Durio 654 (001-134)

Durio - A Bibliographic Review
MICHAEL J. BROWN
Department of Plant Science
MacDonald College, McGill University,
Quebec, Canada
INTERNATIONAL PLANT GENETIC RESOURCES INSTITUTE

Office for South Asia, c/o NBPGR, Pusa Campus
New Delhi 110 012, India
First published 1997
© International Plant Genetic Resources Institute
The International Plant Genetic Resources Institute (IPGRI) is an autonomous international

scientific organization operating under the aegis of the Consultative Group on International
Agricultural Research (CGIAR). The International status of IPGRI is conferred under
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Congo, Costa Rica, Cote d’lvoire, Cyprus, Czech Republic, Denmark, Ecuador, Egypt,
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Mauritania, Morocco, Pakistan, Panama, Peru. Poland, Portugal, Romania, Russia,
Senegal, Slovak Republic, Sudan, Switzerland, Syria, Tunisia, Turkey, Uganda and
Ukraine. IPGRI’s mandate is to advance the conservation and use of plant genetic
resources for the benefit of present and future generations. IPGRI works in partnership
with other organizations, undertaking research, training and the provision of scientific
and technical advice and information, and has a particularly strong programme link
with the Food and Agriculture Organization of the United Nations. Financial support
for the research agenda of IPGRI is provided by the Governments of Australia,
Austria, Belgium, Canada, China, Denmark, Finland, France, Germany, India, Italy,
Japan, the Republic of Korea, Luxembourg, Mexico, the Netherlands, Norway, the
Philippines, Spain, Sweden, Switzerland, the UK and the USA, and by the Asian
Development Bank, CTA, European Union, IDRC, IFAD, Interamerican Development
Bank, UNDP and the World Bank.
This publication has been funded by the International Plant Genetic Resources Institute.
Citation
Michael J. Brown. 1997. Durio - A Bibliographic Review (R.K. Arora, V. Ramanatha
Rao and A.N. Rao, Editors). IPGRI office for South Asia, New Delhi.
ISBN 92-9043-3 18-3
Copies can be had from:

IPGRI Office for South Asia
c/o NBPGR, Pusa Campus,
New Delhi 110012, India
IPGRI Regional Office for Asia, the Pacific and Oceania

PO Box 236, UPM Post Office
43400 Serdang, Selangor Darul Ehsan
Malaysia
IPGRI Office for East Asia

C/o CAAS, No. 30 Bai Shi Qiao Road, Beijing 100081,
Peoples Republic of China
Foreword
Under the IPGRI Project on ‘Promoting Conservation and Use of

Tropical Fruit Species in Asia,’ much information on status of plant genetic
resources has been synthesized for major (mango, citrus, rambutan) and minor
(jackfruit, litchi, durian) fruits of South, Southeast and East Asia. This information
gathering particularly relates to distribution, extent of diversity, status of germplasm
collection, characterization, evaluation, documentation, conservation and
utilization. Fifteen such reports prepared by national experts for key national
programmes/countries holding rich diversity in these tropical fruits have been
brought out by the IPGRI-APO; six on mango (Bangladesh, India, Indonesia,
Philippines, Thailand and China), three on citrus (India, China, Japan), two
each on rambutan and durian (Thailand and Malaysia), one on jackfruit
(Bangladesh), one on litchi (China). Dissemination of this well synthesized
information by IPGRI will promote further the conservation and use of these
crop genepools.
Among the native tropical underutilized fruits of promising potential,
durian assumes great importance particularly in Malaysia, Thailand and
Indonesia. Global interest in this fruit is catching up fast and its commercial
prospects increasing. With promising clones now available and considering
the export potential of durian, more area under its cultivation has led to loss
of primitive diversity and wild/semi-wild/domesticated species. Further, research
and development efforts are still required to understand durian flora1 biology
and physiology of fruit set, and assessing wild and domesticated primitive
genepools for their usefulness in diseases/pest resistance, physiological stress
and other attributes imparting better characteristics. Specific treatises dealing
with studies on taxonomy, genetic diversity, flora1 biology, breeding,
improvement, conservation and utilization aspects are wanting.
In the above context, the detailed synthesis provided on durian by
Dr. Michael J. Brown is welcomed. It is perhaps for the first time that such
a comprehensive account has been attempted on an underutilized fruit crop
genepool. This publication on Durian which is a review of bibliography of
about 1000 references, provides a well-synthesized information on different
aspects of Durio. It deals with its origin, history, taxonomy of wild and
cultivated species, identification, morphology of flower and fruit and other
organs, its edibility, composition and uses, nutritional aspects, medicinal and
iii
toxicological prospects, seed physiology, pollination biology, ecology, forest
resources, crop improvement efforts, agronomic requirements, propagation,
cultivation and maintenance, post harvest technology, genetic resource and
conservation.
IPGRI-APO has put in tremendous efforts to bring out this monograph
suitably edited by Drs. R.K. Arora, Ramanatha Rao and A.N. Rao, and
published by IPGRI Office for South Asia, Pusa Campus, New Delhi. IPGRI
is happy to be able to publish this work by Dr. Michael J. Brown, as it greatly
enriches knowledge on the durio genepools. Also, IPGRI supports the
dissemination of such information on underutilized fruits, their R&D needs
and above all promoting their conservation and use.
I am confident that this publication will generate further interest in
this crop, and will be useful to researchers and concerned institutes to strengthen
their research and development needs on durian.
Dr. Kenneth W. Riley

Regional Director
IPGRI Regional Office for Asia, the Pacific and Oceania
Serdang, Malaysia
iv
Preface
My desire to write this book grew out of an innate and deeply rooted
concern to understand the durian. In the late 1980s, a vigorous search of
scientific abstracts led me to believe that a bibliography on the subject would
only fill 1 or 2 type-written pages; the idea that a bibliographic review
resulting in a book would be necessary to fully examine the subject was
unthinkable. Today, I would estimate that approximately 1200 research
articles, book chapters, conference abstracts etc. have been produced which
pertain to durian. At the current rate of increase, this figure will very likely
double over the next 20 years. Thus, we stand at a crossroad. If our future
researches are really to advance our knowledge forward, we need to come to
terms with what we already know.
Despite the preponderance of information that exists on durian,

numerous obstacles are present for those who wish to study them. Much of
the literature presented in the bibliography of this book has never been
abstracted. This, and many of the items that are abstracted, have been
exceedingly difficult (and sometimes prohibitively expensive) to obtain. Thus,
in many areas, a researcher cannot be reasonably expected to have discovered
and read much of the pertinent background information. In numerous instances,
this has led to the repetition of experiments and has stood in the way of an
appropriate focus around which further research projects could be built up.
Currently, no comprehensive bibliography on durian literature exists

for the researcher, which is difficult to comprehend given the economic
importance of this fruit to several nations. The goal of this work is two-fold.
Firstly, it is intended to remedy the information management problems just
described. Secondly, it is hoped that, by means of a review, those issues
which are most in need of further study will be brought to light. The
diversity of research conducted has raised several very interesting lines of
enquiry; while the obscurity and fragmentary nature of many studies has led
to the premature acceptance of hypotheses.
The research conducted on some of the world’s major crops (rice,

wheat, maize, etc.) compared with that on durians is like a mountain to a
mole-hill. The number of fields of scientific enquiry that have been touched
upon by this mole-hill is rather staggering, and leaves me to wonder what
marvels we could unearth had we the whole mountain.
V
In this work, great pains have been taken to give full credit to all the
authors. Much time and effort have been devoted to obtaining original research
articles to confirm statements and ideas presented in more general reviews. In
cases where original results were published using the Imperial system of
measurements, a metric equivalent is presented parenthetically. Where research
on species, that are no longer taxonomically recognized, is discussed, parenthetic
notes are given to help clarify their identity.
I can in no way take full credit for the massive amount of effort that
went into compiling the bibliography. I wish to extend thanks to the numerous
libraries and institutions without whose aid in supplying original works and/
or photocopies, this book would not have been possible: University of Guelph
Library, Canada; Universiti Pertanian Malaysia Library; National Library of
Malaysia; Malaysian National Agricultural Library; Management Information
Systems Division, MARDI, Malaysia; University of Los Banbs Library,
Philippines; Kasetsart University Library, Thailand; Thailand Institute of Scientific
and Technological Research; Hunt Botanical Library, Pennsylvania; The Library
of the Herbarium Universitatis Florentinae, Italy; The Library of the New
York Botanical Gardens; The Library of the Jardin Botanique National de
Belgique, Belgium; The British Library (Oriental and India Office Collections);
Centre for Scientific Documentation and Information, Indonesia; SEAMEO-
BIOTROP, Bogor, Indonesia; Forest and Nature Conservation Research and
Development Centre, Bogor, Indonesia; Library of The Royal Botanic Gardens,
Kew, England; Lloyd Library and Museum, Cincinnati USA; and The Solomon
Islands National Library.
I am grateful to the many authors who supplied reprints of their
research articles. I am indebted to the friendship and excellent library skills
of Mr. David Bantroch, which greatly enhanced the scope and content of the
bibliography. I am also indebted to those who graciously supplied or helped
obtain copies of articles from different locations across the world: Dr. Gordon
Brown, Carol Bowes, Paul d’Amboise, Peter Toorop and Dr. Andrew Powell.
I also express my gratitude to the International Plant Genetic Resources Institute
for supporting the publication of this work, and to Drs. R.K. Arora, V. Ramanatha
Rao and A.N. Rao for editing the manuscript.
Finally, I would like to acknowledge the invaluable assistance of
those who helped translate articles or parts thereof: Nusin Brown for translation
of Turkish items; Aldo De Moor, Dr. Annette Nassuth and Peter Toorop for
translation of Dutch and German items; Dr. Wataru Mitsuhashi for the translation
of Japanese items, Chumnum Wongmanee for translation of Thai articles and
my wife Nathalie Bourgouin-Brown for translation of several French documents.
Michael J. Brown
vi
Contents
Foreword iii
Preface V
Contents vii
Introduction
Taxonomic History 2
Origin of the word durian; Durian poetry; Historical works;
Authority for Durio zibethinus; Formation of the modern
concept of Bombacaceae; Wild species of durian; Post
Kostermans (1958b) treatment; Keys to the species; Future
taxonomic work
Morphology 22
Durian theory; Floral morphology; Pollen morphology; Fruit
morphology; Fruit teratology; Other teratologies; Ovule and
seed morphology and development; Leaves; Roots; Tree
architecture; Chromosome number
Edibility, Composition and Uses of the Fruit 34
Edibility; Nutritive constituents; Fatty acids; Smell;
Miscellaneous uses of fruit
Medicinal and Toxicological Properties 51
Durians and alcohol; Medicinal properties; Febrifugal and
anti-malarial properties; Vermifugal properties; Treatment
of jaundice; Diabetes; Aphrodisiac; Miscellaneous medicinal
properties
Seeds 56
Mature seed constituents; Culture of seeds and seed size;
Viability; Germination
Pollination Biology 61
Anthesis; Natural pollinators; Early ovary abscission vs
premature fruit drop; Self-incompatibility; Heterostyly;
Mechanisms of self-incompatibility; Empirical evidence;
Premature fruit drop; Manipulating premature fruit drop;
Leaf flushing; Species which feed upon durians; Natural
dispersal of seeds
Fruiting Seasons 69
Maturation of buds; Environmental effects on flowering;
vii
Manipulating seasonality
Ecology, Origin . and Distribution 72
Centre of diversity; Wild form of Durio zibethinus; Attempts
at introduction
Clones 77
Clonal selection and hybridization; Clonal identification
Nursery Care and Cultivation 81
Seeds; Branch pruning; Root pruning; Application of fertilizer;
Soil conditions; Water relations; Transplanting; Intercropping
Post-harvest Technology
Grading; Shipping and cold storage of fruit; Packaging; 87
Ripening of fruits; Effects of atmosphere; Plant growth
regulators; Post-harvest technology; Processed food products
and their packaging
Forestry Aspects 94
Timber characteristics of durio species and close relatives;
Boschia griffithii Mast. [=D. griffithii (Mast.) Bakh.];
Coelostegia griffithii Mast.; Cullenia excelsa Wight; Durio
carinatus Mast.; Durio dulcis Becc.; Durio kutejensis (Hassk.)
Becc.; Durio ?lowianus Scort. ex King; Durio malaccensis
Planch. ex Mast.; Durio ?oblongus Mast.; Durio oxleyanus
Griff.; Durio testudinarum Becc.; Durio zibethinus L.,
Durability of ‘Durian’ timber; Uses of ‘Durian’ timber;
Properties of ‘Durian’ timber; Wood anatomy
Major Diseases, Parasitism and Associated Organisms 100
Bacteria; Fungi; Lichens; Algae; Ferns; Angiosperms; Insects;
Nematodes; Other animals; Hyperparasitism in durians
Vegetative Propagation of Durians 111
Etiolated shoot method; Double root system; Approach grafting;
Inarching; Top grafting; Budding techniques; Other asexual
propagation techniques; Grafting to other species; Advanced
planting material; ‘Hybrid’ Durians
Economics and Prospects for Development 117
Genetic Resources and Conservation 120
Bibliography 126
viii
Durio — A Bibliographic Review
Introduction
The genus Durio is native to South East Asia with its centre of
diversity in Borneo (Mendoza 1941; Lim 1990). The genus comprises
approximately 30 known species, of which only Durio zibethinus is cultivated
for its fruits to a great extent (Lim 1990). As a rain-forest tree, it typically
attains heights of 30-40 m (Tidbury 1976) and diameters of 2-2.5 m, but the
cultivated varieties in an orchard, especially when grafted, grow no higher
than 12 m (Malo and Martin 1979). Although relatively unknown to the
western world, the durian is a valuable commodity in South East Asia, and
has had a profound effect on the history and culture of that part of the world.
James Low (1836) recorded that the king of Ava had fruits transported to him
at Amerapoora, ‘by relays of horsemen, and by boats pulled by 40 or 50
men’. The durian fruit's reputation precedes it wherever it goes: ‘durian is to
fruit what limburger is to cheese and pornography is to literature’ (Anon.
1979a).
In Malaysia, the value of durian exports alone accounted for over

40% of total fruit exports in 1989 (Ali 1993). Subhadrabandhu et al. (1991)
state that in Indonesia, the rice harvest suffers if it happens to coincide with
durian season, the harvesters being more interested in the consumption of
durians than in the harvest of rice. There are quite a few recognized clones
of durian, whose fruits vary in size, shape, smell, colour, texture and taste.
Some clones (D2, D98, etc.) are much sought after and fetch a very high price
in the market.
Although little research has been carried out in the past on durian,
this trend is slowly starting to change as plans to improve the quality and
consistency of durian fruits develop. This is evidenced by the success of the
recent release of the first ‘hybrid’ durians in Malaysia (Othman 1991).
Studies, which have been conducted on durian, have been scattered

through various disciplines including chemistry, ecology, entomology, food
science, forestry, medicine, pathology, systematics and even zoology. These
studies have often been published in obscure or difficult to locate publications,
and not properly abstracted. Despite what can only be described as exhaustive
efforts to obtain and examine every book or article that supposedly pertained
to the genus Durio, some items were unobtainable. A fair number of erroneous
or non-existent research papers have also been referenced for their relative
Durio — A Bibliographic Review 2
importance. For these reasons, a comprehensive bibliography for durian is

presented at the end of this work. This bibliography contains only items of
which originals or copies have been obtained and examined by the author.
Taxonomic History
The durian has influenced various cultures of South East Asia for
millennia, but has only been known to the western world for about 600 years.
In this section, attempt is made to trace the scientific description of durian
from its earliest origins to the present day. Early descriptions and knowledge
were largely of a morphological and taxonomic nature. Although accounts on
the effects of the fruit on human physiology abounded, our knowledge of
these have expanded enormously in the last few decades and thus this aspect
will be discussed in a separate section. Tracing the origins of durian research
to their beginnings not only allows us to place certain information in proper
historical perspective, but also an understanding of how the current taxonomic
state of affairs, and perhaps much of the confusion, has arisen.
Origin of the word durian : The word durian (Durio) without doubt originates
from the Malay word ‘duri’ which means spine (Don 1831). The word ‘zibethinus’
is a reference to the Indian civet cat Viverra zibetha. Don (1831) provided the
following information: ‘the fruit is used as bait to entrap the civet-cat, which
is very fond of it; hence the specific name’. Others have suggested that zibethinus
refers to the smell of the fruit which is it’s (and the civet cat’s) most legendary
characteristic (Hawson 1983; Watson 1984). The accuracy of this comparison
is perhaps best summed up by Barrett (1912). ‘According to the specific name
zibethinus, the fruit should osphresiologically remind one of the civet cat; the
writer, however, after having seen and smelled live civets in Mozambique,
does not concur in this idea.’ Nevertheless, the fruit is occasionally referred
to as the civet cat fruit (Gamble 1881; Watt 1890; MacMillan 1909, 1912;
Anon. 1952; Singh et al. 1983). The Latin name was coined by Linnaeus who
apparently never encountered an actual specimen of a durian, and based his
description entirely upon that provided in the ‘Herbarium Amboinense’ (König
1804). As Rumphius (1741) refers to the use of durian fruits to catch civet
cats in his ‘Herbarium Amboinense’, it seems likely that this was the reason
for the name. De Candolle (1824) also states that the name arises from the
fact that civets eat durians.
De Clercq (1909) enumerated many vernacular names for durians

and Heyne (1950) enumerated well over 50 names used in various parts of the
Malay archipelago. Most of these are close variants of ‘durian’. Endert (1927b)
suggests that the conservation of the word ‘durian’ in different native Indonesian
languages probably indicates its early spread by the Malays.
Many authors give Malay vernacular names for the various species.
Malay names can often be quite useful in aiding identification (Corner 1988);
the taxonomy of many of the wild species of Durio is still in some disarray,
and it is quite likely that the Malay names are more reliable in some instances.
By far the most comprehensive, and probably most accurate, list of vernacular
names for different species of durian are those given by Kostermans (1958b).
In this book, the word ‘durian’ will be used to refer only to D.

zibethinus, except in its use as a forestry term. In this sense, it has historically
been applied to a mixture of different species, and perhaps even related genera.
Where any confusion could arise, I have endeavoured to refer to exact species.
Furthermore, where the taxonomy is confusing, I have referred to species as
defined in most comprehensive monograph by A.J.G.H. Kostermans (1958b)
[=sensu Kostermans 1958b], or otherwise, as appropriate.
Durian poetry : For whatever reason, the durian has appealed to the artistic
side of people for a long time. Many Malay idioms contain references to
durians (Kostermans 1958b). There are also several published poems pertaining
to durians (Whiteside 1914; Slate 1974; Chin 1979; Chin 1980a; Bantroch,
1995). A short story simply entitled ‘Durian’ by A.R. Roces (1949) delightfully
conveys the not-so-subtle nature of durian fruits to the reader.
Historical works : The durian appeared in pre-Linnaean literature as early as

the 16th century although erroneous information abounded. In 1741, Rumphius’
‘Herbarium Amboinense’ was finally published, and provided the most thorough
account of durians for over 100 years. Descriptions of durians in Linnaean
works, and those that followed it, have relied almost entirely on information
gleaned from this work.
The earliest European description of the durian is perhaps that of

Nicolo Conti who travelled in South East Asia at the beginning of the 15th
century (Bracciolini 1857). Fragoso’s ‘Discursos’ of 1572 offered a two-page
Spanish description of durians ‘Doriones’. Some details about durians were
also given in several works of Garcia De Orta in the late 16th century. Acosta’s
(1585) ‘Trattato’ briefly described durians and contained a very stylized figure
of a tree bearing fruit. Daléchamps’ (1586) ‘Historiae Generalis Plantarum’
contained a Latin description which is a near translation of the description
from Acosta’s Trattato of 1585 and bears a figure of the same stylized tree.
This figure was also redrawn in Boym’s ‘Flora Sinensis’ of 1656. Paludan in
Linschoten (1592) published a two-page description which appeared again in
one form or another in several later publications. An English translation of

this work is also presented (Anon. 1851).
Dodoens (1608) published his ‘Cruydt-Boeck’ which included a German
translation of Acosta’s description of durians from his ‘Trattato’ of 1585.
Pyrard (1619) stated that ‘The durion tree nearly resembles a pear
tree in size; the fruit is as big as a melon. The Indians esteem this fruit to be
one of the best and daintiest in the Indies. To those who are unaccustomed to
it, it is disagreeable, having a stink like that of our onions, but the taste is far
more excellent.’
Bauhin (1623) listed durians (duryaoen) in his ‘Pinax’, and De Bondt
(1658) presented a Latin description of the physiological effects of eating
durian and a drawing of the fruit which was by far the most accurate published
to date. Tavernier mentioned that durians are found growing in Siam (ca.
1676), and John Ray’s ‘Historia Plantarum’ of 1693 contains a one page Latin
description of durian fruits, largely copied verbatim from Acosta’s ‘Aromatum’,
with a few added notes. According to Hamilton: ‘The Durean is another
excellent fruit, but offensive to some peoples noses, for it smells very like
human excrements, but when once tasted, the smell vanishes. The skin is
thick and yellow, and within is a pulp like thick cream in colour and consistence,
but more delicious in taste. The pulp or meat is very hot and nourishing, and
instead of surfeiting, they fortify the stomach and are a great incentive to
Wantonness’ (Hamilton 1727).
Rumphius’ ‘Herbarium Amboinense’ was finally published in 1741.
This encyclopedic work contains several pages devoted specifically to durian
fruits. Although the identity of several plants described in this work remain
controversial, the plant described as ‘durioen’ in this book definitely represents
D. zibethinus (Buchanan-Hamilton 1824). The text of the ‘Herbarium
Amboinense’ was originally written in Dutch, and was published side by side
with a Latin translation. No English translation ever appears to have been
published. The different species mentioned by Rumphius are merely varieties.
Rumphius did manage to distinguish successfully between the durian and the
soursop (Annona muricata), which had been muddled together since the time
of Garcia De Orta. Rumphius’ description included a lengthy discussion on
the digestive effects of eating durian and the germination of the seeds. His
description was accompanied by a very accurate plate depicting the flowers,
fruit and a branch of a tree. The genus was rendered into Linnaean systematics
by Adanson (1763) in his ‘Familles des Plantes’, based on Rumphius’ description
in the ‘Herbarium Amboinense’.
Authority for Durio zibethinus : The taxonomy of durians brings to light
many of the taxonomic problems. The genus Durio, as stated by Chevalier

(1934), was created by Rumphius (1741) in his ‘Herbarium Amboinense’, and
was rendered into Linnaean systematy by Adanson (1763).
The type species Durio zibethinus is attributed to (Murr.); J.A. Murray,

however, some authors also attribute the species to Linnaeus (L.). Infrequently,
other (erroneous) authorities are encountered, i.e., Chattaway (1933) referred
to Durio zibethinus DC. De Candolle (DC.) is obviously not the correct
authority as A.P. De Candolle’s Prodromus of 1824 is predated by over 50
years of valid taxonomic references to this species. Furthermore, in this
work, De Candolle himself cited Linnaeus as the authority for the species.
Thus, the question remains as to the correct valid authority. As there has
been some confusion of this matter in taxonomic and other works, and no
consensus appears to have been reached, a full investigation of the issue
was warranted.
The authority L. appeared several times in early taxonomic literature.

