapter 10
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
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 Polos 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
(C22 H34 O4), devoid of acid character as in copaiba (a
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
191
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, C24 H42 O2, M.P. 125o-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
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
(C10 H17OH, M.P. 209oC) 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 1428% 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 Buddhas 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 collectors 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.
Acknowledgements
We would like to thank Manuel Ruiz-Perez (Center for
International Forestry Research) for comments and suggestions for the improvement of this chapter.
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Scientific Index
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
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
200
Scientific Index
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
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
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
201
Scientific Index
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
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
Ehretia laevis, 165

ELAEOCARPACEAE, 9, 10
Elaeocarpus stipularis, 167
Endogone, 103
Endospermum malaccense, 167
Scientific Index
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
FAGACEAE, 95
Ficus, 91
Fragraea fragrans, 157, 161
FUNGI IMPERFECTI, 120, 122
Furcraea, 194
Fusarium, 120
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
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
203
Scientific Index
IMBRICATE group, 27, 28, 31, 32, 34

Imperata cylindrica , 161, 165
Indigofera teysmanii, 161
Isauxis section, 32
Isauxis sub-genus, 8
Isoptera section, 32
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
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
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
OCHNACEAE, 9, 10, 26
OCHNALES, 25
Oecophylla smaragdina, 117
Ougeineia oojeinensis, 165
204
Scientific Index
Ovalis section, 8, 32
Ovalis sub-group, 8, 32
Ovalis, 24
Ovoides section, 32
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
Quercus lucida, 167
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
Scientific Index
Rubroshorea, 8, 31, 32, 33

Rubroshoreae section, 15
Rugosae section, 32
Russula amatic, 155
Russula sp., 106
Russula, 104, 158
RUSSULACEAE, 104
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
Scientific Index
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
207
Scientific Index
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
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
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
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
208
Scientific Index
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
White Meranti group of Shorea, 59

Woburnia porosa, 16
Woburnia, 19
Yellow Meranti group of Shorea , 59
Xyleborus declivigranulatus, 119

Xyleborus pseudopilifer, 119
Zygomycetes, 103
General Index
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
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
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
210
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
pests, 117
silviculture, 151-2
burning, see fire
butter fat, 193
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
211
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
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
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
212
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
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
butter fat, 193
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
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
213
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
planting stock, 157
site preparation, 161-2
stand density regimes, 155
thinning regimes, 166
see also survival
gums, 191, 194
gurjan oil, 188-91
Guyana, 12, 14
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
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
agroforestry, 170
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
General Index
distribution, 15
oleoresins, 191
pests, 117
Indonesia, 15, 153
agroforestry, 170
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
Java, 192
diseases, 121
distribution, 19, 22
fruiting age, 168
mycorrhizas, 100, 101, 102, 103
underplanting, 161
Johore, 192
Kalimantan, 4, 12
agroforestry, 170
enrichment planting, 141
mycorrhizas, 100-3, 104
pests, 116, 118
stump plants, 160
wildings, 159
kanyin oil, 188, 189
Katanga, 14

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
215
General Index
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
agroforestry, 170
dammars, 192
Departmental Improvement Fellings, 143-4
diseases, 119, 121
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
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
underplanting, 161
weed vegetation, 165
wildings, 158-9
Malesia, 1, 134
distribution, 13
oleoresins, 191
seed research, 69
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
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
mixed forests, 89-98
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, 1434, 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
stump plants, 160
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
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
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
outplants, 103
ovaries, 7, 9, 11, 24, 34
overtopping vegetation, removal of, 164
oxygen levels in storage, 81
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
oleoresins, 190
pests, 117, 118, 119
seed research, 69
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
217
General Index
preservatives, 119, 122-3, 124

processing seeds, 79, 83-4
procurement of seeds, 75-7
propagation of seedlings, 157
rain-weeding, 165
re-establishment by natural regeneration, 134-5, 16870
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
mixed forests, 90-4
nursing phenomenon, 107
plantations, 151-64
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
plantations, 172-3
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
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
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
diseases, 119-20
dispersal, 48-9, 89
pests, 116
plantations, 156-7
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, 16970
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
218
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
planting stock, 157
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
219
General Index