Its first appearance seems to have been in Willdenow (1800) in the 5th edition
of ‘Species Plantarum’, which lists ‘Durio zibethinus Syst. Veg. 698.’ as a
species. This is a reference to page 698 of the 14th edition of ‘Systema
Vegetabilium’ of 1784 by J.A. Murray upon which Durio zibethinus is listed.
However, this species was described several years earlier on page 581 of the
13th edition of Linnaeus’s ‘Systema Vegetabilium’ published in 1774, also
edited by J.A. Murray. This 13th edition is without doubt the first ‘valid’
publication of this species, however, Willdenow’s error of accrediting the first
publication to the 14th edition was copied by several future authors, and
found its way into several important taxonomic works: A.P. De Candolle
(1824) described ‘Durio Linn. Syst. 698. non Adans.’ in his ‘Prodromus’.
George Don (1831) lists ‘Linn. Syst. 698. but not of Adans. DC. prod. 1 p.
480.’, a reference most-likely copied directly from De Candolle’s work.
Furthermore, Endlicher (1840) in ‘Genera Plantarum’ cited ‘Linn. Gen. n.
698.’; and Koorders and Valeton (1895) refer to ‘Durio zibethinus Linn Syst.
698’. Kurz (1874) also referred to Durio zibethinus L. Sp. Pl. 698, however,
the tree he was most likely describing was Cullenia ceylanica; furthermore,
he describes the same species as ‘D. zibethinus DC.’ in his ‘Forest Flora of
British Burma’ (1877).
As stated previously, the earliest valid publication on the species is

that appearing on page 581 of the 13th edition of Linnaeus’s ‘Systema
Vegetabilium’ published in 1774, and edited by J.A. Murray. This work has,
in fact, often been cited as the first valid publication with a twist. Ridley
(1922), for instance, referred to ‘D. zibethinus Linn. Syst. Nat. edn. xiii. 581’;
and Wyatt-Smith (1953a) states that ‘D. zibethinus Murr. [Syst. Nat. Veg.
edn. 13, 581 (1774)]’ is the earliest described species. They are both correct.
They differ in that Ridley (1922) attributes this work to Linnaeus, while
Wyatt-Smith (1953a) attributes the same work to Murray.
As Durio is not mentioned in the 12th or earlier editions of ‘Systema

Naturae’ (later ‘Systema Vegetabilium’), the only remaining question is to
whom new species appearing in the 13th edition of ‘Systema Vegetabilium’
of 1774 should be attributed, Murray or Linnaeus. Linnaeus himself was
solely responsible for these changes, and thus is the correct authority 1.
Formation of the modern concept of Bombacaceae : In early times, there

was some confusion between durian and the soursop (Annona muricata), both
of these species having spiny green fruit. Weinmann (1739) considered the
durian to belong to the Castaneae as its fruit was reminiscent of the horse
chestnut. Rumphius (1741) was more astute and recognized similarities between
the flowers of durian and those of kapok (Ceiba pentandra), another
Bombacaceous tree.
De Jussieu (1789) placed durian in the Capparideae, largely due to

the presence of scales on the underside of the leaves, and the erroneous belief
that the ovary is stipitate. König (1804) was the first botanist to examine the
flowers of durian in detail. This led him to transfer Durio to the Malvaceae.
What is now the family Bombacaceae was originally treated as a

tribe (Bombaceae) of the Malvaceae (Bentham 1862). This group of trees
remained as a tribe of the Malvaceae consisting of three subtribes (including
the Durioneae) in Baillon’s Natural History of Plants and Maxwell T. Master's
1874 monograph. Schumann (1895) created the family Bombacaceae, elevating
1
According to what is unquestionably the most thorough source of information on taxonomic
literature, "the botanical novelties in this [13th] edition still stem from Linnaeus and must be
attributed to him. Murray acted here as editor" (F. Stafleau and R.S. Cowan 1981), Taxonomic
Literature. 2nd edition. Volume 3). Yet, Farr et al. (1979) in the ‘Index Nominum Genericorum’,
who attempted, after indepth research, to compile a list of valid plant genera, type species and
their authorities, cites D. zibethinus Murray (Syst. Veg. edn. 13:581. 1774) as the type.
Murray was unquestionably the editor of the 13th edition of ‘Systema Vegetabilium’;
Murray received a copy of the manuscript for the 13th edition from Linnaeus in 1771, who
asked him to find a publisher for it in Germany (H. Goerke 1976), Linnaeus and the Murray
family, Taxon 25(1), 15-19). Although Murray was awarded the status of Editor, his function
was only to obtain a publisher for this work. Accordingly, the title page bears the Latin
inscription "Accessionibus et emendationibus novissinus manu perillustris auctoris scriptis" .
Thus, the common durian is correctly referred to Durio zibethinus L.
the three subtribes of the former Bombaceae (Adansonieae, Matisieae, Durioneae)

to tribal status. The genus Durio belongs within the Durioneae. These divisions
have been supported by differences in leaf morphology between them.
Adansonieae have palmately compound leaves; Matisieae have simple leaves
with palmate venation (termed Quararibees by Dumont 1887); and the Durioneae
is characterized by simple entire penninervate leaves. This system has been
more or less followed to the present day, although the durian is occasionally
(and erroneously) included within the Sterculiaceae; the Bombacaceae actually
share more anatomical similarities with the Malvaceae (especially the genus
Hibiscus) than they do with the Sterculiaceae.
Wild species of durian : The taxonomy of the common durian is convoluted

enough, but the real confusion belongs to the wild species. Several attempts
have been made to sort out the species, each in turn has in time been thoroughly
reworked. The most recent monograph, and probably not the last, is that of
Kostermans (1958b). Korthals (1842) introduced the new genus Boschia
containing one species, which differed from Durio most significantly in having
anthers that dehisced by means of pores, whereas D. zibethinus has anthers
that dehisce by slits. Bentham and Hooker’s ‘Genera Plantarum’ of 1862
listed one species of Durio, one species of Lahia and 2 species of Boschia (all
of which are now considered under Durio). One hundred years later, Masters
(1874b) recognized 7 species of Durio as well as Lahia kutejensis and four
species of Boschia.
In 1889, Beccari produced volume three of ‘Malesia’ which included

a monograph of the genus Durio. Beccari’s monograph was undoubtedly the
best and most thorough treatment yet produced containing lengthy descriptions
and excellent diagrams. The value of this work was undoubtedly enhanced by
his familiarity with field material as well as herbarium specimens. This monograph
described 14 species of Durio, 4 of Boschia, 7 of Neesia, 3 of Coelostegia
and Cullenia excelsa Wight. The genus Lahia was done away with and the
species Lahia kutejensis Hassk. became Durio kutejensis (Hassk.) Becc., and
has remained so until the present.
King (1891) described only 6 species of Durio (D. zibethinus, D.

lowianus, D. malaccensis, D. testudinarum, D. wrayii, D. oxleyanus and Boschia
griffithii) in his enumeration of the flora of the Malay peninsula. The new
species D. wrayii King was first described in this work. Ridley (1922) enumerated
9 distinct species of Durio and Boschia griffithii.
The genus was thoroughly reworked by Bakhuizen Van Den Brink in

his ‘Revisio Bombacacearum’ of 1924. He described 14 species of Durio as
did Beccari, however, only 9 of these species were the same. Of the several
new species enumerated by Beccari, 5 were dropped in Bakhuizen Van Den

Brink’s work (namely, D. graveolens Becc., D. dulcis Becc., D. gratissimus
Becc., D. affinis Becc., D sumatranus Becc.); additionally, 5 new species
were erected (D. griffithii, D. excelsus, D. mansoni, D. ceylanicus and D.
lowianus). Unfortunately, Bakhuizen Van Den Brink used only herbarium
specimens for the creation of his monograph and, as stated by Kostermans
(1958b), many well defined species were lumped together in this work. Many
specimens were also misidentified. Most of the major changes in classification
described in this work have now been overturned by several authors. His
grouping of the genus Boschia into Durio, however, still stands.
Corner (1939) offered a tentative scheme to untangle some of the

resultant confusion, although most of his suggestions were not taken up in
later works. Wyatt-Smith (1953a) unravelled much of the convoluted history
surrounding the wild species of Durio and set the stage for Kostermans’
monograph.
Kostermans' (1958b) book stands as the most recent and useful

monograph of the genus. This monograph is a combination of Kostermans
and Soegeng-Reksodihardjo’s 1958 work on Bornean species and Kostermans’
(1958a) work on non-Bornean species. This combined work describes and
depicts 27 species of Durio. In this work, the genus is divided into two
subgenera, Boschia and Eu-Durio. The subgenus Boschia, based on the
aforementioned difference in anther dehiscence, contains 6 species while Eu-
Durio contains the rest.
Post-Kostermans (1958b) treatment : Since the publication of his monograph,

Kostermans changed his opinion on the validity of D. cupreus Ridl., which
he had merged with D. carinatus Mast. [sensu Kostermans 1958b]. He re-
erected it to species status after examining new material (Kostermans 1961).
Furthermore, some of the specimens he included ‘reluctantly’ under D.
graveolens Becc. in his monograph (formerly considered as D. conicus Becc.
by Wyatt-Smith) were considered to most likely represent D. Wyatt-smithii
Kosterm. (a new species erected by Kostermans 1958b). This is further
complicated as Kostermans (1961) cites the specimen of Wyatt-Smith’s (Kep.
80131) as the one which he has reconsidered. No mention of this specimen
is made by Wyatt-Smith (1953a). Furthermore, the delineation of what
constitutes D. graveolens Becc. in Kostermans (1958b) from which he now
wishes to separate this species includes only D. dulcis (non Becc.) from
Wyatt-Smith (1953a) (listed by Wyatt-Smith as a synonym of D. graveolens
Becc.), D. conicus (non Becc.), and specifically not D. conicus Becc. sensu
Wyatt-Smith (1953a). Although, it appears that further clarification is necessary,
I am not convinced that further subjective analysis of herbarium material

can add anything more to our understanding.
Soegeng-Reksodihardjo (1965) described a new species, D. burmanicus,

which is allied to D. oxleyanus. This species was erected from an herbarium
specimen (lacking fruit) collected in the South of Burma (now Myanmar). To
date, no further information on this species has been published.
Kochummen et al. (1970) proposed that the species D. macrolepis

Kosterm. consisted of a mixture of two species, namely, D. pinangianus (Becc.)
Ridl. and D. macrophyllus (King) Ridl. Kostermans (1958b) separated D.
macrolepis from the other two species based on its production of flowers at
the base of the stem (cauliflory) rather than on the branches (ramiflory),
which is characteristic of the other two species. Kochummen et al. (1970) felt
that this difference was not sufficient enough to warrant the erection of a new
and separate species. Further, they noted that some of the specimens included
under D. macrolepis by Kostermans (1958b) consisted of flowers collected
from ramiflorous branches (not from cauliflorous inflorescences), or flowers
of which the position on the tree was not recorded. Kochummen et al. (1970)
suggested that most, but not all, specimens of D. macrolepis be reassigned to
D. pinangianus (Becc.) Ridl., the remaining known specimen being reassigned
to D. macrophyllus.
An incomplete specimen collected from Johore, Malaysia (FRI 8677)

has been described, but so far not named (Kochummen 1972). It is known as
Durio sp. ‘A’. It bears some morphological resemblence to D. lanceolatus and
D. kutejensis (Kochummen 1972).
Kostermans has recently published descriptions of two new species,

namely, D. bukitrayaensis Kosterm. (Kostermans 1990) which bears tiny fruits;
and D. macrantha Kosterm. which bears large edible fruits (Kostermans 1992a,b).
However, from the published description and pictures (Kostermans 1992a), it
is difficult to understand how D. macrantha differs from D. zibethinus.
Numerous species and varieties of durian have been named over the
years, over half of which are no longer recognized. Despite this, several of
these ‘defunct’ names continue to be used. This is hardly surprising considering
the complexity surrounding the taxonomy. Table 1 attempts to present all the
published names of species of Durio and their synonyms. The authority(ies)
of all published names along with the date of publication have been listed.
Where name changes have occurred, details are provided as far as possible.
The actual details of the current composition of several species (with respect
to particular herbarium specimens) is far more complex than can actually be
tabulated. The relevant works for further details should be consulted. The
genus Cullenia is also included as it has, in part, been previously included
within the genus Durio and may well be again. As diagrams have, and still
do have, a large role to play in the naming and identification of species, a list
of sources for figures of plant parts for currently recognized species is presented
in Table 2.
Table 1. Published species and synonyms of Durio
Boschia acutifolia Mast. (1874b)

-reassigned to D. acutifolius (Mast.) Kosterm. (sensu Kostermans 1953)
B. excelsa Korth. (1842)
-most specimens are now reassigned to D. excelsus (Korth.) Bakh.
(sensu Kostermans and Soegeng-Reksodihardjo 1958)
-Note: B. excelsa [sensu Merrill 1921, 1929] is now D. grandiflorus
(Mast.) Kosterm. and Soegeng. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
B. grandiflora Mast. (1874b)
-most specimens reassigned to D. excelsus (Korth.) Bakh. (sensu
Kostermans and Soegeng-Reksodihardjo 1958)
-some specimens reassigned to D. grandiflorus (Mast.) Kosterm. &
Soegeng. (sensu Kostermans and Soegeng-Reksodihardjo 1958)
B. griffithii Mast. (1874a)
-is now D. griffithii (Mast.) Bakh. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
B. mansoni Gamble [Anon. 1908]
-is now D. mansoni (Gamble) Bakh. (sensu Kostermans 1958a)
B. oblongifolia Ridl. (1933)
-reassigned to D. acutifolius (Mast). Kosterm. (sensu Kostermans and
Soegeng-Reksodihardjo 1958)
Cullenia ceylanica (Gardn.) K. Schum. (Kostermans 1956)
-includes part of C. excelsa Wight. (sensu Robyns 1970)
-includes D. ceylanicus Gardn. (sensu Robyns 1970)
C. exarillata A. Robyns (1970)
-includes part of C. excelsa Wight (sensu Robyns 1970)
C. excelsa Wight (1852)
(Contd....)
Table 1. Contd.
-is a mere synonym of D. ceylanicus Gardn. (sensu Kostermans 1958b)
-split among C. ceylanica, C. rosayroana and C. exarillata (sensu Robyns
1970)
C. rosayroana Kosterm. (1956)
-includes part of C. excelsa Wight. (sensu Kostermans 1956)
C. zeylanica (Gardn.) Wight ex K. Schum. (1895)
-transferred to D. ceylanicus Gardn. (sensu Bakhuizen Van Den Brink
1924a)
Durio acuminatissima [Kostermans and Soegeng-Reksodihardjo 1958]
=D. acuminatissimus Merr.
D. acuminatissimus Merr. (1924)
-included in D. zibethinus L. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. acutiminatissimus Merr. [Lim 1990]
=D. acuminatissimus Merr.
D. acutifolia (Mast.) Kosterm. (1953b)
-now included in D. acutifolius (Mast.) Kosterm. (sensu Kostermans
and Soegeng-Reksodihardjo 1958)
D. acutifolius (Mast.) Wyatt-Smith (1953a)
-includes B. acutifolia (Mast.) (sensu Wyatt-Smith 1953a)
-includes D. griffithii (Mast.) Bakh. var. acutifolius (Mast.) Bakh. (sensu
Wyatt-Smith 1953a)
D. acutifolius (Mast.) Kosterm. & Soegeng. (1958)
-includes D. acutifolius (Mast.) Wyatt-Smith
-includes B. oblongifolia Ridl.
-includes D. griffithii (Mast.) Bakh. var. acutifolius (Mast.) Bakh.
D. affinis Becc. (1889)
-includes D. malaccensis Planch. ex Mast. (sensu Kostermans and
D. beccarianus Kosterm. & Soegeng. (1958)
D. bukitrayaensis Kosterm. (1990)
D. burmanicus Soegeng. (1965)
D. carinatus Mast. (1874b)
-includes D. cupreus Ridl. (sensu Kostermans and Soegeng-Reksodihardjo
1958)
(Contd....)
Table 1. Contd.
D. carinatus Mast. var. bintulensis Becc. (1889)
D. ceylanica Gardn. [Wight 1852]
= D. ceylanicus Gardn.
D. ceylanicus Gardn. (1847)
-includes D. zibethinus Moon (sensu Gardner 1847)
-is now C. ceylanica (Gardn.) K. Schum. (sensu Robyns 1970)
D. conicus Becc. (1889)
-is now D. dulcis Becc. (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. crassipes Kosterm. & Soegeng. (1958)
D. cupreus Ridl. (1938)
-included in D. carinatus Mast. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
-re-erected to species status (Kostermans 1961)
D. dulcis Becc. (1886)
-reduced to D. conicus Becc. (sensu Bakhuizen Van Den Brink 1924a)
-synonym of D. graveolens (sensu Wyatt-Smith 1953a)
-includes D. oblongus Mast. (sensu Kostermans & Soegeng-Reksodihardjo
1958)
-includes D. conicus Becc. (sensu Kostermans & Soegeng-Reksodihardjo
1958)
D. excelsus (Korth.) Bakh. (1924a)
-includes part of D. excelsus (Korth.) Bakh. var. typicus Bakh. (sensu
-includes D. griffithii (Mast.) Bakh. var. heteropyxis (Griff.) Bakh.
-includes most of B. excelsa Korth. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. excelsus (Korth.) Bakh. var. typicus Bakh. (1924b)
-part is now D. excelsus (Korth.) Bakh.; other part is D. griffithii
(Mast.) Bakh. (sensu Kostermans and Soegeng-Reksodihardjo 1958)
D. excelsus (Korth.) Bakh. var. grandiflorus (Becc.) Bakh. (1924b)
-transferred to D. grandiflorus (Mast.) Kosterm. & Soegeng. (sensu
(Contd....)
Table 1. Contd.
D. falcata [Stadelman 1966]
-name of no taxonomic standing
D. foetida Thunb. (1796)
-name of no taxonomic standing, synonym of D. zibethinus L.
D. grandiflorus (Mast.) Kosterm. & Soegeng. (1958)
-includes part of B. grandiflora Mast. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
-includes B. excelsa Korth. [sensu Merrill 1921, 1929] (sensu Kostermans
and Soegeng-Reksodihardjo 1958)
-includes D. excelsus (Korth.) Bakh. var. grandiflorus (Becc.) Bakh.
D. gratissimus Becc. (1889)
-is now included in D. oxleyanus Griff. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. graveolens Becc. (1889)
-reduced to D. conicus Becc. (sensu Bakhuizen Van Den Brink 1924a)
-includes D. dulcis (sensu Wyatt-Smith 1953a) [this inclusion not valid
under Kostermans and Soegeng-Reksodihardjo 1958]
-re-erected to D. graveolens Becc. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. griffithii (Mast.) Bakh. (1924a)
-includes some of D. griffithii (Mast.) Bakh. var. heteropyxis (Griff.)
Bakh. (sensu Kostermans and Soegeng-Reksodihardjo 1958)
-includes some of D. excelsus (Korth.) Bakh. var. typicus Bakh. (sensu
D. griffithii (Mast.) Bakh. var. heteropyxis (Griff.) Bakh. (1924b)
-a synonym of D. excelsus (Korth.) Bakh. with the exception of a
Sumatra specimen (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. griffithii (Mast.) Bakh. var. acutifolius (Mast.) Bakh. (1924b)
-included under D. acutifolius (Mast.) Kosterm. & Soegeng. (sensu
D. kinabaluensis Kosterm. & Soegeng. (1958)
-includes D. kutejensis (Hassk.) Becc. forma kinabaluensis Bakh.
(Contd....)
Table 1. Contd.
D. kutejensis (Hassk.) Becc. (1889)
-includes L. kutejensis Hassk. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. kutejensis (Hassk.) Becc. forma kinabaluensis Bakh. [Wyatt-Smith 1953a]
-is now D. kinabaluensis (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. lanceolatus Mast. (1874b)
-reduced to D. singaporensis Ridl. (sensu Wyatt-Smith 1953a)
-re-erected to species status (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. lissocarpus Mast. (1874b)
-reduced to D. carinatus Mast. (sensu Bakhuizen Van Den Brink 1924a;
Wyatt-Smith 1953a)
-re-erected to species status (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. lowianus Scort. ex King (1891)
-includes D. wrayii King. (sensu Kostermans 1958a)
-includes D. zibethinus L. var. roseiflorus Corner (sensu Kostermans
1958a)
D. lowii Hook. [Sutisna and Soeyatman 1985]
-name of no taxonomic standing, possibly D. lowianus Scort. ex
King
D. macrantha Kosterm. (1992a)
D. macrolepis Kosterm. (1958a)
-specimens fractured into D. pinangianus (Becc.) Ridl. and D.
macrophyllus (King) Ridl. (sensu Kochummen et al. 1970)
D. macrophyllus (King) Ridl. (1922)
-reduced to D. oblongus Mast. (sensu Bakhuizen Van Den Brink 1924a)
-re-erected and includes D testudinarum Becc. var. macrophylla King
(sensu Kostermans 1958a)
D. malaccensis Planch. ex Mast. (1874a)
-includes D. sumatranus Becc. (sensu Kostermans 1958a)
-part has been removed to D. affinis Becc. (sensu Kostermans and
D. mansoni (Gamble) Bakh. (1924b)
-includes B. mansoni Gamble (sensu Kostermans 1958a)
(Contd....)
Table 1. Contd.
D. oblongus Mast. (1874b)
D. oxleyanus Griff. (1845)
-includes D. gratissimus Becc. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
-includes D. griffithii Planch. ex King. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. perakensis King (1891)
-a name of no taxonomic standing, possibly D. lowianus Scort. ex
King
D. penangianus [Kochummen and Wyatt-Smith 1979]
=D. pinangianus (Becc.) Ridl.
D. pinangianus (Becc.) Ridl. (1922)
-includes D. testudinarum Becc. var. pinangianus Becc. (sensu Kostermans
1958a)
-includes some of D. macrolepis Kosterm. (sensu Kochummen et al.
1970)
D. purpureus Kosterm. & Soegeng. (1958)
D. singaporansis Ridl. [Lim 1990]
=D. singaporenis Ridl.
D. singaporensis Ridl. (1922)
-reduced to D. oblongus Mast. (sensu Bakhuizen Van Den Brink 1924a)
-re-erected by Wyatt-Smith (1953a) as a possible synonym of D.
sumatranus
-retained as D. singaporensis Ridl. (sensu Kostermans 1958a)
D. singapurensis [Corner 1978]
=D. singaporensis Ridl.
D. spontaneus Bakh. [Van Steenis 1949]
-is now D. lowianus Scort. ex King. (sensu Kostermans 1958a)
D. stercoraceus Noronha (1790)
-synonym of D. zibethinus L.
D. sumatranus Becc. (1889)
-may include D. singaporensis Ridl. (sensu Wyatt-Smith 1953a)
-synonym of D. malaccensis Planch. ex Mast. (sensu Kostermans
1958a)
(Contd....)
Table 1. Contd.
D. testudinarium Becc. [Lim 1990]
=D. testudinarum Becc.
D. testudinarum Becc. (1889)
D. testudinarum Becc. var. pinangianus Becc. (1889)
-now D. pinangianus (Becc.) Ridl. (sensu Wyatt-Smith 1953a and
Kostermans 1958a)
D. testudinarum Becc. var. macrophylla King (1891)
-lifted to D. macrophyllus (King) Ridl. (sensu Wyatt-Smith 1953a and
Kostermans 1958a)
D. testudinarum Becc. var. macrophyllus King [Corner 1939]
=D. testudinarum Becc. var. macrophylla King
D. wrayi [Ridley, 1922; Kostermans and Soegeng-Reksodihardjo 1958]
=D. wrayii King
D. wrayii King (1891)
-reduced to D. lowianus Scort. ex King (sensu Kostermans 1958a)
D. wyatt-smithii Kosterm. (1958a)
D. zeylanica [Worthington 1959]
=D. ceylanicus Gardn.
D. zibethianus Murr. [Kanehira 1935]
=D. zibethinus L.
D. zibethinus L. (1774)
D. zibethinus L. var. roseiflorus Corner (1939)
-now D. lowianus Scort. ex King (sensu Kostermans 1958a)
D. zibethinus Moon (1824)
-synonym of D. ceylanicus Gardn.
D. zibethinus Murr. (1774) improper attribution of authority = D. zibethinus
L. (1774)
D. sp. “A” [Kochummen 1972]
-a new, as yet un-named and incompletely known species with similarities
to D. lanceolatus and D. kutejensis
Lahia kutejensis Hassk. (1844)
-is now Durio kutejensis (Hassk.) Becc. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
(Contd....)
Table 1. Contd.
Neesia griffithii Planch. ex King
-is now D. oxleyanus Griff. (sensu Kostermans and Soegeng-Reksodihardjo
1958)
Currently valid species are in bold; species of dubious, controversial or unknown status are in
italics, invalid species are in normal type. Dates in round brackets ( ) after a species name
represent the date of publication of the species by the preceding authority. Authors and dates
in square brackets [ ] represent publications in which an invalid or unrecognized name was cited
either by typographical error or erroneously, or cases where the author of the first published
description is not the authority.
Table 2. Sources for figures of currently valid species of Durio and close
allies
Species Fruit Flower Seed Tree Leaf
Cullenia ceylanica LN5LN13 LN5BW13 LN13BW17 BW10 (BW,LN)13

BW15LN33* BW35
Cullenia exarillata LN25LN33* LN25LN33* LN25 LN25LN33*
Cullenia rosayroana LN33* LN33* LN13LN33*
Durio acutifolius LN6 LN4c LN4c BW15 LN6

LN C N15,16
15,16 26
LN15,16
Durio affinis LN6LN15,16 LN4LN15,16 LN4LN6

LN31* LN15,16
Durio beccarianus LN15,16 LN15,16
Durio bukitrayaensis LN18 LN18 LN18
Durio burmanicus LN28 LN28
Durio carinatus LN4d LN4d LN4d LN4d

LN15,16 LN15,16 LN15,16
Durio crassipes LN15,16 LN15,16 LN15,16
Durio dulcis LN4,4b LN15,16 LN4 LN4,4b

15,16 26
LN C LN15,16
(Contd....)
Table 2. Contd.
Durio excelsus LN12* LN2*LN12* LN7* LN12*

LN C LN15,16
15,16 26
LN15,16
Durio grandiflorus LN6BW6 LN6LN15,16 LN6BW6

LN15,16BW22 LN15,16
BW22
Durio graveolens C 1LN4LN6 LN15,16 LN4LN6

LN15,16 LN15,16
Durio griffithii BW14 LN15,16 BW14 LN15,16
Durio kinabaluensis LN15,16 LN15,16 LN15,16
Durio kutejensis LN15,16 LN4LN15,16 LN4LN23

LN23C 26 LN23BW27
Durio lanceolatus LN4LN15,16 LN4LN15,16 LN4LN15,16

C26
Durio lissocarpus LN15,16 LN15,16
Durio lowianus LN16 LN16
Durio macrantha BW19 BW19,20 BW19 BW19 BW19
Durio macrolepis LN16 LN16
Durio macrophyllus LN16 LN16 LN16
Durio malaccensis LN4aLN16 LN4BW8 BW8 LN4,4aBW8

LN24* LN16LN24* LN16LN24*
Durio mansoni LN16,29 LN16,29 LN16,29
Durio oblongus LN4LN15,16 LN4LN15,16 LN4 LN4LN15,16
Durio oxleyanus LN15,16C 26 LN15,16 LN7* LN3*LN6

LN15,16
Durio pinangianus LN16 LN16 LN16

(Contd....)
Table 2. Contd.
Durio purpureus LN15,16 LN15,16
Durio singaporensis LN16 LN16 LN16
Durio testudinarum LN4LN6 LN4LN15,16 LN4LN15,16 LN4LN6

LN15,16C 26* LN15,16
LN31
Durio wyatt-smithii BW16 BW16 BW16
Durio zibethinus LN11 LN2LN4 LN4 LN11 LN4LN6

LN15,16 LN6 BW30 LN11
LN21BW30 LN9LN11 LN15,16
LN31 LN15,16 LN21
LN21LN34
Kostermansia BW32
malayana
[Published diagrams of durian species: BW=black and white photo, C=colour
photo, LN=line drawing. Citations with 2 superscripts indicate the same figure
appears in two separate publications. An * indicates that additional clarifying
information is given in the following list of sources.]
Sources for data presented in Table 2:

1
Anon. (1986b).
2
Baillon (1875) *Note: Figure 174 and 175 of this work are labelled Boschia excelsa = D.
excelsus (Korth.) Bakh. sensu Kostermans (1958b).
3
Bakhuizen Van Den Brink (1924a) *Note: According to Kostermans (1958b), Figure D (a
flower) of Table 38 labelled D. oxleyanus is incorrect. The rest of the figures (including
leaves) in the table are accurate.
4
Beccari (1889) Changes to the classification system of Beccari by Kostermans (1958a) reflected
in the above table. (a) Species described as D. sumatranus Becc. are now considered to
be another specimen of D. malaccensis Planch. ex Mast.; (b) D. conicus Becc. is now
considered to be another specimen of D. dulcis Becc.; (c) Boschia acutifolia is now
considered as D. acutifolius (Mast.) Kosterm.; (d) D. carinatus Mast. is split into D.
carinatus Mast. and D. lissocarpus Mast. under this scheme Table: 17 Figs. 6-7 of D.
carinatus are called D. lissocarpus Mast., and Table: 17 Figs. 1-5,8-9, 18 Figs. 1-5
remain D. carinatus. Mast.
5
Beddome (1869).
6
Cockburn (1976).
7
De Vogel (1980) *Note: Diagrams are of seedlings with attached seeds.
8
Foxworthy (1927).
9
Griffith (1854a) Plate 596
10
Kadambi (1954).
11
Köing (1804).
12
Korthals (1839-1842) *Note: What is depicted as Boschia excelsa is now interpreted by
Kostermans (1958b) as D. excelsus (Korth.) Bakh.
13
Kostermans (1956), 14Kostermans (1953b), 15Kostermans (1958a), 16Kostermans (1958b).
17
Kostermans (1958c), 18Kostermans (1990), 19Kostermans (1992a), 20Kostermans (1992b).
21
Lamarck (1823).
22
Meijer (1969).
23
Ochse (1927).
24
Ridley (1922) *Note: Part of the figure representing D. malaccensis was copied from Masters
(1874b). The copied parts include the anther at the top right, and the two bottom left
diagrams (longitudinal section of an ovary, and a spine bearing a peltate scale). According
to Kostermans (1958b), of these 3 copied diagrams, only the anther belongs to D.
malaccensis Planch. ex Mast.
25
Robyns (1970).
26
Setiadi (1991) *Note: The photo of D. testudinarum is captioned ‘Durian kura-kura’.
27
Soegeng-Reksodihardjo (1962), 28Soegeng-Reksodihardjo (1965).
29
Sprague (1915).
30
Stanton (1966).
31
Wettstein (1935) *Note: Figure 3 on page 806, showing the flower of D. affinis, is copied
from volume III, Table 24 of Beccari’s Malesia of 1889. This diagram was also copied
by Kostermans 1958b, appearing as part of Fig. 28.
32
Whitmore (1990).
33
Wight (1852) *Note: This figure is titled Cullenia excelsa Wight. which, according to Kostermans
(1958b), is equivalent to Durio excelsa Gardn.=Cullenia ceylanica (Gardn.) K.Schum.
However, Kostermans (1956) and Raizada (1957) consider Wight’s figures to be drawn
from a mixture of specimens of C. ceylanica and C. rosayroana, most of them being
C. rosayroana with the exception of the fruit depicted in figures 14-16 which are
probably C. ceylanica. More recently, Robyns (1970) has described a new species
Cullenia exarillata A. Robyns. which he claims is what is depicted in Wight’s original
figures, the interpretation of Kostermans (1956) and Raizada (1957) being in this regard
erroneous.
34
Winkler (1905).
35
Worthington (1959).
Keys to the species : As numerous new species have been described in the
last 50 years, and major reworking of species have also occurred, keys prior
to Kostermans (1958b) are now of little use. An exception to this might be
Wyatt-Smith (1953a), whose key to 19 species, based mainly on floral
characteristics, is largely in agreement with the classification of Kostermans
(1958b). Kostermans (1958b) presents a key in his monograph based largely
on floral characteristics. Soegeng-Reksodihardjo (1962) provided keys to six
edible species based on floral, fruit or vegetative characteristics.
Kochummen (1972) presents two keys to 10 species of Durio, one

based on leaf characteristics, and the second on flower and fruit characteristics.
Cockburn (1976) has published a key to 12 species based on leaf and fruit
characteristics.
Future taxonomic work : Despite all the taxonomic work which exists on
the genus Durio, it is quite clear that much remains to be clarified. Some
of the wild species are known only from very little and incomplete herbarium
specimens. For example, D. crassipes Kosterm. & Soegeng. is only known
from one herbarium specimen, consisting of a few flowers which are missing
the epicalyx (Kostermans 1958b). Some species have not been collected in
many years and may have already been extinct. An examination of the various
treatments of the genus, which have been published, reveals that the major
cause of the collapse of old species and the erection of new ones is the
subjective decision as to how physically different two herbarium specimens
have to be in order to be given different names. Further, monographing of
species in the style so far established for them seems unlikely to provide
more insight on the matter. It is difficult to imagine how such further
taxonomic shuffling can really address the most pressing and meaningful
questions that are in need of answers. Future work involving proper cladistical
analysis of as many characters as can be obtained from the relevant herbarium
(and other) specimens, perhaps coupled with RFLP (restriction fragment length
polymorphism) mapping and isozyme analysis would be valuable. Isozymes
have already proved useful in distinguishing clones of D. zibethinus (Salma
1993). It has also been suggested that the position and morphology of the
leaf trichomes may also be of taxonomic value (Salma 1991). Furthermore,
chromosome counts might be of use in addressing taxonomic questions
surrounding durian. Many of the species of Durio are known from living
specimens, thus crossing experiments are possible. The structure of many
durian flowers lends itself to artificial pollination, and the life span of durian
pollen has been demonstrated to be sufficiently long under appropriate
conditions to allow such crossing. Some crossing experiments have been
conducted and offer the exciting prospect of improved trees, especially with
regard to disease resistance.
Morphology
A fair amount of morphological and anatomical information on durians
has been published. The majority of this information deals with the structure
of the flowers and fruits, but some information is available on the leaves,
roots, wood anatomy and seeds. One of the most fascinating developments
from the study of the morphology of durian fruits has been E.J.H. Corner’s
durian theory (Corner 1949). A full examination of this theory is well beyond
the scope of this work, as much of it involves species extraneous to the topic,
but as this theory is forever linked to the durian fruit, it will be briefly
discussed. Additionally, the tree architecture of durian has been analyzed. The
anatomy of the wood will be discussed in a later chapter in relation to the use
of durian in forestry, as this is the field in which such information is most
useful. As chromosome counts have often been used in conjunction with
taxonomic and morphological analysis, this information is also included under
this section.
Durian theory : In 1949, Corner elaborated his ‘Durian Theory’ which, among
other things, predicted that the ancestral angiospermous fruit was large, spiny
and dehiscent, bearing large seeds covered in colourful fleshy arils. This type
of fruit is more or less typified by that of the durian tree (Durio zibethinus L.).
This theory was spurred by his observations of species with strikingly similar
fruit morphologies in numerous unrelated angiosperm families. These species
are usually very rare, and their fruits atypical of the other more common
members of the family. Through a series of arguments, he proposed it was
unlikely that this peculiar fruit type had evolved independently numerous
times, and more likely that it represents a relic. Several arguments have been
raised against this theory, most notably by Parkin (1953) and Van Der Pijl
(1952, 1955). The main points of contention are the subjectivity of Corner’s
observations, and the denial that rarity necessarily represents antiquity. An
attempt to refute this theory was published by Datta and Biswas (1969), but
their ‘argument’ was based on an obvious misunderstanding of Corner’s theory,
and thus provides neither support nor evidence against it. Corner has since
expanded upon his theory (Corner 1953, 1954a, 1954b), and it has been given
some support by others (Mabberley, 1974a,b; Von Teichman and Van Wyk
1991, 1994). The most objective and perhaps most useful new evidence comes
from Von Teichman and Van Wyk (1991) whose use of statistical character
associations revealed that ‘durian fruit and seed-like’ characteristics (i.e.,
recalcitrance, arils, pachychalazy, etc.) were significantly correlated with the
occurrence of other character traits generally regarded as primitive. More
recently, Von Teichman and Van Wyk (1994) have elaborated on the idea
hinted at by Corner (1949) that recalcitrance (high seed moisture content and
a short period of seed viability) is the ancestral state of seeds, orthodoxy (the
ability to withstand desiccation) having evolved later under selection pressure.
This avenue of investigation suggests that recalcitrant seeds may share
morphological and physiological commonalities by descent. This concept may
be of substantial importance in the understanding of recalcitrant seeds and
definitely opens new lines of enquiry. However, to this date, the validity of
Corner’s durian theory remains in dispute, and its domain of applicability is
yet to be firmly established.
Floral morphology : According to Hawkins (1986), the durian is one of the

most beautiful flowering trees. The flowers do have a certain appeal (apart
from their smell of sour milk). The common durian is ramiflorous (Fig. 1),
and very rarely cauliflorous (Lim 1990), the flowers being borne on older
branches in di- or tri-chasial cymes (Davis and Bhattacharya 1974) consisting
of 10-25 (Croft 1981), 5-30 (Davis and Bhattacharya 1974) flowers. Species
of durian with small fruits (D. griffithii) have flowers that are borne in the leaf
axils (Corner 1988). In the developing flower bud, the sepaline, petaline,
staminal and carpellary primordia develop acropetally and at approximately
the same rate (Soepadmo and Eow 1977). The floral bud of D. zibethinus is
completely enclosed within a bracteole which eventually splits into 2 or 3
sections near maturity.
The flowers are generally pentamerous with a fused calyx usually
consisting of five lobes (Fig. 2). Some variation does occur, flowers of clone
D88 occasionally have 4 petals, while those of clones D8 and D104 occasionally
have 6 (personal observations). Flowers with more or less than 5 petals do not
always have corresponding changes in the number of other floral parts. For
example, flowers of clone D88 occasionally have 4 petals, but always have 5
staminal phalanges (personal observations). In some species of Durio, the
sepals separate at anthesis (Kostermans 1958b), however, this is not the case
in D. zibethinus.
Flowers normally possess five staminal groups (bearing varying numbers
of filaments). To quote Van Heel (1966), ‘the stamens are generally placed in
concave more or less antepetalous phalanges’, ‘staubblattbnndel’ of Winkler
(1905). The filaments near the middle of the staminal phalange are longer
than those on the edge (Fig. 2d). The filaments vary with respect to the degree
with which they are fused with neighbouring filaments. Some filaments are
connate just at the base, others are fused along almost their entire length to
adjacent filaments, the number of filaments per stamen is positively correlated
with floral size (personal observations). The filaments of D. acutifolius are
apparently exceptional within the genus Durio in not being united into staminal
phalanges. In some other species, the phalanges themselves are united more
or less into a tube. The anthers are unilocular, midfixed, strongly recurved
and twisted, and dehisce via a single slit (Croft 1981). According to Davis and
Bhattacharya (1974), the anthers are two-celled or occasionally one-celled.
An extremely detailed examination of the stamens of several species of Durio
is presented by Van Heel (1966), and the development of the anther-wall has
been described in detail by Soepadmo and Eow (1977).
The single ovary of D. zibethinus is superior, and normally 5-loculed.