habitats, 12
mycorrhizas, 100, 101, 104, 109
oleoresins, 191
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
stump plants, 160
subfamilies, 5-6, 7, 8, 30-3
subspecies, criteria for definition of, 29
suckers, 118
sucrose, 83
Sulawesi, 13, 134
Sumatra, 134
agroforestry, 170
dammars, 192
diseases, 121, 122
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
outplants, 103
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
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
agroforestry, 170
diseases, 119, 121, 122
mycorrhizas, 100, 101, 102, 103, 104, 108-9
oleoresins, 190, 191
pests, 117, 118, 119
phenological studies, 75
seed research, 69
thinnings, 165-8, 172
thiram, 81, 82, 120
thread blights, 121
threatened species, identification of, 52
thrips, 24, 48
timber, 1, 166
diseases, 122-3
pests, 118-19
tissue chemistry, 94-5
tissue culture, 67, 83, 160
topography, 92, 93
see also slopes
220
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
underplanting, 3, 161, 162

understorey, 23, 94, 163
Uniform System, 136
United Kingdom, 16, 69
United States of America, 16, 19, 23
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
Wallaces 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
yield, 139-41
camphor, 192
dammar gardens, 192
seed production stands, 156-7
Yunnan, 134
Zaire, 14
Zambia, 14, 101
zonation, 11, 12
zygotic embryos, 30

A Review Dipterocarps

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A Review Dipterocarps

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A Review of

Taxonomy, ecology and silviculture

1998 by Center for International Forestry Research

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

Center for International Forestry Research

Chapter 1. Biogeography and Evolutionary Systematics of Dipterocarpaceae

Chapter 2. Conservation of Genetic Resources in the Dipterocarpaceae

Chapter 3. Seed Physiology

Chapter 4. Seed Handling

Chapter 5. Seedling Ecology of Mixed-Dipterocarp Forest

Chapter 6. Root Symbiosis and Nutrition

Chapter 7. Pests and Diseases of Dipterocarpaceae

Chapter 8. Management of Natural Forests

Chapter 10. Non-Timber Forest Products from Dipterocarps

International Union of Forestry

Species Improvement Network

Timber Stand Improvement

The Center for International Forestry Research (CIFOR)

Appannah at Forest Research Institute Malaysia (FRIM),

Prof. Jeffrey Sayer

As a family of plants, Dipterocarpaceae may perhaps hold

Wyatt-Smith 1963), most other studies remain

As a consequence, much of the knowledge on

efficient use of resources? Several issues require further

Biogeography and Evolutionary

The history of Dipterocarpaceae botany, as understood

rise of ambient temperature and desiccation. Faced with

Presentation of the Family

Biogeography and Evolutionary Systematics of Dipterocarpaceae

Asian sub-division Dipterocarps

Monotoideae in Africa and South America. The position

Morton 1995). During the past decade the numbers have

Biogeography and Evolutionary Systematics of Dipterocarpaceae

Maguire et al. 1977, Maguire and Ashton 1980

Maury 1978, Maury-Lechon 1979a, b*

Kostermans 1978, 1985, 1989

Londoo et al. 1995

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

the data now available, and the collaboration of still active

aliform short sepals, capsule at length dehiscent or

Biogeography and Evolutionary Systematics of Dipterocarpaceae

Meijer and Wood

Biogeography and Evolutionary Systematics of Dipterocarpaceae

paniculate (compound raceme)

2-celled anthers generally dehiscing longitudinally

persistent sepals and calyx

generally 2 ovules/ cell

aliform fruit sepals

short-sepal fruit calyx

possibility of peltate scales on the twig

seeds with scanty endosperm

Biogeography and Evolutionary Systematics of Dipterocarpaceae

entirely transcurrent and

presence of columns of sclerenchyme

paired basal leaf nerves

hypodermis (+papillose lower epiderms)

1-2 layered hypodermis

hair (various within a section)

indumentum + complex anatomy petiole (Malvales type)