It is not stipitate as claimed in the original Latin description (Linnaeus 1774).
The ovules are anatropous (Winton and Winton 1935), bitegmic and
crassinucellate. The micropyle is formed by both the inner and outer integuments
(Soepadmo and Eow 1977). Embryo sac development is of the Polygonum
type, and the antipodal cells are ephemeral (Soepadmo and Eow 1977). The
stigmatic surface is heavily papillate (Soepadmo and Eow 1977). There are
differences in colour and shape of durian stigmas (Ochse 1961; Lye 1980).
Some clones (D8, D24, D104) have 5-lobed stigmas, whereas the other clones
show no trace of lobes (personal observations). Some stigmas are top shaped
(D2, D88, D96) while others are broad and flat (D16, D66). Stigma shape is
highly consistent within a clone. Clones D8, D66 and D104 have bright orange
stigmas, whereas several clones have yellow stigmas (personal observations).
Figure 1. The common durian is ramiflorous. The flowers/fruits are borne

along big branches that are capable of bearing the weight of the
mature fruit.
Figure 2. An open flower from D24 durian and its parts. The flower was
collected in the evening of anthesis, the male and sterile floral
organs abscind during the night. Figures B-E represent parts collected
the next morning.
A) A mature flower, projecting from the epicalyx which normally
splits into 2 or 3 sections. The sepals are fused to form a short tube
with a swollen base. The 5 petals emerge through the calyx tube
and reflex, exposing the numerous filaments and the stigma.
B) The gynoecium of the flower. All of the floral parts except the
gynoecium have abscinded. The ovary is covered in peltate scales,
the long style is often kinked in flowers from this particular clone.
The stigma is also visible. The receptacle bears visible scars left
from the abscision of the other floral parts.
C) The calyx of the flower. The fused sepals form a swollen base
which holds the nectar produced by nectaries inside the calyx.
D) A staminal phalange consisting of 7 filaments. The filaments
are connate at the base. Note that the filaments near the centre of
the phalange are longer than those at the sides.
E) A petal collected from under the tree the morning after anthesis.
Note that the petals are reflexed to expose the numerous filaments
enclosed within them.
According to Chin and Phoon (1982), there is heterostyly between

the different durian clones, however, based on evidence which is discussed in
a later section of this work, this is unlikely the case. Davis and Bhattacharya
(1974) note that some buds have the anthers packed in a right-handed twist
while others have a left-handed twist. Likewise, the aestivation of the petals
is either right- or left-handed2.
The average nectar volume of a durian flower is 0.36 ml (Gould

1977, 1978), the caloric value of durian nectar has been calculated at 869
calories per ml (Gould 1978).
Pollen morphology : The pollen grains of durian were probably first described
by Van Der Pijl (1936) who described them as 90 mm in diameter (wet), and
covered in a sticky mass which stains red with Sudan III. The pollen grains
are approximately spherical (Soepadmo and Eow 1977). Davis and Bhattacharya
(1974) measured pollen grain size from left and right twisted flowers, and
discovered that left twisted flowers had slightly, though consistently, larger
pollen grains from those of right twisted flowers (see also footnote 1). According
to Soepadmo and Eow (1977), the pollen grains of Durio zibethinus are
approximately 80-150 µm in diameter, 3-4 or rarely 6 porate with a smooth
and sticky exine. Davis and Bhattacharya (1974) calculated durian pollen
grains to vary from 20 to 80 µm with a mean of approximately 55 µm for dry
pollen and 67 µm for water soaked pollen. Erdtman (1972) states that the
equatorial diameter is 50-100 µm. A study by Salakpetch et al. (1992) included
the measurements of the equatorial and polar axes of pollen grains from four
Thai cultivars. This study showed that there were slight differences between
cultivars, however, the average length of both axes of all cultivars was of the
order of 15 µm. I have no explanation for the much smaller dimensions
provided by this study compared with those mentioned previously.
Durian pollen is binucleate when shed, and remains viable for

approximately 48 hours after shedding if kept at room temperature (Soepadmo
and Eow 1977). Erdtman (1972) briefly described the pollen morphology of
eight species of Durio as 3(-4) colporate, colpoidorate, and usually suboblate
with a thin sextine. According to Fuchs (1967), D. oblongus and D. testudinarum
stand out among other species of Durio in that their pollen grains are not
smooth but bluntly macropositively sculptured. The pollen grains of Cullenia
2
Davis studied thirteen species of Bombacaceous trees (excluding the genus Durio), and found
that all bore both left and right handed flowers (Davis 1967), stamen number and pollen size
in levo- and dextro-rotary flowers of Bombacaceae. Review of Palaeobotany and Palynology,
3, 133-139. Water soaked pollen grains of Bombax ceiba from left handed flowers had significantly
bigger pollen grains, while no significant difference in size was found in the other 12 species.
are very similar to those of Durio (Fuchs 1967).
The pollen of D. griffithii is shed in thread-like chains (Ha et al.

1988). According to Soepadmo and Eow (1977), the pollen of D. zibethinus
is shed singly or in clumps. The pollen of several Thai cultivars of D. zibethinus
have been reported to be shed in clumps (Salakpetch et al. 1992).
Fruit morphology : There is some degree of exaggeration in the available

literature as to the maximum size attained by durian fruits. It has been stated
that the fruits can reach 30 cm in length, 15 cm in diameter and attain weight
of 20 kg (Anon. 1975). The stated physical dimensions are more than reasonable;
however, the weight is a gross exaggeration; a durian of the given dimensions
and mass would have to have a density greater than that of titanium to weigh
20 kg. This possibility is negated as the specific gravity of durian fruits is
known to be less than 1 (Chattavongsin and Siriphanich 1990a,b), i.e., durians
float. Perhaps the claim by Everett (1968) that the fruits can weigh up to 100
pounds should remain unmentioned.
Large varieties of durian do exist, and are known as durian kepala

gajah3, and may weigh 4.5-7 kg (Ridley 1902; Chevalier 1934; Grist 1936);
the size of an average durian fruit is much less. Pratt and Del Rosario (1913),
for example, records the average weight of durians in their study as 2.25 kg.
Coronel (1986) cites Mon Thong durians as weighing 3.2 kg. Mohamad Idris
(1987) lists the average weights of fruits of several registered Malaysian
durian clones, none of which exceeded 3 kg. Lim et al. (1992) recorded a fruit
of approximately 4 kg as a result of crosses between some Thai cultivars.
Winton and Winton (1935) described 5 distinct pericarp layers as

seen in cross-section: (1) an epicarp of thin walled rounded-polygonal cells,
and scales; (2) a hypoderm of one or more layers of thin walled round cells;
(3) fibre zone of white porous fibres; (4) mesocarp of thin walled parenchyma
containing numerous oil drops; and (5) endocarp of thin walled elongated
narrow celled parenchyma with occasionally sclerenchymatized walls.
Different cultivars can be distinguished to some extent by variations

in fruit morphology. For example, the shape of the spines has been described
for several Thai varieties (Hiranpradit et al. 1992a). The overall gross morphology
of the fruit is useful in grading them for market (Hiranpradit et al. 1992).
Hiranpradit et al. (1987) claim that fruit shape and spine morphology are
3
‘Kepala’ is Malay for head, while ‘gajah’ is the Malay term for the Indian Elephant ( Elephas
maximus).
useful for characterizing durians into distinct groups, and both are highly
heritable characters.
Durian fruits develop over a period of 3 to 4 months. Despite this

long period of development, most of the growth occurs late in development
(Fig. 3). The ovary of a durian flower is covered with peltate scales, however,
spines begin to appear early in development (Fig. 3). Spineless durians are
occasionally encountered. These are produced artificially by scraping the scales
off the immature fruits (Mohamad Idris 1987); the bases of the scales normally
develop into the spines as the fruits mature. Rodrigo (1968) claims that a
naturally spineless variety of durian growing wild in Davao, Phillipines was
discovered, and that fruits borne on trees grown from seeds of this fruit at the
Bureau of Plant Industry Experiment Station are also spineless. I have found
no other mention of these or other naturally spineless durians in the literature
other than that of Coronel et al. (1983) who briefly mention a spineless
variety from Davao.
The shape of durian fruits is affected by the presence of seeds. Locules

containing unfertilized ovules tend not to develop, and thus the fruits become
uneven in shape. Fruit shape greatly affects marketability and thus an
understanding of pollination and its effect on durian fruit development is
important for the improvement of durian.
The anatomy of the mature fruit stalk has been described by

Chattavongsin and Siriphanich (1987). It consists of an outer periderm surrounding
a cortex consisting of parenchyma, tannin containing cells and scleroids. The
vascular tissue is contained in the centre of the stalk. Chattavongsin and
Siriphanich (1987) showed that the number of cortical sclerids in the stalks
of Mon Thong durians increased as the fruits matured. This study provided a
useful and non-destructive method of estimating fruit maturity by measuring
the stiffness of the stalk with an ‘Effigi’ firmness tester (Chattavongsin and
Siriphanich 1990a,b). The mature fruit is a loculicidally dehiscent capsule
(Ketsa and Pangkool 1994).
Fruit teratology : The first published description of durian teratology was in

the ‘Herbarium Amboinense’ of Rumphius, in which the rare occurrence of
a small durian fruit within a locule of a durian fruit was described. This
occurrence was further elaborated by Joger (1814). Penzig (1921) briefly
described such a fruit, and Ochse (1925-1926) provided a photograph of a
specimen with a teratology similar to that described by Rumphius, which he
termed ‘doerian si bakoel’. This specimen was further described by Costerus
and Smith (1932). Venema (1937) provided a description and a figure of a
new specimen with this teratology collected in 1933. He described it as ‘perhaps
a case of median prolification with a tendency to apocarpy.’
Ridley (1902) also gave an account of a variety of durian in which

an entire fruit actually developed within the ovary of a durian, ‘I have met
with a curious variety in which the fruit which was very large had a hole in
the top and inside was another small durian complete with the spiny husk
replacing the placenta of the fruit.’ Ridley (1922) described ‘a very small
fruited variety with only one carpel and one seed’.
Rao and Singh (1964) reported that occasionally the floral meristem
continued to be active after formation of all the regular floral parts resulting
in the production of stamens and carpels within the ovary on the central axis
of the fruit. The anthers were found to contain pollen, but none of the carpels
contained mature ovules. No petals or sepals were seen to be produced. The
authors noted that these anomalous structures do not take the place of previous
structures, but are superfluous in nature.
A brief description of the flowers, and some scanning electron

micrographs of some floral parts of Durio zibethinus is given by Chin and
Phoon (1982), although no evidence of stamens or carpels was found within
the ovaries of their specimens.
‘A fruit of which the margins of the carpels with their warts and
hunches at the inner-side interlock like a cog-wheel’ was described by Costerus
and Smith (1932).
Other teratologies : Apart from fruits occurring within fruits, petaloid stamens
have also been described in durian (Winkler 1905). Soepadmo and Eow (1977)
related that they had twice observed the formation of binucellate ovules. These
ovules both shared a common outer integument, but each had its own inner
integument.
Ovule and seed morphology and development : Inside each developing

locule of a durian ovary, two alternate rows of 5-7 ovular primordia develop
on the central placental column (Soepadmo and Eow 1977). The development
of these primordia into ovules is described and depicted in detail by Soepadmo
and Eow (1977). They also described megasporogenesis and embryosac
development in detail. The initial stages of embryogenesis of D. griffithii have
been described by Ha et al. (1988).
The ovules of durian are bitegmic. Although the ovule primordia of

both integuments develop simultaneously, the outer integument grows faster,
and overtops the inner integument and micropyle is formed by both integuments
(Soepadmo and Eow 1977).
The aril develops from the funiculus/funicular end of the ovule (Soegeng-
Reksodihardjo 1962; Soepadmo and Eow 1977). In D. zibethinus and the
majority of wild species, the aril eventually completely surrounds the seed; D.
griffithii, D. oblongus and D. malaccensis have incomplete arils, and D.
singaporensis has no aril (Kostermans 1958b).
The inner integument is eventually crushed by the developing embryo

and does not persist, the cells of the outer integument develop considerably
(Soepadmo and Eow 1977). The endosperm of the seed does not persist until
maturity.
The major storage reserves of durian seeds are starch and protein. No
lipid bodies are present in the storage parenchyma cells of mature durian
seeds (Brown 1995b).
The storage protein present in mature durian seeds is unrelated to the

7S and 11S globulins common to many seeds (Charbonneau et al. 1991;
Brown 1995b). This storage protein exists in two ungenically related forms of
slightly differing electrophoretic mobility; these proteins have been termed
‘zibethinins’ (Brown 1995b).
The development of cotyledon storage parenchyma cells and the

deposition of storage reserves were studied by Brown (1995b). It was shown
that some of the storage protein and starch was mobilized prior to fruit dehiscence.
Furthermore, the deposition of storage proteins within the vacuoles of the
storage parenchyma cells during development does not seem to involve the
golgi apparatus as is typical of many dicotyledonous seeds, rather the protein
appears to be deposited in large swellings of the endoplasmic reticulum which
presumably fuse with the vacuoles (Brown 1995b).
Some measurements of gibberellins in the seeds of developing durians

have been published (Mamat and Wahab 1990, 1992).
Leaves : Durian leaves are oval-oblong in shape with an acuminate tip. The
leaves of D. zibethinus are hypostomatic (Shanmukha Rao and Ramayya 1981).
The stipules of durian leaves are sub-falcate and deciduous (Davis and
Bhattacharya 1974). Durian leaves have several rather noteworthy characteristics.
Rumphius (1741) noted that durian leaves have distinctly swollen petiole
bases, the anatomy of which has been investigated by Funke (1931). The
vascular system of durian petioles is very complex (Solereder 1908).
Clear spots occur on the upper and lower surface of durian leaves.
Bottle shaped mucilaginous cells in the upper and lower epidermis of durian
leaves are responsible for this phenomenon (Radlkofer 1886, 1890; Solereder,
1908). Only the narrow neck of these cells lies in the epidermal layer, the
bulbous inner portion lies in the hypodermis (Radlkofer 1886). Radlkofer
(1886) reported similarly constructed clear spots in the leaves of Boschia
(now Durio).
Lamarck (1786) stated that the bottoms of the leaves and young
stems are covered by scales similar to those of Capparis. This, among other
things, led him to ally Durio with the Capparaceae. Similar parallels were
echoed by Radlkofer (1884) because of similarities in flower construction,
scales, and leaf folding between Durio and Capparis. A brief description of
the peltate scales ‘schildharre’ of durian was first given by Bachmann (1886).
Schumann (1895) described peltate scales ‘schuppe’ in Durio, and stellate
hairs in Adansonia. He reported that both forms could be found in the tribe
Matisieae. Some typical Malvaceous and Bombacaceous hairs are described
and depicted by Solereder (1908). Salma (1991) recorded five different types
of trichome morphology on the leaves of Durio, which can be useful in the
identification of species; unfortunately, no key was given. The adaxial (upper)
surface of durian leaves are smooth and apparently hairless.
Until recently, the only published description of the morphology of

young Durio and Neesia leaves was that of Burger Hzn (1972), who described
them both as conduplicate-induplicate; indicating that not only are the two
halves of the lamina folded together lengthwise (conduplicate), but that it is
the adaxial surfaces which lie against each other (induplicate). Whatley (1992)
noted that the young durian leaves (20-35 mm) used in her study of plastid
development were folded along the midrib. More recently, Brown (1994)
showed that the leaves of Durio and Neesia share this characteristic folding.
These leaves remain folded for an extended period of time, while the leaflets
of other genera unfold relatively early in development. More noteworthy is
the discovery that, when immature, the outer edges of the adaxial surfaces of
the leaves of Durio and Neesia are rimmed with stellate hairs. These hairs
differ from the stellate hairs and peltate scales of the abaxial surface in that
their arms are much longer. Close examination of the adaxial leaf margin of
immature folded leaves shows that the hairs intertwine to hold the two edges
of the leaf together in a manner similar to that of a Velcro® fastener. Examination
of adaxial leaf margins of mature leaves of Durio zibethinus failed to reveal
the presence of these hairs, indicating that they are lost after unfolding. This
mechanism appears to be a modification on an already extant theme, the
leaflets of several compound-leaved Bombacaceous species examined were
conduplicate-induplicately folded, but were not held together by hairs (Brown

1994). The two halves of the lamina probably separate from each other by
differential growth between the upper and lower leaf surfaces.
Although one would be tempted to speculate that the covering of the

abaxial surfaces of the leaves with various assorted trichomes would protect
the young leaves from herbivorous insects, especially since the hairless adaxis
is hidden by the induplicate-conduplicate folding, Gadug and Hussein (1987)
noted that the durian carsidarid makes slits beneath scales on the underside of
durian leaves.
The anatomy of leaves of Kostermansia malayana Soegeng has been

described (Baas 1972). The most striking feature of the leaves of this species
is the circular arrangement of stomata around the base of scales on the undersides
of the leaves.
A very brief account of the shoot apices of several Bombacaceae

including Durio is given by Johnson (1961). Tunica corpus zonation is apparently
less pronounced in Durio than in other non-Durioneae Bombacaceous species.
Whatley (1992) examined plastid development during leaf development

of durian and several other woody perennial tropical species. Mesophyll cells
of very young durian leaves contain chloroplasts with well developed grana.
Thyalkoids become more extensive and more deeply stacked as the leaves
develop. The plastids of fully expanded leaves often contain starch granules.
Membrane bound bodies occur in the plastids of epidermal cells.
Ashton (1978) recorded the moisture content of fresh leaves of D.

griffithii as 63%.
Roots : Differences in root development between marcotted, inarched and

seed grown trees have been noted (Polprasid 1961a). Original anchor roots
were observed only in trees grown from seed or produced by inarching, but
not found in trees produced by marcots. Further, the roots of marcotted trees,
although well distributed, did not extend as deeply into the soil as those of the
other two types (Polprasid 1961a).
D. carinatus Mast., which is endemic to marshes of Malaya, Borneo

and Sumatra, forms ‘knees’ or breathing roots. These roots emerge up to 30
cm from the water and are up to 10 cm in diameter (Thorenaar 1927). Thorenaar
(1927) notes that the ‘knees’ are looped, but the loop does not become fused
as occurs in some other marsh tree species. The bark of D. carinatus is
distinctively red with large pale lenticels (Kostermans 1958b).
Masri (1990) developed a method to quantify the root distribution

pattern of durian trees using root length density (RLD). A study of the root
distribution of durian trees (clone D24) was carried out on trees growing in
different soils types (Masri 1991). Results of this study demonstrated that the
RLD of budded D24 durian varies with environment and orchard management
techniques. Generally, 72-87% of the RLD is found in the top 45 cm of the
soil. Furthermore, 85% of the RLD was contained within the canopy radius
of the tree. These results may be of use in developing ideal fertilizing strategies
for durian. It has been reported that durian roots may form mycorrhizal
associations (Nanthachai 1994).
Tree architecture : Durian trees are large forest trees, which can reach heights
of 37 m (Foxworthy 1927), the first branch can be as much as 18-21 m off
the ground (Foxworthy 1927). The trunks of most species are normally buttressed
in mature specimens. Out of over 300 trees measured, an average diameter of
56 cm and a maximum diameter of 107 cm was recorded (Foxworthy 1927).
Larger specimens of Durio often become buttressed (Soegeng-Reksodihardjo
1962). Many Bombacaceous trees are classified as having the architectural
model of Massart. D. zibethinus has been assigned to the model of Roux
(Hallé et al. 1978), as has D. griffithii (Ashton 1978). The model of Roux is
characterized by having a monopodial orthotropic trunk and plagiotropic lateral
branches. Roux’s model differs from that of Massart largely due to diffuse
growth rather than rhythmic growth. Detailed description of the architecture
of D. zibethinus with diagrams is presented by Buisson (1986). The orthotropic
axis of durian seedlings is fast growing, producing numerous strongly plagiotropic
laterals; some orthotropic laterals are also produced which eventually compete
with the main axis (Subhadrabandhu et al. 1991).
An article by Gültekin et al. (1983) provides a dendrological description

of what is claimed to be a specimen of Durio kutejensis Becc. found growing
in the Cukurova region of Turkey. From the authors description of the tree,
and an examination of the accompanying plates, the tree is most certainly not
D. kutejensis, in fact, it does not belong to the genus Durio. The tree is most
likely a specimen of the Bombacaceous genus Chorisia or Ceiba, at any rate
a member of the tribe Adansonieae and not the Durioneae.
Chromosome number : Members of the Bombacaceae are characterized by

high chromosome numbers (Baker and Baker 1968) with 2n=72 being very
common (Baum and Oginuma 1994). It is also characteristic for the nucleolus
to persist throughout the process of mitosis (Baker and Baker 1968). Durian
stands out as an exception in having a diploid number of 2n=56 (Mangenot
and Mangenot 1958, 1962). Datta and Biswas (1969) reported a diploid number
of 2n=28 for durian. The authors attribute this difference to the sample of
Mangenot and Mangenot (1958) being tetraploid. As neither groups of authors
mentions the clone (if any) from which sample material was obtained, it may
be that different durian clones have different levels of ploidy. To the best of
my knowledge, this has never been investigated, however, it is certainly worthy
of further investigation. Soepadmo (1979) reported a chromosome number of
56 in D. zibethinus and 60 in D. griffithii. Cullenia excelsa Wight [=Durio
ceylanicus Gardn.] has a chromosome count of n=28 (Pushparajan et al. 1986),
thus there may be variation in chromosome numbers between the species.
This question never appears to have been directly addressed, but could possibly
be of value in the classification of the wild species of Durio.
Differences in chromosome number are but one of several characteristics

that distinguish the Durioneae from the other Bombacaceous tribes. Chromosome
counts of durian are lower than those of Bombacaceous species from other
tribes and closer to those found in the related Sterculiaceae, Tiliaceae and
Malvaceae (Baum and Oginuma 1994), suggesting a primitive stature of the
Durioneae within the Bombacaceae, at least in this regard.
Edibility, Composition and Uses of the Fruit

The greatest economic use of the durian is as a source of fruit, although
it is also used as a timber tree. The most frequently eaten part of the durian
is the fleshy aril which surrounds each seed. The aril is a fleshy outgrowth of
the funicular end of the seed coat (Soepadmo and Eow 1977). The aril of D.
zibethinus is commonly white, cream coloured or yellowish (Soegeng-
Reksodihardjo 1962), or even orange (Barrett 1928). Some species have bright
red arils (D. acutifolius, D. excelsus, D. carinatus, D. graveolens); the arils of
D. griffithii are orange to red and those of D. pinangianus are pinkish (Kostermans
1958b). A great variety of studies have provided information on various chemical
constituents of the fruit, from components of nutritional value to those involved
in its very characteristic smell and odour.
Edibility : Of all the known species of durian, the six commonly listed as
producing edible fruit are D. dulcis, D. grandiflorus, D. graveolens, D. kutejensis,
D. oxleyanus and D. zibethinus (Soegeng-Reksodihardjo 1962). Several other
lesser known species also bear edible fruit, such as D. testudinarum and D.
lowianus (Kostermans 1958b; Ogata 1978). Fruits of the newly discovered D.
macrantha are also edible (Anon. 1992; Kostermans 1992a,b). In addition to
these species, the fruits of D. excelsus (Korth.) Bakh. and D. pinangianus
(Becc.) Ridl. may be edible (Kunkal 1984), as are the seeds of D. carinatus
Mast., the arils of which are inedible. Kostermans (1958b) mentions that the
bright red arils of D. excelsus (Korth.) Bakh. and the pink arils of D.
pinangianus are tasteless, and the yellow arils of D. lanceolatus are almost
tasteless. The fruits of D. wyatt-smithii Kosterm. have been incompletely
described. It is possible that they are also edible as this species is very closely
allied with D. zibethinus. Beccari (1889, 1921) described his discovery of a
single tree of what he termed D. carinatus var. bintulensis Becc., which had
edible fruit (D. carinatus itself has inedible fruit) (Endert 1927a). Kostermans
has apparently made no comment on the nature or affinity of this specimen
and, according to Soegeng-Reksodihardjo (1962) since Beccari’s description,
no tree has ever been found. It is possible that other recognized species have
edible fruit as the fruit is completely unknown, or has never been described
for D. burmanicus, D. crassipes, D. kinabaluensis, D. macrolepis and D.
purpureus. Additionally, the mature fruit has never been collected or described
for several other species.
The amount of mass that the aril contributes to the entire fruit (i.e.,
the amount of edible portion of a durian fruit) varies greatly with the clone
or variety of fruit. Some recorded values are 30% (Adriano 1925), 19% (Abdullah
and Ragab 1970), 29% (Joachim and Pandittesekere 1943), 30% (Pratt and
Del Rosario 1913) and 14.5% (Intengan et al. 1955).
Numerous authors record that the seeds of D. zibethinus are also

eaten after roasting (Parsons 1932b; Ochse and Bakhuizen Van Den Brink
1977; Quisumbing 1978), or boiling (Ochse and Bakhuizen Van Den Brink
1977; Quisumbing 1978; Ridley 1902). Uphof (1968) and Usher (1974) state
that the almost tasteless seeds of D. carinatus are also eaten. According to
Ochse and Bakhuizen Van Den Brink (1977), the petals of durian are occasionally
consumed. Morton (1987), in a short review on the durian, states that the
young leaves and shoots are sometimes eaten as greens, however, there is no
original report confirming this information.
Apart from being eaten fresh when ripe, the aril is also traditionally
mixed with coconut juice, sugar, rice flour and eggs to make a cake like
concoction (dodol) (for recipes, see Soegeng-Reksodihardjo 1962; Coronel et
al., 1983; Anon. 1986b). Durian cake is now a popular contemporary commercial
product (Paweenakarn et al. 1992). Nutritional information for durian cake
has also been published (Leung et al. 1972; Ismail and Seow 1982; Paweenakarn
et al., 1992; Seow 1994). Several other commercial durian products are produced
including candy and jam. The author has also encountered durian filled swiss
rolls and donuts. At the present time, durians are not specifically grown for
the production of processed products (Maneepun et al. 1994), most commercial
durian products are likely produced using lower grade, damaged or surplus
fruits. Unripe arils are apparently eaten after roasting by the Bataks of Sumatra
(Ochse and Bakhuizen Van Den Brink 1977). Unripe arils are also recorded
as being used as an ingredient in soup (Ochse 1961). The ripe arils can be
prepared by fermenting them inside bamboo-joints, or earthenware vessels
usually for 3 to 4 days, but in some locations up to 2 weeks (Ochse and
Bakhuizen Van Den Brink 1977). The bamboo joints are either buried during
this time, or suspended over a source of smoke. The following comment from
Burkill (1966) would seem to adequately summarize the previous recipe:
‘Apologists say that the fruit should be eaten before the garlic flavour is at all
apparent. It must be added that to the depraved taste of the Besisi fermented
durian pulp, obtained by burying the aril in a time of glut, appeals’. Wallace
(1856, 1869) stated that durians were sometimes preserved with salt in jars or
bamboo joints by the Dyaks of Borneo. In the Moluccas, fish is sometimes
flavoured by smoking it above empty durian husks (Ochse and Bakhuizen
Van Den Brink 1977).
According to Rambo (1988), the fruits of durian are probably the

second most important source of carbohydrates for the Semang (aboriginal
group from central peninsular Malaysia). Favre (1848), in describing the native
Jakun, relates the following: ‘For six weeks or two months, they eat nothing
but durians. When the season is over, the place is abandoned until the next
year.’ Logan (1847) stated that the Binua of Johore actually travelled for up
to two days to favourable locations for durians, at which sites they erect
temporary huts, not returning to their homes for several weeks, until the last
durian was eaten.
Nutritive constituents : Numerous studies have reported on the nutritive

constituents of durian (Table 3). A comparison of many of the individual
estimates for a particular constituent will reveal a fair amount of variation.
This is in part due to the different and continually improving methods of
analysis employed. However, a large part of this variation is likely due to
variations between different durian fruits themselves. This is evidenced by
several studies in which more than one variety of durian have been analyzed
by the same method.
Table 3. Some nutritive constituents of durian fruits
Constitutent Fresh ripe arils of Fresh arils of

D. zibethinus D. oxleyanus
Total ash 1.242; (1.2,1.2,1.5)3*; 1.26; 1.1115; 3% of DW26

(% of fresh weight) 0.816; 1.2417; 0.818, 1.0423
Total solids 44.502
Fibre (1.7,1.7,0.9)3*; (4.40,3.35,3.47)4*; 5.9% of DW26

(% of fresh weight) 1.715; 1.416; 1.918; 0.923; 1.8710
Moisture (62.8, 56.3,66.6)3*; (64.1,58.3,57.4)4*; 30%26

(% of fresh weight) 58.05; 656; 66.07; 54.910; 68.711;
62.915; 66.816; 55.517; 59.918;
68.020*; 64.223; 60.627
Protein 2.312; (2.6,3.2,0.9)3*; (2.33,2.58.2.81)4*; 7.7% of DW26

(% of fresh weight) 2.85; 2.66; 2.57; 2.36%11; 2.515; 2.516;
2.3117; 2.018; 2.723; 2.627
Nitrogen 0.211%10
Carbohydrates (total) (29.4,34.7,23.9)3*; (21.33,29.32,27.81) 4*; 65% of DW26

(% of fresh weight) 34.15; 27.86; 25.0111; 30.415; 28.316;
36.118; 28.223
Starch (% of fresh 11.117; 4.020*

weight)
Sugar total 13.5517; (10.02, 11.14, 13.38)19* 7.7% of FW26

(% of fresh weight) (11.1, 13.4)22*; 11.020*
Reducing sugars 4.79%2; 2.7%11
Sucrose 8.762; (12.6, 19.8, 19.4)4*; 5.811;

(% of fresh weight) 9.16, 10.22, 12.7019*; 10.2, 12.722*
Glucose 0.30, 0.51, 0.48%19*; 0.5,0.5%22*
Fructose (0.55, 0.41, 0.20%)19*; 0.4,0.2%22*
(Contd....)
Table 3. Contd.
Lipid (2.3,2.9,4.2)3*; (6.29, 5.38, 7.34)4*; 3.95; 19.0% of DW26

(% of fresh weight) 3.26; 5.007; (5.1,5.2,3.8,4.2)8*
(0.71, 0.91)10*; 2.1011; 3.115; 1.616;
1.218; 3.023
Caloric value (149, 178, 145 kcal/100g)3*

(151.2, 176.0, 188.5 kcal/100g)4*
183 cal/100g5 128.411
144 cal/100g15 124 cal/100g16
147.0 cal/100g18 154 cal/100g23
Beta carotene (710, 600.5, 1398.3 IU/100g)4*
Vitamin A (-,-,1025)3*; (-,-,30.0)4*; 205; trace12;

(IU per 100 g) 3015; 10µg/100g16; trace18; trace23;
unless otherwise noted 3.8mg/g24*
Vitamin B1 (0.52, 0.67, 0.47)3*; (0.24, 0.36, 0.39)4*;

(thiamine) 0.36610; 46 IU/100g14; 0.2415;
(mg/100g) unless 0.2716; 0.3218
otherwise noted
Vitamin B2 (0.49,0.53,0.17)3*; (0.07, 0.13, 0.14)4*;

riboflavin 0.17210; 0.2015; 0.2916; 0.2818
(mg/100g)
Niacin (1.17, 1,17, 1.37)4*; 1.1310; 0.715;

(mg/100g) 1.216; 1.118
Vitamin C (32.5 ± 5.4)1; (32,43,58)3* 2.08 mg/100ml26

(mg/100g) (31.0, 43.3, 41.3)4*; 50.0010; 2511;
23.413; 2415; 3716; 44.018; 22.920*
10721*
Vitamin E (mg/100g) (1.50 ± 0.26)9
Minerals 1.2%5
Calcium (5.6, 4.5, 5.9)3*; (5.35,4.64,5.10)4*; 105; 0.03 % of DW26

(mg/100g) 12.1410; 41.511; 915; 2016; 1818; 4023
(Contd....)
Table 3. Contd.
Phosphorous (27.7,28.3,19.6)3*; (42.0, 36.3, 36.6)4*; 0.13% of DW26

(mg/100g) 505; 65.3910; 40.011; 4415; 6316;
56.018; 4423
Magnesium (mg/100g) 3316; 33023 0.08 µg/g26
Iron (1.0,1.1,0.8)3*; (0.80, 0.38, 0.55)4*; 1.05; 17 µg/g26

(mg/100g) 1.1310; 0.915; 0.916; 1.118; 1.923;
1.1 mg/g25
Cobalt 0.4 mg/100g16; 0.03mg/g25
Chlorine 4 mg/100g23
Sodium (mg/100g) (0.57, 0.59, 0.67)4*; 116; 4023
Potassium (mg/100g) 474.6, 431.3, 488.14*; 60116 1.59% of DW26
K2O 70 mg/100g23*
SO4 180 mg/100g23
Arsenic (0.0007, 0.0004, 0.0002 mg/100g)4*
Copper 2.2 g/100g23; 1.0 mg/g25 21 mg/g26
Manganese 7.2 g/100g23; 0.81 mg/g25 12 mg/g26
Iodine 2.8 g/100g23
Cadmium <0.01 mg/g25
Chromium <0.05 mg/g25
Nickel <0.02 mg/g25
Lead <0.03 mg/g25
Zinc 1.4 mg/g25 17 mg/g26
Mercury <0.01 mg/g24
(Contd....)
Table 3. Contd.
pH 7.0 ± 0.041; (6.77, 6.66, 6.60)19*; 6.66,

6.0022*; 6.620*
Acidity (as citric) 0.192
Sugar: acid ratio 112.520*
Acidity (others) (0.09 ± 0.02 g/100g)1; 52.3 cc N/10

per 100 g11; 0.1%20*;
0.6, 0.8 meq/100g22*
1
Abdullah and Ragab (1970).
2
Adriano (1925) [Note: Data originally from Pratt and Del Rosario 1913].
3
Anon. (1973) *Note: This original Thai article has several entries for durian, data for only three
cultivars Kradum Thong, Kan Yao and Chanee are reprinted in Woller and Idsavas
1981. Only these three respectively are presented in this table.
4
Anon. (1989b) *Note: Data on three cultivars Chanee, Mon Thong=Golden Pillow, and Ocean
Petal are given respectively. Data is reprinted in Maranet (1991).
5
Aykroyd (1963).
6
Bauchau (1972).
7
Berry (1980b).
8
Berry (1980c) *Note: Data on four clones is given-D24, D2, D66, D8 respectively.
9
Candlish (1983).
10
Intengan et al. (1955) *Note: Two values for lipid are given fat and fat ash respectively.
11
Joachim and Pandittesekere (1943).
12
Leong (1939a), 13Leong (1939b), 14Leong (1940).
15
Leung et al. (1952).
16
Leung et al. (1972).
17
Manas Y Cruz et al. (1939).
18
Martin (1980) [Note: Data originally from Abdon et al. 1980].
19
Niyomporn et al. (1984) *Note: Data on three cultivars Kan Yao, Chanee and Ruong are given
respectively.
20
Pauziah et al. (1992) *Note: All values are from analysis of fresh fallen mature fruits of clone
D24.
21
Phang (1976) *Note: Authors state that their result is likely artificially high.
22
Preungvate (1982) *Note: Data on two varieties Chanee and Ruong respectively are given.
23
Rosedale (1935) [Note: Data is reprinted in Willimot 1949.] *Note: The estimate of potassium
content based on amount of K2O is likely a gross underestimate.
24
Speek et al. (1988) *Note 3.8 mg/g total carotenoid determined by spectrophotometry was
reported with 1.9 mg/g as b-carotene as determined by reverse phase HPLC.
25
Wong and Koh (1982).
26
Wong (1992).
27
Zanariah and Noor Rehan (1987).
The edible aril of the common durian (D. zibethinus) is a good source
of vitamin C (Table 3), which is of the order of 33 mg per 100 g of aril
(Abdullah and Ragab 1970). This is roughly equivalent to the vitamin C
content of many citrus fruits. Phang (1976) estimated that durian arils contain
100 mg of vitamin C per 100 g of aril, however, he also warned that this
estimate was likely inflated due to interference from mercaptans, naturally
present in the aril, with his assay. This is likely a problem associated with all
published estimates of the vitamin C content of durians, thus all published
values are likely overestimates. Rosedale (1935) used a Guineapig bioassay to
estimate the vitamin C content of durian and various other foodstuffs. He
assigned an antiascorbic value to durian of 10 grams (the amount of durian
necessary to alleviate the symptoms of scurvy) which was approximately only
one third that of citrus fruits. This result supports the suggestion that published
vitamin C contents for durian are overestimations.
The average mature fruit weighs in excess of a kilogram (Abdullah

and Ragab 1970), yet the edible arils generally make up less than one third
of the mass, and the seeds only about 15% (Abdullah and Ragab 1970). The
aril contains about 33% carbohydrates on a fresh weight basis (Table 3), of
which about one third is probably starch. The carbohydrate of durian apparently
consists of mannans (Martin 1980), and an erythrodextrin, which gives a clear
red colour upon reaction with iodine (Pratt and Del Rosario 1913). Two
relatively old studies provided estimates of the reducing sugar content of
durian arils (Table 3). These estimates (4.79 and 2.7%) grossly exceed the
amounts of the two major reducing sugars (glucose and fructose, together
totalling only 1% of the fresh weight at most) reported in more recent studies.
These estimates of reducing sugars were in both cases produced by an unspecified
copper reduction assay which strictly speaking measures total reducing substances
present, not just sugars. In the case of durian, this would include mercaptans
and ascorbic acid (vitamin C) which are known to be present in substantial
quantities within the mature fruit. Thus, the published values for reducing
sugars are large overestimates. Measurements of percentage of fructose, glucose
and sucrose from ripe durian fruits, obtained by high-performance liquid
chromatography (HPLC), have recently been published (Freeman and
Worthington 1989).
Fresh durian arils contain 2-2.5% protein (Table 3). The total and
essential amino acid composition of durian arils has also been investigated
(Zanariah and Noor Rehan 1987) on a ‘per gram fresh weight’ basis. The data
show that durian is a better source of all the essential amino acids than dates,
peaches, oranges, mangoes, cempadak or papaya (Table 4). However, it should
be remembered that durian has a much higher percentage of total protein than
all these fruits.
Table 4. Amino acid composition of durian fruits
Amino acid composition Essential amino acids

(mg/100g FW) (g/16g N)
Isoleucine 85.8 Lysine 4.8

Leucine 143 Histidine 2.0
Lysine 124.8 Arginine 2.1
Methionine 44.2 Aspartic acid 9.3
Histidine 52 Threonine 2.6
Cystine 78 Serine 3.9
Phenylalanine 78 Glutamic acid 11.9
Tyrosine 57.2 Proline 3.8
Threonine 67.6 Glycine 4.1
Valine 122.2 Alanine 8.4
Cystine 3.0
Valine 4.7
Methionine 1.7
Leucine 5.5
Isoleucine 3.3
Tyrosine 2.2
Phenylalanine 3.0
Source : Zanariah and Noor Rehan (1987).
Leung et al. (1952) found that fresh arils are high in several B vitamins.
Durians are also an excellent source of vitamin E (Martin 1978). Visetbhakdi
(1988), in a brief description of durians and the economics of their production
in Thailand, mentions that they are high in cholesterol; but authentic reports
are lacking as to the cholesterol measurements of durian fruits.
The fruit has been examined and tested positive for thermostable
anti-thiamine (thiaminase) activity (Rattanapanone 1979). Bate-Smith (1959)
described the presence of caffeic acid and small amounts of leuco-anthocyanins
in durians (although this was probably from an analysis of the leaves). Baldry
et al. (1972) listed several alcohols present in the aril including ethanol,
methanol and n-propanol. Wong and Tie (1995) have recently questioned the
presence of ethanol and methanol. Numerous other compounds have been
identified in fresh durians (Table 5). The creamy consistency of the aril is
attributed to gums, pectins and hemicelluloses (Martin 1980).
Phenolic compounds have been reported in durian (tissue unknown,

probably from leaves), these include flavanols and caffeic acid. Ferulic and
Table 5. Volatile compounds identified in the aroma profile of durian

fruits
Volatile compound Source

Acetaldehyde 1,not 3
Alkyl hydropolysulphides 2
Butan-1-ol 3
Butane-2,2-diol 3
Butanedione 3
Butyl acetate 3
Butyl propanoate 3
g-butryolactone 3
Dialkyl polysulphides 2
1,1-diethoxyethane 2,not 3
Diethyl carbonate 3
Diethyl disulphide 1,2,3
Diethyl tetrasulphide 2,not 3
Diethylthioether (diethylsulphide) 1,not 3
Diethyl trisulphide 2,3
Dimethylthioether (dimethylsulphide) 1,not 3
Cis-3,5-dimethyl-1,2,4-trithiolane 3
Trans-3,5-dimethyl-1,2,4-trithiolane 3
Dodecan-1-ol 3
Ethanol 1,not 3
Ethanethiol (ethanediol) 1,3
Ethyl acetate 1,2,3
Ethyl benzene 1,not 3
Ethyl butanoate (ethyl butyrate) 1,3
Ethyl (E)-but-2-eonate 3
Ethyl decanoate 3
Ethyl dodecanoate 3
1-(ethylthio)ethanethiol 3
Ethyl heptanoate 3
Ethyl hexanoate 3
Ethyl 3-hydroxybutanoate 3
Ethyl 2-hydroxypropanoate 3
Ethyl isovalerate 1
Ethyl methacrylate 1,not 3
ethyl(E)-2-methylbut-2-enoate 3
Ethyl 2-methylbutanoate
(ethyl a-methylbutyrate) 2,3
Ethyl 3-methylbutanoate
(ethyl isovalerate) 1,3
(Contd....)
Table 5. Contd.
Ethyl methyl disulphide 3

Ethyl 2-methylpropanoate
(ethyl isobutyrate) 1,3
Ethyl (methylthio)acetate 3
Ethyl methyl trisulphide 2,not 3
Ethyl octanoate 3
Ethyl pentanoate 3
Ethyl propanoate
(ethyl propionate) 1,3
Ethyl propyl disulphide 2,3
Ethyl propyl trisulphide 2,3
S-ethyl thioacetate 3
Heptan-1-ol 3
Hexadecane 3
Hydrogen sulphide (not due to
microbial action) 1,2,not 3
3-hydroxybutan-2-one 3
4-hydroxyhexan-3-one 3
3-hydroxypentan-2-one 3
2-hydroxypentan-3-one 3
Methanethiol 1,not 3
Methanol 1,not 3
Methyl acetate 1,not 3
Methyl butanoate 3
2-methylbut-2-enal 3
2-methylbutan-1-ol 3
3-methylbutan-1-ol 1,3
Methyl hexanoate 3
Methyl 3-hydroxybutanoate 3
Methyl 2-methylbutanoate
(methyl a-methylbutyrate) 1,3
Methyl octanoate 3
2-methylpropan-1-ol 3
Methyl propanoate
(methyl propionate) 1,3
Methyl propyl disulphide 3
Nerolidol 3
Propanethiol 1,3
Propan-1-ol 1,3
Propionaldehyde 1,not 3
Propyl acetate 3
Propyl butanoate 3
(Contd....)
Table 5. Contd.
Propyl 2-methylbutanoate
(N-propyl a-methylbutyrate) 1,3
Propyl 2-methylpropanate 3
Propyl propanoate
(N-propyl propanoate) 1,3
S-propyl thioacetate 3
S-propyl thiopropionate 3
2,4,6-trimethyl-1,3,5-trithiane 3
1=Baldry et al. (1972), 2=Moser et al. (1980), 3=Wong and Tie (1995).
sinapic acids are reported absent and leuco-anthocyanins present in low amounts
or absent (Bate-Smith 1959).
Fatty acids : The fresh arils of durian fruits consist of about 3-5% fat (Table 3).
The exact amount of fat seems to vary between clones as does the nature of the fats
themselves. The aril of the durian was first analyzed using gas chromatography by
Aspiras and Tocino (1971), but no conclusions about its constituents were made.
The doctoral thesis of Greve (1974) included an analysis of the fatty acid
composition of durian seeds, arils and husks collected from two locations in
Thailand (Table 6). Not only does the fatty acid makeup differ between these three
tissues, but it differed markedly between the two varieties of durian. Most notably,
20:0 fatty acids constitute approximately 33% of the total fatty acids found in the
husk (pericarp) and almost 10% of that of the seeds in Prajeen Rayong durians, but
were found in negligible amounts in all tissues of Chanthaburi durians (Greve
1974); variance in the fatty acid makeup of the arils was much less between these
two varieties than that found in the pericarp and seeds. Berry (1980a,b,c) reported
on the lipid constituents of both the arils and seeds of durian, in particular the
presence of cyclopropene fatty acids which are present in many related genera,
and which may be carcinogenic (Berry 1980b). He concluded that cyclopropene
fatty acids are present in the seeds in unusually high quantities, 65% of the total
fatty acids in durian seed oil are cyclopropenoids (Berry 1980a). Cyclopropenoid
fatty acid content was reduced, but not eliminated upon cooking. Sterculic,
dihydrosterculic and malvalic acids were present in the uncooked seeds but not in
the aril. Due to the toxic and perhaps carcinogenic nature of these substances,
Berry (1980b) concludes that it would be unwise to ingest uncooked durian seeds.
Berry (1981) compared fatty acid composition of four durian clones and linked the
palmitic:palmitoleic acids ratios with taste. The lower this ratio, the higher fruits
were ranked by a panel of tasters. He also noted that the degree of saturation of
fatty acids varies between clones, which may influence retention of volatile
flavouring compounds, and hence the flavour itself. However, at present, this
relationship remains correlative rather than causative.
Smell : Although the fruits of Durio zibethinus are infamous for their foul
smell, they apparently are not the most offensive of all the known species.
Most species of Durio produce odourless, or nearly odourless fruits, however,
the odour of the ripe (and edible) fruits of Durio dulcis is apparently so vile
and pervasive that they can be smelled for miles (Soegeng-Reksodihardjo
1962); Kostermans (1958b) describes the smell of these fruits as ‘simply
nauseating’. Hayes (1957) recorded that it is believed that the smell of durian
fruits can be eliminated by soaking the flesh in coconut milk overnight. I
have, however, seen no recorded experiments on this procedure.
Capus and Bois (1912) described the smell of durians as alliaceous

and stercoraceous. Barrett (1912) suggested that the smell might be due to
sulphur compounds with some base perhaps related to butyric acid. In recent
years, the exact nature of the volatile components that compose the infamous
smell have been subjected to some very sophisticated analysis including mass
spectroscopy, gas chromatography and 1H nuclear magnetic resonance.
Stanton (1970) and Baldry et al. (1972) divided the smell into two
distinct categories, a very strong onion-like smell (alliaceous), probably caused
by thioethers; and a second fruity smell due to the presence of esters. Stanton
(1966) suggested that indoles and skatols may be present in the fruit accounting,
in part, for its malodorous nature. Stanton (1970) presented evidence that the
arils do not release thiols, and thus thiols are not responsible for the smell.
However, esters and thioethers were implicated. Baldry et al. (1972) analyzed
durians obtained from Singapore which gave similar results, but a durian
obtained from Kuala Lumpur was found to owe a large portion of its odour
to thiols with minimal contribution by thioethers. The major volatile components
were found to be ethanol, n-propanol and ethyl a-methylbutyrate [=ethyl 2-
methylbutanoate]. This issue was further addressed by Moser et al. (1980)
who provided evidence that durian arils do not contain thiols and that mature
arils do release hydrogen sulphide, while those of immature durians do not.
They also provided evidence that the release of hydrogen sulphide is not due
to bacterial action in the fruit (as has occasionally been suggested), since
tissue from immature arils did not yield any colonies when cultured on substrates
for H2S forming species. Martin (1978) stated that hydrogen sulphide and
diethyldisulphide are responsible for the foetid odour. Durians smell more as
they mature; analysis of H2S production by 500 grams of durian at different
stages of maturity produced the following results: immature durian arils produce
0.009 g H2S, ripe durian arils 0.023 g and very ripe durian arils 0.047 g
(Greve 1974).
Moser et al. (1980) ascribed the characteristic odour of durian to

H2S, hydrodisulphides, dialkyl polysulphides, ethyl esters and 1,1-diethoxyethane.
Indoles were not detected as was predicted by Stanton (1966). The polysulphides
identified by Moser et al. (1980) were mainly ethyl polysulphides which
apparently rarely occur in fruits. Unfortunately, Moser et al. (1980) obtained
their durians from Thailand. Thai durians differ substantially in odour and
flavour from Malay durians, the Thai durian being sweeter and considered
less aromatic than Malay clones (Mohamad Idris 1987). Thus, the differences
in constituents found between the study of Moser et al. (1980) and Baldry et
al. (1972) may either reflect clonal differences in aroma composition or
differences due to technique.
Very recently, a third analysis of the aroma composition of durian

fruits has been published. This study by Wong and Tie (1995) reported the
presence of 63 volatile constituents (including 30 esters, 5 ketones and 16
sulphurous compounds), which contribute to the smell. More importantly, this
study sheds further light on the work of Baldry et al. (1972) and Moser et al.
(1980). Interestingly, this work demonstrated several differences from the
earlier analyses (all the volatile compounds as yet described in durian fruits
are listed in Table 6).
Wong and Tie (1995) were unable to confirm the presence of several
of the volatile compounds reported in both previous studies including ethanol,
methanol, ethyl methacrylate and several sulphur containing compounds. They
attribute these differences to better technique, and to using ripe durians and
a minimal time between harvest and analysis. There are three interesting
aspects of this study, asides from a long list of new volatile components.
Firstly, Wong and Tie (1995) sampled and compared the composition of
fruits from three named clones, D15, D28 and D74. Not only did they show
considerable differences in the total yield of volatile compounds between
clones, but numerous large differences in quantities of particular volatiles
exist between these clones. Some volatiles were not detectable in all of the
clones. Secondly, Wong and Tie (1995) demonstrated the presence of three
different a-hydroxyketones which they claim have never before been reported
from an analysis of a fruit. Thirdly, these authors stated that no trace of
H 2S was detectable in the aroma profile of the fruits from any of the three
clones tested. This result is at variance with several previous works (Baldry
et al. 1972; Greve 1974; Martin 1978; Moser et al. 1980) and is likely to
be related to change in aroma composition over time during ripening of the
fruit. The H2S production increases markedly as the fruit ripens (Greve 1974).
Thus, although Wong and Tie (1995) used ripe fruit, it may not have been
ripe enough to produce H2S. Apart from the study of Greve (1974), the change
in aroma composition of durian fruits over time has not been examined. Such
Table 6. Fatty acid composition of durian
Tissue Clone Saturated Palmitic 14:0 15:0 16:0 16:1 17:0 18:0 18:1 18:2 18:3 20:0 22:0 Dihy- Mal- Ster- Cis
unsatur- Palmit- drost- valic culic vaccenic
ated oletic erculic
Aril D241 1.4 4.7 0.5 <0.1 39.8 8.5 <0.1 0.8 45.8 1.8 2.7
Aril D2 1 1.7 6.9 0.5 <0.1 35.9 5.2 <0.1 1.0 51.0 2.4 4.1
1
Aril D66 1.8 5.3 0.8 <0.1 33.8 6.4 <0.1 0.9 48.7 2.8 7.0
Aril D8 1 1.9 13.9 0.5 <0.1 32.3 2.3 <0.1 2.2 53.6 3.2 6.0
Aril Unknown 0.91 34.13 7.1 1.21 42.14 7.85 5.69 <0.01
12
Aril Unknown 0.34 28.94 5.16 1.23 5.98 3.16 2.21 <0.01
22
Whole Unknown2 82% 0.12 12.20 1.15 1.42 8.42 6.50 11.3 1.21 2.52 15.72 38.53
fresh unsaturated
seeds
Pericarp Unknown 30.9 4.2 1.8 52.8 4.2 4.0 <1
Thai#13
Pericarp Unknown 14.9 1.4 5.1 17.8 7.6 4.9 33.2
Thai#23
Aril Unknown 33.9 4.9 1.0 51.2 3.0 5.1 <1
Thai#13
Aril Unknwon 31.9 6.9 1.0 51.0 3.0 5.4 <1
Thai#23
Seed Unknown 26.8 8.4 3.3 38.8 5.9 3.0 <1
Thai#13
48
(Contd...)
Table 6. Contd.
Seed unknown 23.0 3.1 1.2 41.0 11.5 5.3 9.5
Thai#23
Aril unknown 0.6 31.3 3.4 2.1 (47.1, 1.3 7.1 8.4
Thai4 4.3)*
Fatty acid compositon of various parts of durian fruits. Note: all values are in percent GLC peak area except for that of Shibahara et al. which is
in % wt from GC analysis.
1
Berry (1980c) and Berry (1981).
2
Berry (1980b) (Unknown#1 was from a Malay durian with yellow arils, #2 was from a Malay durian with cream coloured arils).
3
Moser et al. (1980) (Thai#1 was a durian from Chandburi, Thai#2 was from Prajeen Rayong.) Note: This data is originally from the thesis of Greve
(1974).
4
Shibahara et al. (1987) *Note: Vaules are for 18:1 n-9, n-7 respectively.
49
a study would be enlightening, as it is possible that the aromas of fruits

from different durian clones may not just simply differ in the presence or
absence or relative amounts of certain volatiles, but there may be timing
differences in the production and release of these compounds that exist
between clones.
Though numerous compounds have now been identified in the aroma

profile of durians (Table 6), these contribute only marginally to the overall
smell. The major problem that still remains is the determination of the components
most responsible for the distinctive smell of durians. This is confounded by
four problems. Firstly, durians from different clones appear to vary markedly
in aroma composition, and the aroma composition may vary with ripeness,
thus there would be no fixed ‘recipe’. Secondly, of the three major published
studies on aroma composition, very different conclusions regarding the major
contributing compounds have been reached. Thirdly, how much a volatile
substance contributes to the aroma of a fruit is not necessarily indicated by its
relative abundance, some substances simply smell much more than others.
Finally, all other concerns aside, the relative abundance of a compound itself
is likely to be misleading, and must be interpreted with caution. Relative
abundances are usually expressed as an area percentage of the total response
area, taken by the response peak for a particular compound. These areas have
not been corrected for differences in detector response that exist between
different compounds. Thus, a compound that appears to exist ten times the
quantity of another substance because its peak area is ten times greater would
actually be present at the same concentration if the sensitivity of the detector
to the compound was ten times greater. I could not sum up the wonderful
complexities of the smell and taste of durians better than Martin (1980) who
stated: ‘Thus, there appear to be a wide variety of durian fruits, each combining
foul, sulphurous compounds in its own delightful way...’.
Miscellaneous uses of fruit : Durian husk extracts have been studied for
their suitability as aqueous binders for granules and tablets (Umprayn et al.
1990a,b). These authors showed that husk (pericarp) extracts are suitable
for the preparation of granules and to have desirous binding properties for
the manufacture of tablets (Umprayn et al. 1990b). The ashes of D. zibethinus
are used to extract a dye which, in turn, is used to prepare batik dye
(Kostermans 1958b). The ashes have also been used for whitening silk
(Kostermans 1958b; Morton 1987). The empty husks are also used in Java
as a source of fuel (Ochse and Bakhuizen Van Den Brink 1977; Morton
1987).
Medicinal and Toxicological Properties
Durians and alcohol : A common Malaysian belief is that it is harmful to

drink alcohol after consuming durians. This belief dates back at least to the
time of Rumphius (1741) who states that one should not drink alcohol after
eating durians as it will essentially cause indigestion and bad breath. Gimlette
(1929) in his ‘Malay Poisons and Charm Cures’ records: ‘It is said that the
durian fruit must not be eaten with brandy’. More recently, Croft (1981)
states that ‘...a feeling of morbidity often follows the consumption of alcohol
too soon after eating durian’. Some scientific studies have actually been
conducted to investigate the validity of this belief.
For the record, there is, in fact, a published medical instance of a

middle aged Indian woman dying after eating a durian and consuming alcohol
(Singh 1941). In this case, the woman was admitted to hospital complaining
of intense epigastric pain, and persistent vomiting. Despite attempts to save
her, she died several hours after admission. Autopsy revealed fat necrosis
on the peritoneum, a reddish and swollen pancreas, a swollen main pancreatic
duct, and 2 pints of blood stained fluid in the abdominal cavity. It was
suggested by one physician that the death was due to acute haemorrhagic
pancreatitis, the ingestion of durians that morning being merely coincidental;
the patient had a history of epigastrium pain and discomfort for 7 months
preceding her death. A second physician concurred and added that these
conditions may have been caused by alcoholism. Oddly, the opinion of the
author of the report was that the ingestion of durians and the death due to
acute haemorrhagic pancreatitis was not coincidental, but perhaps causative.
No explanation for this opinion was presented.
Two Singaporean researchers attempted to probe the effects of durian

and alcohol mixtures, in two experiments, by force feeding mice durian-
alcohol homogenates containing 0.8 g durian and 10%(v/v) ethanol (Ogle
and Teh 1969). Mice were fed homogenates at different times after preparation
in the first experiment, controls were fed either just durian or just 10% ethanol.
The alcohol-alone mice and the alcohol-durian mice behaved similarly, all
had an unsteady gait. Mice fed just durian behaved similarly to untreated
mice. When alcohol was administered at different times after force-feeding
of durian, it was found that the sooner alcohol was administered, the fewer
mice fell asleep, the lower the mean sleeping time, and the quicker the onset
of sleep. The authors proposed that durian may inhibit the absorption of
alcohol. Furthermore, the authors cited personal experience with persons who
have consumed durians with alcohol:
‘The alcohol was taken together with, or 1 to 6 hours after a durian

meal. None of them felt any ill effects, except for one person who experienced
discomfort from fullness of the abdomen and flatulence.’ -- Ogle and Teh
(1969).
In a second paper, Ogle and Teh (1971) investigated the possibility

that the ill effects of durian and brandy cited by Gimlette (1929) might be due
to some constituent in the brandy other than alcohol. In these experiments,
mice were given homogenates consisting of 0.8 g of durian, and then fed 1
ml of 10% brandy or whisky 2, 4, or 6 hours later. Durian controls showed
no ill effects, while those receiving alcohol and durian showed an unsteady
gait. The authors report that those receiving alcohol after 2 hours appeared to
have a more steady gait than the others. The authors again suggest that durian
may inhibit the absorption of alcohol.
Techapaitoon and Sim (1973) have also investigated the effects of

durian and alcohol mixtures. These experiments have corrected some of the
problems present in the first study (the effects of lethal dose was studied,
larger sample sizes were used, and the variety ‘Koh’ of durian chosen was
recorded). These authors performed a series of experiments on mice and rats.
In one experiment, they orally administered the lethal dose of alcohol (1 ml
of 37% alcohol) to 20 mice. The authors noted that, after administering the
lethal dose of alcohol, all mice subsequently died. More interestingly, in a
group of mice fed the same amount of alcohol together with durian homogenate,
only 11 of the 20 mice died. Furthermore, only 7 of 20 mice fed similar
concentrations of Hennessey V.S.O.P. reserve brandy-durian homogenates died.
Attempts to feed sleeping mice durian homogenates after feeding them the
lethal dose of alcohol resulted in the deaths of all mice involved. Further
experiments by the authors revealed that mice given durian together with
alcohol, or immediately after, reduced the number of mice that fell asleep,
and shortened the average time spent sleeping. These effects were noticed
when mice were given alcohol orally or intraperitoneally. Finally, mice or rats
fed durian and alcohol homogenates appeared to have a more steady gait than
did alcohol controls. The experiments of Techapaitoon and Sim would appear
to confirm and expand the findings of Ogle and Teh (1969, 1971).
More recently, a Thai research paper has again studied the effect of
alcohol-durian mixtures on mice and rats (Nilvises and Saengsirinavin 1986).
Similar results to those of the previous studies were found. Furthermore, this
study revealed that similar results are obtained with durian fruits of several
different clones.
Despite the important contributions to our knowledge of durians

and human/mouse physiology provided by these experiments, several critical

flaws are evident. And a priori assumption made by these authors is that the
effects of durians and alcohol on humans is equivalent in mice. Mice might
be less sensitive, or not sensitive at all. Ogle and Teh (1969) did, however,
give some consideration to the mouse human scaling factor, thus durian
doses given to mice were roughly equivalent to an adult male human consuming
6 pounds of fruit. Most importantly, all of the aforementioned experiments
were lacking a control treatment in which animals were fed an equivalent
mass of some food item other than durian. Thus, any effects on alcohol
absorption might be due to the presence of food in the stomach, and not
durian per se.
In considering the available evidence, experiments with mice and

rats, observations of humans (by Ogle and Teh), and the recorded medical
case of Singh (1941), it is fair to say that consumption of durians with alcohol
has not been shown to be harmful.
It should be noted that deaths or maladies apparently do occasionally

occur due to durians, but usually from being struck by one of these large
spiny fruits as it falls from a tree, rather than by ingestion (Craig 1973). If the
much quoted article by Wallace on durians can be relied upon, actual death
by striking is in fact quite unusual:
‘When a durian strikes a man in its fall, it produces a fearful wound,

the strong spines tearing open the flesh, while the blow itself is very heavy;
but from this very circumstance, death rarely ensues, copious effusion of
blood preventing the inflammation which might otherwise take place’ (Wallace
1856).
Medicinal properties : The effects on human physiology of the ingestion of

durians has been discussed for centuries. James Low (1836) wrote: ‘He who
can eat and digest a durian, and not find his liver stirred up by a host of blue
imps, may well despise the anti-dispeptic precepts of a Kichener, a Sinclair
or a Johnstone’. The durian fruit is often cited as having a ‘warming effect’
on the body (Rumphius 1741; Malo and Martin 1979), but this property does
not seem to ever have been investigated.
Paludan in Linschoten (1579-1592) (Anon. 1851, an English translation)

claimed that betel leaves when enclosed in a room with durians will cause
them to rot. He also claims that a betel leaf laid upon the stomach, or eaten
after eating durians will ease digestion. This belief has been echoed through
several works since Paludan.
Febrifugal and anti-malarial properties : There are numerous suggestions

that different parts of durian trees have been traditionally used in remedies for
fevers. The juice of fresh leaves has been used as an ingredient in a lotion for
fevers (Gimlette and Burkill 1930). The juice from the bark has been used to
attempt to treat malaria (Panggabean 1975). Ridley (1906, 1907) states that
the roots are used for treating fever, either ground up and rubbed on the body
or as a decoction. According to Brown (1954), a decoction of the roots is
made for fevers when the fever has lasted 3 days; also a compound is made
out of the leaves and roots for fever. The most complete description of this
medicinal use is a native Malayan prescription for fever, collected by Burkill
and Haniff (1930): (1) Boil the roots of Hibiscus rosa-sinensis with the roots
of Nephelium longan, Durio zibethinus, Nephelium mutabile and Artocarpus
integrifolia, (2) drink the decoction, (3) also, boil the leaves of all of these
species and use as a poultice. Gimlette and Burkill (1930) state, as a remedy
for fever, that the leaves of Curculigo latifolia, Gleichenia linearis, Nephelium
lappaceum and Durio zibethinus should be squeezed by hand, and water poured
on them. The patients head should be bathed in this water for three days.
Finally, in this regard, Heyne (1950) states that a concoction made from the
bark of D. oxleyanus is used to treat malaria in Sumatra.
Although durians obviously do not cause malaria, it has been noted

that many malarial infections can be traced to individuals who have stayed
up, unprotected, through the night in durian orchards to collect fruit as it falls
(Ponnampalam 1975).
Vermifugal properties : Hurrier and Perrot (1907), and Morton (1987) both
report that durian is used as a vermifuge. De Padua et al. (1978) reported it
to be a vermifuge, vermicide and an anthelmintic. I have, however, been
unable to locate any prescription for deworming based on durians, or to find
any record of what part of the plant is supposed to have this property.
Treatment of jaundice : It appears that the leaves have been used in the
traditional treatment of jaundice (Brown 1954). Burkill and Haniff (1930)
have recorded the following traditional prescription for the treatment of jaundice:
‘Boil leaves in water and bathe in it.’
Diabetes : An investigation of the post-prandial glucose and insulin responses

of diabetes patients to 5 tropical fruits including durian revealed that the
insulin response, and insulin area was the greatest after ingestion of durian.
Thus, they are not optimal food items for persons with non-insulin dependant
diabetes mellitus (Roongpisuthipong et al. 1991). The reason for the greater
effect of durian on insulin levels in the blood compared to other fruits is still
unknown.
Aphrodisiac : The durian fruit is frequently claimed to have strong aphrodisiac

properties, as do many foul things (Rumphius 1741; Baillon 1875; Ridley
1902; Popham 1979 and others). In fact, an often quoted Malay saying states
‘When the durians come down, the sarongs go up’. Despite the widely held
nature of this belief, no experiments to determine its empirical validity have
been conducted.
Miscellaneous medicinal properties : Morton (1987) states that ashes of the

pericarp are taken after childbirth. The fruit is also said to be an abortivum
and to improve menstruation (Kostermans 1958b). Kostermans also relates
that the valves of the fruit are rubbed on the abdomen against constipation.
Kostermans (1958b) and De Padua et al. (1978) report that durian husks are
used externally to treat skin diseases. The leaves probably contain hydroxy-
tryptamines and mustard oils (Stanton 1966; Morton 1987). The alleged action
of durian leaves and husks in reducing swelling and aiding skin diseases may
be due to the vaso-constrictor properties of hydroxy-tryptamines and bactericidal
action of mustard oils (Stanton 1966).
Ridley (1902) states that durian is believed to be strengthening for

children. De Padua et al. (1978) claim that durian is used as a general tonic,
however, no specific information is given. Heyne (1950) mentions that crushed
seeds of D. oxleyanus are used to treat sores and wounds.
De Padua et al. (1978) recorded the presence and relative amounts

of several components in the leaves and stems of durians, which may, in
part, play a role in suspected medicinal qualities. They found detectable
levels of tannins in durian leaves, and abundant to very abundant amounts
in the stem. Furthermore, there are detectable to abundant levels of saponins
in the leaves and stems, detectable levels of fats in the leaves and stem,
detectable to abundant quantities of calcium oxalate in the stem, and detectable
levels of formic acid in the leaves; abundant amounts in the stem. Ashton
(1964) records that leaf extracts of D. zibethinus and D. graveolens react
with FeCl2 causing a blue colouration, indicating the presence of tannins.
Furthermore, the leaves of both species tested negative for protein precipitating
compounds.
According to many authors, durian seeds contain a poisonous substance

that makes one short of breath (Rumphius 1741; Kostermans 1958b; Stanton
1966). Seeds contain sterculic acid which may be responsible for this effect
(Berry, 1980b).
Seeds
Mature seed constituents : The moisture content of fresh durian seeds is
enumerated by Chin et al. (1989). The moisture content of the embryo is
given as 65% (w/w) on a fresh weight basis, while the value for the ‘whole
seed’ is only 50%. The authors do not define what is meant by whole seed,
so it is unclear if this refers to the embryo and seed coat, or the embryo,
seed coat, and the large fleshy aril. Berry (1980b) gave measurements of
durian seed moisture as 77%, Soepadmo and Eow (1977) stated that the
average moisture content of fresh seeds is 51%, Chin et al. (1984) listed the
moisture content of freshly harvested seeds as 39%. The large variance in
these estimates is likely due to differences in drying regimes. Estimates of
seed moisture content of durian apparently vary greatly when the drying
time and temperature is varied (Grabe 1992). Fresh durian seeds contain
some small amounts of protein, 1.57% (Berry 1980b), but consist largely of
starch (Table 7). Mature seeds appear to be metabolically active when shed;
polysomes are evident in mature seeds and specific enzymatic activity is
detectable (Brown 1995b).
Oil content is reported to be 0.5% (Berry 1980b). On the lipid

constituents of both the arils and seeds of durian, it was reported that
cyclopropene fatty acids including sterculic, dihydrosterculic and malvalic
acids were present in the uncooked seeds but not in the aril (Berry 1980a,b,c,
1981). Due to the toxic and perhaps carcinogenic nature of these substances,
it would be unwise to ingest uncooked durian seeds. It is of interest that no
indication exists in the literature that durian seeds are ever eaten uncooked
by any native peoples.
Culture of seeds and seed size : The seeds of durian are large. According to
Troup (1921), about 1 dozen seeds weighs 454 g. I think this may be an
overestimation unless the author made his calculations from a variety with
unusually large seeds. From my own measurements, mature durian seeds weigh
approximately 20 grams. Davis and Bhattacharya (1974) give an average
weight of 12 to 15 grams from measurements made on the seeds of two
fruits4. Chin et al. (1984) state that the weight of 1000 durian seeds is 14783
g (14.7 g per seed). There is a great variation in seed size within a single
4
These estimates include the smaller semi-aborted and full sized seeds.
Table 7. Components of durian seeds
Component Amount and source
Moisture (77.0%)1* (51.5, 51.1%)3*

Fat (0.5%)1* (0.23%)2* (0.4, 0.2%)3*
Protein (1.57%)1* (2.6, 1.5%)3*
Total carbohydrates (43.6, 46.2%)3*
Crude fibre (0.71%)2* (-, 0.7%)3*
Nitrogen (0.297%)2*
Ash (1.00%)2* (1.9, 1.0%)3*
Calcium (88.80mg/100g)2* (17, 39mg/100g)3*
Phosphorus (86.65mg/100g)2* (68, 87mg/100g)3*
Iron (0.64mg/100g)2* (1.0, 0.6mg/100g)3*
Sodium (3, - mg/100g)3*
Potassium (962, - mg/100g)3*
Beta carotene equivalents (250, - µg/100g)3*
Riboflavin (0.052mg/100g)2* (0.05, 0.05 mg/100g)3*
Thiamine (0.032mg/100g)2* (-, 0.03mg/100g)3*
Niacin (0.89mg/100g)2* (0.9, 0.9 mg/100g)3*
1
Berry (1980b) Note: data is for fresh durian seeds. 2 Intengan et al. (1955) Note: This data is
for cooked durian seeds. 3 Leung et al. (1972) Note: two values are given, the first is for raw
seeds (minus seed coat) and the second for cooked seeds (minus seed coat). In some instances,
only one of these values is given, the missing value is represented by “-”.
durian fruit, some seeds abort at different stages of development. Full sized,
but flattened and completely empty seed coats may often be found inside the
arils of mature fruits. Fully mature and filled durian seeds sink when immersed
in water, but ‘aborted’ or ‘partly-filled’ seeds float (personal observation).
Partly filled seeds can often be successfully germinated (personal observation).
When mature, the seeds are non-endospermous, the endosperm becoming
exhausted several weeks before fruit abscision. The endosperm remains in a
free nuclear state until late in embryogeny at which time it becomes cellular
(Soepadmo and Eow 1977). According to Ho (1972), there is a great deal of
genetic variability in endosperm abortion in Durio, however, I think that he
is actually referring to seed abortion. The seeds of durian are described and
portrayed diagrammatically by Corner (1976) with particular attention to the
vascular supply, in his encyclopedic ‘Seeds of Dicotyledons’.
It has recently been possible to culture excised durian embryos (Chin

et al. 1988). Wounded seeds exude a mucous like secretion and are highly
susceptible to oxidative browning, thus, excised embryos fared best when first
soaked in antioxidant, and subsequently grown on charcoal containing media,
containing 1.0 mgl-1 NAA (a-napthalene acetic acid) or IAA (indole-3-acetic
acid) and 1.0 mgl-1 kinetin, BAP (benzylaminopurine), or 2iP (2-isopentyladenine)
(Chin et al. 1988). Embryos of D. lowianus cultured on woody plant medium
supplemented with BAP produced callus (Kongnakorn et al. 1985). Apparently,
some success in callus formation from excised cotyledonary tissue of D. zibethinus
has also been achieved (Rao et al. 1982; Johri and Rao 1984), however, as of
yet no literature has been uncovered on plant regeneration from such callus
cultures of durian5.
Viability : It has long been known that the seeds of durian have a short period
of viability (MacMillan 1909; Main 1909a,b; Troup 1921; Chevalier 1934).
The seeds of durian are now categorized as recalcitrant (Hofmann and Steiner
1989); seeds with a relatively high moisture content at maturity, which cannot
withstand desiccation (Hanson 1984; Roberts et al. 1984), and thus have a
relatively short period of viability. In durian, this is correlated with the inability
to withstand chilling or freezing. The recalcitrant nature of durian seeds presents
major problems in the storage of durian seeds both for commercial purposes,
and for conservation. As durian is undoubtedly suffering some degree of
genetic erosion (Sastrapradja 1975), research into the physiological response
of durian seeds to chilling and drying is assuming a new importance.
Seeds stored at 36oC lose viability after 6 days, while surface sterilized
seeds sealed in an airtight container under nitrogen can maintain viability for
32 days if the temperature is lowered to 20oC (Soepadmo and Eow 1977).
Further experiments by Teng (1980) showed that fresh durian seeds, stored
for 3 months on wet tissue paper at 15oC, maintained 79% germinability,
although some problems with fungal infection and radicle protrusion were
encountered. Storage of durian seeds at 5oC for 10 days resulted in a reduction
of viability to 80%. Viability was completely lost at this temperature after 20
days of storage. Seeds stored at 15oC and 29oC maintained their viability for
20 days; however, seeds stored at the higher temperature had all germinated
after 10 days. Hanson (1981) performed some preliminary drying and storage
experiments on some tropical recalcitrant seeds including durian. Durian was
shown to have a viability period at 27oC of 3-4 weeks; seeds stored at 15oC
exhibited only 15% germination after 2 months.
5
Numerous attempts at tissue culturing durian seeds have been attempted by myself and Miss
L. Sreedhar, in the Department of Botany, University of Guelph. Although vigorously growing
callus is somewhat easy to produce, various combinations of plant growth regulators at different
concentrations have proven unsuccessful in regenerating plants.
The critical moisture content of durian seeds as determined by drying

over silica gel was found to be 45-50% and durian seeds lose viability at
about 20% moisture (Hanson 1981). Several drying regimes were carried out
by Teng (1977); 14 days of drying (to a final moisture content of 36%)
reduced the percentage germination of seed from 100 to 70%. Drying for
longer time periods (28 days), or to moisture contents of 23.2% or less,
resulted in the complete loss of seed viability.
The moisture content at viability loss after slow and fast drying was
reported as 71 and 63% respectively (Boyce 1989; Grabe 1989). These values
are much higher than numerous other accounts of the minimal moisture content
of viable durian seeds. The differences between this report and those of others
are undoubtedly related to the exact method of moisture determination which
is known to greatly affect moisture determination in durian seeds (Grabe
1992).
Hor et al. (1989, 1990) investigated the behaviour of durian seeds

and embryos upon drying and freezing in liquid nitrogen. The critical moisture
content (the value below which rapid loss of viability occurs) of durian embryos
was determined to be 51.0% (Hor et al. 1989); 53.9% (Hor et al. 1990) ,
while that of the seed was determined to be much lower, 26%. Unfortunately,
the threshold moisture content of durian embryos (the level below which there
is no freezable water present) was determined to be 32%. Thus, it was not
possible to lower the moisture content enough to permit successful
cryopreservation without the complete loss of viability due to dehydration.
Today, there is still no method for medium to long term storage of durian
seeds (Lin 1992).
Morton (1987) claimed that durian seeds are rendered inviable by

exposure to sunlight. This is most likely based on a statement by Malo and
Martin (1979, 1980a). In their original context, they might have implied that
direct sunlight would damage seeds because it raised their temperature, not
because of a direct effect of light per se. However (Anon. 1982a) also stated,
‘germination is rapid and easily accomplished, but the vitality is short-- a few
weeks or only a few hours if the seed is exposed to the sun.’ Unfortunately
no data were presented in this study. These remarks are in conflict with those
of Main (1909a,b) who stated that before packaging and shipping seeds, it is
best to thoroughly wash them, then dry them in the sun for two or three days
to help ensure viability upon arrival. It is the experience of the author that
durian seeds left in a moist location on the ground, exposed to sunlight in a
durian orchard, will germinate. It also seems unlikely that light would penetrate
the thick opaque seed coat of durian. However, to the best of my knowledge,
formal experiments on the effects of light on the viability of durian seeds
have never been conducted or published.
Germination : MacMillan (1909) states that durian seeds germinate in 7 to

8 days, but no information on percentage germination was given. Durian
seeds germinate very quickly, 95% germination occurring after about 10 days
(Soepadmo and Eow 1977). The complete removal of the fleshy aril from the
seed coat before sowing greatly enhances the rate of germination (95%
germination after 3 days) (Soepadmo and Eow 1977). Ng (1975) reports 100%
germination after 4 weeks, 2 weeks being necessary for 50% germination in
D. zibethinus, and 87% germination after 3 weeks in D. griffithii, 1 week
being necessary for 50% germination. According to Padolina (1931), 17 days
are necessary for germination of D. zibethinus, but no record of percent
germination after this time was given. As much as 94% germination has been
reported for D. graveolens after 3 weeks, 2 weeks for 50% germination;
100% germination for D. lowianus after 3 weeks, 2 weeks for 50% germination;
66 and 91% germination after 6 and 3 weeks respectively for D. oxleyanus,
2 weeks being necessary for 50% germination (Ng and Mat Asri 1979).
The method of germination of D. zibethinus is unusual, and was

described in detail by Rumphius (1741) in his ‘Herbarium Amboinense’. The
first part of the seedling to emerge from the seed is the hypocotyl, which is
thick and square in cross-section. Eventually, roots emerge from the tip of the
hypocotyl, and the shoot emerges from between the petioles of the cotyledons.
Oddly enough, whether germination is epigeal or hypogeal depends on the
orientation of the seed when planted (Singh and Rao 1963). When the seed
is planted with the micropyle oriented downwards, the germination is epigeal;
upwards or horizontal, the germination is hypogeal. This type of germination
has been termed false epigeal. Such a crude delineation of germination types
(epigeal vs. hypogeal) grossly oversimplifies the processes which actually
occur in this and many other species. The much more comprehensive seedling
classification scheme of De Vogel (1980) further subdivides the method of
germination in the genus Durio into two types. D. excelsus and some D.
zibethinus are ascribed to the exquisitely rare Horsfieldia germination type
Pseuduvaria subtype, or Hor7b, which has only been firmly documented in
three other genera of flowering plants. Other species of durian; D. acutifolius,
D. dulcis, D. griffithii, D. kutejensis, D. oxleyanus, and some D. zibethinus are
listed as Blumeodendron type. Meijer (1968) stated (erroneously) that Durio
is phanerocotylar (cotyledons free themselves from the testa during germination).
D. zibethinus is most certainly cryptocotylar, as are apparently all other species
of Durio whose germination has been described. Durio appears on Burtt’s list
of genera with known cryptocotyly (Burtt 1991).
It has been stated that the cotyledons are shed shortly after germination,
yielding no apparent advantage to the seedling (Fenner 1985); however, from

personal experience, this is more true of the tropical jackfruit (Artocarpus
heterophyllus) than for the durian. Soepadmo and Eow (1977) relate that the
cotyledons shrivel and drop off within 38 days following germination, which
is more in line with personal observations. A thorough morphological description
of a newly emerged seedling (with diagrams) is given by Burger Hzn (1972)
and De Vogel (1980).
Pollination Biology
Anthesis : Durian flowers open in mid afternoon (the flowers of some clones
are reported to open as early as 2:30 PM (Tidbury 1976). Even though the
flowers open, the anthers do not dehisce and release pollen until about 7:00
PM. Polprasid (1969) states that durian pollen grains function maximally at
9:30 PM. By early morning, the calyx, corolla and staminal groups have
abscinded leaving just the pistil attached to the receptacle. Jamil (1966) reported
that the timing of anthesis and anther dehiscence varies by several hours
between clones, but no data were presented. Attempts to pollinate durian
flowers with pollen released by crushing from undehisced anthers resultd in
no fruit set (personal observations). Although pollen is not functional before
anther dehiscence, pollen does remain viable up to two days after anthesis
when stored in a refrigerator (Coronel 1966).
Valmayor et al. (1965) and Coronel (1966) reported that the stigmas
of durian are receptive to pollen 36-48 hours before the flowers fully open.
Coronel (1966) reports that hand pollination of 36-48 hours before anthesis
flowers by surgical removal of the outer floral parts leaving just the pistil,
results in a greater percent fruit set than hand pollination of fully mature
flowers. Razak et al. (1992) have shown 87.5% receptivity of D24 durian
stigmas 10 hours before anther dehiscence with 42.1% receptivity remaining
24 hours after anthesis (receptivity was estimated by measuring percentage
fruit set). Salakpetch et al. (1992) found that stigma receptivity in some Thai
cultivars began 24 hours pre-anthesis, receptivity remaining high until noon
the day after anthesis, after which it rapidly declined. Shaari et al. (1985)
reported that receptivity was reduced to approximately 50% by noon the day
after anthesis. Jamil (1966) has suggested that the period of stigma receptivity
varies considerably between clones. Salakpetch et al. (1992) showed that
pollen viability also varies between clones in a study with 4 Thai cultivars,
viability ranged from 77 to 93% two days after anthesis.
Natural pollinators : Today, durian trees are generally regarded as bat pollinated,
although Beccari (1889) was of the opinion that birds could be pollinators.
The first suggestion of bat pollination of Durio is that of Beccari in his book
‘Malesia’ (1889) who records Macroglossus minimus Geoffr. visiting the flowers.
This account apparently did not make a big impression for it was not until
over 40 years later, that Boedijn and Danser (1929) noted a flock of bats
visiting a durian tree, their activity causing the shedding of numerous stamens
and calyxes. Although they didn’t observe or demonstrate actual pollination
of flowers by bats, they postulated that this may have been the case. Interestingly
enough, the affinity of flying foxes for the nectar of durian flowers must have
been common knowledge, an account of their feeding is given in an illustrated
tour guide to the Federated Malay States first published in 1910 (Harrison
1910a). Van Der Pijl (1936) conclusively demonstrated that durian flowers
have typical bat pollinated flower morphology, and that the visitation of flowers
by bats does indeed cause fruit set; he also raised strong arguments against
birds as possible pollinators. As birds have not been shown to visit durian
flowers during the time when pollen has been released, they are unlikely
natural pollinating agents.
Baker (1969) witnessed pollination by an unnamed species of bat and

by an unnamed species of hawkmoth. However, this was on a single specimen
growing in the Honduras, far from its native home. Van Der Pijl (1941) has
recorded bats visiting durian flowers, in the case of Durio zeylanicus6 he
notes that the flowers are also squashed and entirely eaten by bats.
Soepadmo and Eow (1977) documented the visitation of Durio zibethinus

flowers by three species of bats (Eonycteris spelaea, Cynopterus brachyotis,
and Pteropus vampyrus). Their observations were also supported by analysis
of bat guano for pollen grains. Start and Marshall (1977) calculated that
pollen of D. zibethinus and D. graveolens makes up 3.4% of all the pollen
grains extracted from samples of bat guano taken from Batu Caves in Kuala
Lumpur, which are inhabited by Eonycteris spelaea. Soepadmo (1979) reported
bat pollination of Durio zibethinus, D.graveolens, D. malaccensis and D.
oxleyanus by Eonycteris spelaea, Macroglossus maximus and M. minimus.
Whitmore (1990) states that the flowers of the related Kostermansia malayana
are also bat pollinated.
Although bats definitely pollinate durian trees, flying foxes (Pteropus

vampyrus) have often been thought to be responsible for destroying many
young flowers (Anon. 1953a; Browne 1955). In fact, in the early part of this
century, it was recommended that ‘...if you have a gun you are hereby requested
to shoot as many of him [Pteropus vampyrus] as possible, for he is a bitter
6
He may be referring to Durio ceylanicus Gardn. which has now become in part Cullenia
ceylanica (Gardn.) K. Schum. (see Table 1).
curse...’ (Harrison 1910a). Gould (1977), in his study of foraging behaviour,

suggested that the damage to durian flowers by flying foxes is negligible, and
rather than a nuisance, they are important pollinators. Gould (1978) investigated
the foraging behaviour of bats on durian flowers with reference to Parkia and
Musa flowers, and proposes that access to nectar is restricted by the shape of
the flower, which encourages sporadic foraging in some bats. This presumably
favours outcrossing, while favouring territorial behaviour in flying foxes.
Soepadmo and Eow (1977) via several pollination experiments, estimated that
about 45% of successful pollination was contributed to by natural pollinators.
Mardan and Zainal (1986) studied the effects of excluding bats as
pollinators of durian flowers by the use of wire mesh cages. Bat excluded
treatments yielded significantly lower fruit set, suggesting bats were responsible
for 39% of the total number of successful pollinations in the study. No significant
effect on the abortion rate of developing fruits was found due to bat exclusion.
Furthermore, these authors report visitation to flowers by the bee species Apis
dorsata, Apis cerana and stingless bees of the genus Trigona. Only A. dorsata
was seen to forage for nectar in flowers at night. Ferrazzi (1995) also claims
that the bee species A. dorsata, A. cerana and A. melifera pollinate durian
flowers in Thailand.
To some extent, nocturnal moths (Soepadmo and Eow 1977) and
other insects (Jamil 1966) may also be natural pollinators of durian, although
this is not well documented. Jamil (1966) reported that at least 10 insect
species visit durian flowers, and probably affect pollination. Unfortunately,
there is no record as to the time of these visitations (if they occur during the
day, they may well precede anther dehiscence), or of the species involved.
Early ovary abscission vs premature fruit drop : The efficiency of fruit-

set of durian is quite low. Only 5.4% of hand pollinated flowers of 3 durian
trees (clone D8), cross-pollinated with pollen from D24, produced fruits
which persisted on the tree until maturity (personal observations). This is
very close to the result of 5% reported by Soepadmo and Eow (1977) after
artificial pollination experiments. Mardan and Zainal (1986) reported that
bat exclusion had a significant effect on fruit set but that the percentage of
mature fruits finally produced compared to the number of flowers produced
was less than 3 percent, regardless of exclusion of bat pollinators. Namuco
(1978) reported that ovary abscision was high 4 days after anthesis, especially
in self-crosses of a known self-incompatible tree; all self-crosses had abscinded
by the tenth day. Premature fruit abscission, however, persisted up to the
9th week of development in cross-fertilized fruits. Similarly, Shaari et al.
(1985) (data reprinted in Razak et al. 1992) showed that the ovaries of
unpollinated flowers of clone D24 had all abscinded by two weeks after
pollination. Self crosses of clone D24, however, resulted in initial ovary

growth, but all fruits abscinded by the 4th week after pollination (Shaari et
al. 1985). These studies imply two things. Firstly, that premature fruit drop
(which occurs later in development) is an independent process to early ovary
abscission caused by selfing, or lack of pollination. Secondly, they also
suggest that self-incompatibility is probably not due to the inability of pollen
to germinate or pollen tube abortion. If this were the case, it would not be
expected that ovaries of selfed flowers would stay on the tree significantly
longer (2 weeks) than those of unfertilized flowers, which was the case in
the study of Shaari et al. (1985).
Self-incompatibility : It has been known for some time that some durian
trees are self-incompatible (Coronel 1966); although crossing and selfing
tests with named clones have been conducted (Jamil 1965), no details
seem to have been published. This is unfortunate as such information
would be useful in determining good clonal mixtures for orchards and for
the production of better clonal material in the future. The Malaysian clone
D99 has been shown to be self-compatible (Zainal Abidin and Nik Masdek
1992). Clone D24 (Shaari et al. 1985), and the Thai cultivars Gumpun and
Luang (Lim el al. 1992) have been demonstrated to be self-incompatible.
However, compatibility information for the many other clones/cultivars is
not available.
The hermaphroditic flowers of the wild species D. griffithii have

been shown to be self-incompatible (Ha et al. 1988). However, Tinggal
(1993) states that the uniformity in fruits from different individuals of the
wild species D. kutejensis suggests that the species is probably very
homogeneous and self-compatible. As for D. zibethinus, there are several
possible physiological explanations that can be put forward to explain self-
incompatibility.
Heterostyly : Chin and Phoon (1982) claimed that there is heterostyly between
different durian clones, although they presented no evidence for this statement.
My personal observations (unpublished) of the stylar lengths of different durian
clones show a lack of distinct size groupings between clones. Lye (1980)
described several differences in stylar morphology that apparently exist between
clones, but he did not mention stylar length. The study of Shaari et al. (1985),
as mentioned previously, showed that unfertilized ovaries abscinded much
earlier than did those of selfed flowers of a self-incompatible clone. Heterostyly
would not explain this observation. Personal observations, and the work of
Lye (1980) have shown that there is definite heteromorphy in stigma and
styles between clones. Despite these differences in stylar morphology, selfed
pollen has been shown to germinate on the stigmas of selfed flowers (Namuco
1978). Furthermore, pollen grains collected from fallen staminal phalanges
will germinate after storage at room temperature for 48 hours, thus stigmatic
exudate is not strictly necessary for germination. Thus, it seems unlikely that
stigmatic heteromorphy contributes to or is associated with self-incompatibility
mechanisms.
Mechanisms of self-incompatibility : There are two other schools of thought

regarding the mechanism of self-incompatibility in durian. Firstly, Shaari et
al. (1985) stated that incompatibility is due to pollen tube arrest at the base
of the stigma; however, their claim is based on a previous research paper that
could not be located for examination. Furthermore, their own data shows
quite clearly that selfed ovaries abscind more slowly than do those of unfertilized
flowers which suggests that at least the early stages of seed development are
occurring in selfed ovaries. The only supporting evidence that I am aware of
for the pollen tube arrest hypothesis is the observation by Soepadmo and Eow
(1977) that the length of pollen tubes produced in vitro is dependent on
sucrose concentration in the medium. While an incompatibility mechanism
based on differential pollen tube growth due to varying levels of available
sucrose is conceivable, it would not explain the results of Shaari et al. (1985).
Empirical evidence : Namuco (1978) showed that self-crossed pollen from

a known self-incompatible tree actually germinated on the stigma, and that
fertilization and initial endosperm development occurred. Unfortunately, the
growth of the pollen tube through the style and actual syngamy was not
observed in Namuco’s study, which would have added further weight to his
conclusions. However, his observation of zygotes and initial endosperm
development is strong circumstantial evidence that these processes did occur,
especially since there is no evidence that durians can produce seeds
parthenogenically. Thus, a second hypothesis regarding self-incompatibility
invokes early endosperm or embryo abortion as a mechanism rather than
pollen tube arrest. Namuco’s conclusions also support and explain the data of
Shaari et al. (1985). Namuco (1978) showed that seed development started in
the selfed flowers of an unnamed durian clone, but all selfed ovaries had
abscinded by 10 days after pollination. Lim et al. (1992) reported that selfed
fruits of two Thai cultivars (Gumpun and Luang) aborted within 3 weeks,
unfertilized flowers, however, aborted after only 2 days. Thus, several independent
studies show a timing difference between the abscision of unfertilized ovaries
and selfed ovaries, which does suggest a difference between the two events;
further, the exact timing of these events may vary between clones.
If durians have evolved under selective pressures favouring outcrossing,

it is possible that several mechanisms of self incompatibility are exploited,

perhaps even in addition to the two mechanisms suggested previously. Different
mechanisms may function to different extents in various clones. This is somewhat
supported by the varied times taken for fruit abortion using trees of different
clonal origins as mentioned above. Further, Lim et al. (1992) state that, while
selfings of the Thai cultivars Gumpun and Luang resulted in abscision within
three weeks, selfings of the cultivar Kob resulted in mishapened fruit. Thus,
there appears to be degrees of compatibility between clones rather than a strict
compatibility/incompatibility relationship. Again, this is easier to explain by
what must be a continuously varying trait such as the timing of embryo or
endosperm abortion than by a simple two state phenomenon such as pollen tube
arrest.
As previously mentioned, Jamil (1966) reported differences in the

timing of anther dehiscence between clones. Ridley (1922) even reported
encountering specimens of D. zibethinus upon which only male flowers were
produced, the female parts being aborted; although this is definitely not the
case with clones such as D24 as discussed above. Much research (Valmayor
et al. 1965; Coronel 1966; Salakpetch et al. 1992) indicates that the stigmatic
surface is receptive to pollen long before the anthers dehisce, thus stigmas would
be open to pollination on the day of anthesis before their own pollen was released.
Together, these observations suggest that varied mechanisms to promote outcrossing,
and prevent self-fertilization are exploited in durian. Thus, although self-
incompatibility in one durian tree (clone unnamed) reported by Namuco (1978)
was likely due to either embryo or endosperm developmental arrest, other
mechanisms may also be at work. Durian reproduction may be even more
complex than can be explained by a simple compatible/incompatible system;
some research has hinted that crosses using pollen from different clones result
in different percentage fruit drops (Jamil 1965; Polprasid 1969). More research
on the mechanism(s) of self and inter-compatibility in durian is needed. This
is likely an area that will provide interesting and rewarding results.
Premature fruit drop : As mentioned above, premature fruit drop occurs after
all abscissions due to a lack of fertilization or self-incompatibility. Yaacob et
al. (1978) suggested that it may be due to some physiological disorder within
the aborted fruits; however, pathology may not be necessary to explain this
phenomenon. The number of harvestable fruits is not necessarily correlated with
the number of successful pollinations above some basal level. As previously
mentioned, the fruit set of durian flowers is quite low (Soepadmo and Eow 1977;
Mardan and Zainal 1986). Other authors have reported very high percentage
fruit set using artificial pollination (Valmayor et al. 1965; Coronel 1966). The
discrepancy between these high values and the low values previously described
probably lies in the number and distribution of flowers pollinated. Durian trees
produce large number of flowers, but owing to the large size of a mature durian
fruit, it is undoubtedly impossible for all flowers produced to develop into mature
fruits, even if the fruits are all healthy. Polprasid (1969) noted that trees with
more flowers had a lower percentage fruit set. Thus, even if all flowers on a
tree are suitably pollinated, the tree must, and apparently does, have some
mechanism to selectively abort some of its excess fruits. The study of Chandraparnik
et al. (1992) showed that trees treated with paclobutrazol and thiourea had a
more even distribution of flowers throughout the canopy, and had a larger
number of developing fruits at 5 weeks after pollination as compared to control
trees.
There is likely an upper limit to the number of mature fruits which

can be produced, this number being much lower than the total number of
flowers produced. Thus, artificial pollination above this limit is not useful.
Future studies on percent fruit set of durian and the effect of the distribution
of these fruits on the tree are necessary to define the limits of fruit set; this
is likely to vary between clones. Clones with highly desirable fruits are
frequently very poor bearers (Hassan n.d.). The possibility that this situation
might be even slightly ameliorated by chemical spray at appropriate times,
or by strategic hand pollination with pollen from particular clones remains
open to investigation.
Manipulating premature fruit drop : The biochemical mechanism(s) behind

the abortion of immature fruit is/are unknown, but may involve gibberellins
(GA) (Mamat and Wahab 1990, 1992). Application of GA (largely GA3 with
traces of GA1, GA4 and GA7) to fruit stalks of 6 weeks old fruits reduced
percentage fruit drop, endogenous GA levels (measured as GA3 equivalents)
of durians have also been shown to decline during fruit maturation (Mamat
and Wahab 1992). Application of GA to trees has been shown to inhibit early
flowering. The use of paclobutrazol (a suppressor of GA production) increases
the number of inflorescences per branch, the percent branches that are flowering,
and the overall number of flowers per tree (Chandraparnik et al. 1992).
Leaf flushing : Durian trees typically produce a flush of new leaves 3-8
weeks after anthesis (Salakpetch et al. 1992). Leaf flushing in durian is
thought to contribute to premature fruit abscision and hence decrease percentage
fruit set. To test the validity of this assumption, and the capacity of plant
growth regulators to delay the onset of leaf flushing, foliar sprays of several
substances were tested on mature durian trees (Punnachit et al. 1992). 2500
ppm mepiquat chloride (300 g/20 l), KNO3 and 2500 ppm daminozide, and
(500 g/20 l) low nitrogen (0-52-34) foliar fertilizer were found to be effective
in delaying flushing, while GA3 and Multigoldâ (a foliar fertilizer) stimulated

flushing. KNO3 treatment resulted in much higher retention of immature
fruits on the trees until maturity, and an overall increase in the average
weight of mature fruits. Additionally, an increase in the flesh to fruit weight
ratio and an increase in the number of aborted seeds over control trees was
found using this treatment. Cultarâ (paclobutrazol) has previously been shown
to reduce leaf flushing in durian (Kittichontawat 1988). Kittichontawat (1988)
demonstrated that although paclobutrazol did not affect fruit set, soluble
solids and the percentage of fruit set or seeds and the overall fruit weight
increased by such a treatment.
Salakpetch et al. (1992), in an experiment to probe source-sink

relationships, sprayed mature trees with 5 different treatments (0-52-34 fertilizer,
glucose-humic acid, 250 ppm Cultarâ , 500 ppm Cultarâ and controls). All the
chemical treatments caused an increase in the number of harvestable fruits per
tree over the controls. The glucose-humic acid treatment, chosen to increase
the size of the source of nutrients, caused a 54% increase in harvestable fruits.
The low nitrogen fertilizer which was meant to suppress leaf flushing, and
thus reduce competition for nutrients, did increase the number of harvestable
fruits, however, a higher number of deformed fruits were produced, and thus
marketable value was not increased. Treatments with Cultarâ , a GA inhibitor
which retards foliage and fruit growth, were intended to prolong development
and hence reduce competition for nutrients. Treatments at both concentrations
resulted in an increased harvest and marketable yield. Voon et al. (1992)
reported the highest increase in harvestable fruits over control treatments in
Chanee durian from foliar sprays of 1000 ppm Cultarâ in the early season.
Application of Cultarâ in the late (normal) season produced only a minor
increase in yield (at 250 ppm), and actually decreased yield of harvestable
fruits at higher concentrations.
Species which feed upon durians : Some researchers have provided information
regarding the species of animals which naturally feed upon durian fruits.
Hubback (1941) reported that elephants trample the fruits and then eat the
orangutans fed on durians (Wallace 1869; Davenport 1967; Shetford 1985).
Rijksen (1978) records that orangutans also eat the fruit of Durio oxleyanus,
which constitutes an ‘esteemed’ food item of the orangutans of Ketambe.
Orangutans have developed behaviours to aid in the plucking, carrying and
opening of durian fruits (Rijksen 1978). Corner (1964) enumerates rather
graphically several species of animals which feed upon durian. According to
Barrett (1912), wild pigs eat durian fruits. A species of Philippine flying
squirrel (Sciuropterus mindanensis) is known to feed on the fruits of D. zibethinus
(Rabor 1939). Gardner (1847) recorded that monkeys eat the seeds of Durio
ceylanicus Gardn. [=Cullenia ceylanica (Gardn.) K. Schum. sensu Kostermans].
Kostermans (1953b) states that most of the fruits of wild species of durians
are lost by damage from squirrels, monkeys, tupais and hornbills. Hawkins
(1986) claims that durians are eaten by elephants, tigers, deer, rhinoceros and
monkeys. Watson (1984) claims the fruit is even attractive to the domestic
cat.
Natural dispersal of seeds : Ridley (1894) speculated on the dispersal of
several durian species by bears, birds (especially hornbills) and even turtles.
D. zibethinus is perhaps dispersed by bears. The wild bear, Helarctos malayanus,
is known not only to eat fallen fruits, but even to climb trees to obtain durians
(Ridley 1894). Ridley further speculated that D. oblongus seeds are dispersed
by birds, as it is native to Singapore where there are no bears. Ridley related
that upon feeding a fruit of this species to a wild bear (Helarctos malayanus),
it ate the arils with great gusto but refused to eat the seeds. An unnamed
Durio species [Ridley described it as having small red arils=D. graveolens?]
is probably dispersed by hornbills. This sentiment is echoed by Browne (1955):
‘I well remember being seen out of a durian grove by a tiger.’ Whitmore
(1990) said: ‘Tigers are notorious for their passion for durians’. Rutten (1939)
reported the germination of durian seeds in elephant faeces, but none became
established.
Meijer (1968) noted that the fruits of the cauliflorous D. testudinarum,
which bears fruits at the base of the tree, are widely believed by natives to be
eaten by tortoises. Ridley (1894) recorded that this species is known in Borneo
as the tortoise durian.
The seeds of D. oxleyanus are severely damaged by orangutans in
attempts to open the fruit (Rijksen 1978). Rijksen (1978) suggests that the
sun-bear and tigers are probably the main dispersers of seeds of this species.
Fruiting Seasons
Maturation of buds : The floral buds take up to 5 months to become mature
fruits. In my experience, mature fruits of clones D8 and D24 require 95-110
DAP (days after pollination) to mature. D99 requires only 95-100 days (Zainal
Abidin and Nik Masdek 1992). Mon Thong durians are harvested 120-130
DAP (Chattavongsin and Siriphanich 1990a). Durians typically have two seasons
during the year, a major and a minor one. The same trees do not necessarily
bear fruits in both seasons (Browne 1955).
Several authors have recorded, for various locations, the months in
which durian trees typically flower and fruit (Table 8). The maturation of durian
fruits is traditionally indicative of the end of the dry season (Dove 1985).
Floral buds are produced well in advance of flowering and are usually
dormant for at least one year, the breaking of bud dormancy is facilitated by
cool nights (Ong and Lee 1981). Some floral buds do, however, develop from
initiation to anthesis without interruption by a dormant state (Subhadrabandhu
et al. 1991). Occasionally, floral initials can revert back to vegetative growth
(Subhadrabandhu et al. 1991).
D. griffithii has synchronized flowering, although some individuals
produced small numbers of flowers asynchronously (Yap 1980).
Table 8. Durian seasons in different geographic areas
Country/province Flowering/fruiting season
Malaysia
Sabah Fruiting August to December, peak in October (Davenport
1967)
Sarawak Main fruiting season January-March (Mohamad Idris 1987)
Peninsular Main fruiting season June-August (Mohamad Idris 1987)
Peninsular Two fruiting seasons June-July, August (Anon. 1982b)
Penang Fruiting June-August (Low 1836)
Jambu Rias, Pahang Flowering in April (major) and August (minor), fruiting
August and February. The major fruiting season harvest
in one year was 266.6 kg/hectare in February and 2269
kg/hectare in August (Chiow 1976).
Philippines
Mindanao and Sulu Flowering May-June, fruiting August-November (Rodrigo
1968; Galang 1955)
Los Banos Flowering February, fruiting July to August (Namuco 1988)
Davao City Flowering January to May, fruiting May-September (Pascua
and Cantila 1991)
Indonesia Flowering June-September, fruiting October-February

(Rodrigo 1968)
Cambodia Flowers in January, fruits in May (Chevalier 1935)
Sri Lanka Flowering March-April, fruiting July to August (MacMillan

1909; Parsons 1932b)
(Contd....)
Table 8. Contd.
Thailand Fruiting April-July (Visetbhakdi 1988)

Nonthaburi Peak of the fruiting season is June 10-15 (Chimprabha
1964)
India
Burliar Research Station Fruiting July to September (Naik 1949)
Environmental effects on flowering : Watson (1984) stated that flowering is

not photoperiod or temperature responsive in equatorial regions. However,
the diurnal temperature range 23 months before harvest is the major climatic
stimulus for flower induction (Ong and Lee 1981). A 10-14 days dry period
was found to be necessary for the induction of flowering (Chandraparnik et
al. 1992).
Pascua and Cantila (1991) reported that rainfall has a significant

effect on the flowering of durian in the Philippines. The flowers emerge
during or immediately after the driest months. Salafsky (1994a) notes that
‘trees apparently will not flower without a prolonged period of warm dry days
(or associated cool clear nights)’; in his study on the effect of El niño oscillation
events on rural Indonesian agriculture.
Manipulating seasonality : Durians are only available during short day periods
of the year. Attempts have been made to alter the timing of flowering to
spread out the season and, therefore, make durian fruits more readily available
throughout the year. Zainal Abidin et al. (1986) have suggested several ways
of reducing the effects of seasonality on the price and availability of durians.
They recommend exploiting differences in fruiting seasons that exist in different
regions of Malaysia, as well as some useful characteristics of some Malaysian
clones; for example, the ability of some clones to fruit twice a year and the
use of early mid and late producing clones to extend the season. Previous and
unobtainable work cited in Chandraparnik et al. (1992) has apparently shown
that ethephon, daminozide, NAA (a-napthalene acetic acid), or fertilizer
application are unsuccessful in inducing early flowering in durian; GA3
application has an adverse effect on early flowering. Chandraparnik et al.
(1992) investigated the effect of foliar sprays of paclobutrazol (Cultar ®), a
suppressor of gibberellin (GA) production, on early flowering and fruit production
on the Thai variety Chanee. They found that increasing concentrations of
paclobutrazol positively affected the number of flowers produced per tree.
Flowering of treated trees (using the highest concentration of paclobutrazol)
began up to 43 days before that of control plants. However, the number of
harvestable fruits produced, and average fruit weight was negatively affected
by this treatment, treated trees also took 2 extra weeks to mature fruits.
Thus, even at the highest concentrations of paclobutrazol used, a maximum
of one month advance on mature fruit production over untreated trees was
gained due to the extended developmental time. Chandraparnik et al. (1992)
also investigated the effects of foliar sprays of thiourea on previously
paclobutrazol treated trees. A somewhat linear relationship between the
concentration of thiourea and the number of flowers produced was shown.
Flower density was increased up to 400% over control trees, and the distribution
of inflorescences throughout the tree was found to be more even than in the
control treatment. The average number of fruits per tree at 5 weeks after
anthesis was significantly increased, however, no indication of the effects on
the numbers, size or quality of harvestable fruits was given. Lin (1992) proposed
an irrigation forcing model to delay the production of durian inflorescences.
This model has not been field tested.
Ecology, Origin and Distribution
Durio species are found growing in lowland and hill primary forests in
Malaysia, up to 1000 m, at a density of not more than 3-4 trees per hectare
(Soepadmo and Eow 1977). Durians (species not named) were recorded at a
density of 3.7 trees per 40 ha in Ulu Kelantan forest, Malaysia (Whitmore
1990). Foxworthy (1916) states that 1.07% of the forest trees of the East coast
of Borneo are Durio sp. based on an examination of almost 690 ha. In 1967,
durian made up approximately 1 tree per 2.6 per km2 in Sabah (Davenport
1967). The relative density of D. griffithii Mast. at Bukit Sebelah in Sumatra
is recorded at 4.16 tree/ha (Mukhtar et al. 1990). Durio sp. tree densities of
4.4, 3.0 and 2.2 trees per hectare are recorded at Pasoh Forest reserve, Taman
Negara and Krau games reserve respectively (Soepadmo 1979).
A map of the distribution of durian in Thailand is given by Kishimoto

and Polprasid (1976), and a map of the suitability of different regions of
peninsular Malaysia for the cultivation of durians can also be found (Anon.
1982b).
D. carinatus is an important species in peat swamp-forest of eastern

Malaysia (Whitmore 1988), and Corner (1978) listed it as a fresh water swamp
species. Additionally, Corner (1978) stated that D. singapurensis [=D.
singaporensis] is common in fresh water swamp forests, and D. griffithii and
D. graveolens occur in hillock-forests of this ecosystem. Corner also recorded
the 2 related species Coelostegia griffithii and Kostermansia malayana as
fresh water swamp forest trees; for the former, he estimated a frequency of 2
large trees per hectare while the latter was a dominant species with up to 30
large trees in 2 hectares.
Sutisna and Soeyatman (1985) examined a logged-over peat swamp
forest, five years after logging, in 3 locations in Eastern Sumatra, Indonesia;
Suakandis (Jambi Province), Sei Teban (Riau province), and Sei Lalan (South
Sumatra province). In this study, D. carinatus was found at a density of
100.0 sapling stage individuals/hectare, 9.2 pole stage/hectare, 10.3 tree
stage/hectare at Suakandis, no individuals of this species were recorded at
the other two locations. D. lowii (the authors most likely mean D. lowianus)
possibly was present only at Sei Lalan, measurements of number of individuals
per hectare are 21.43 sapling stage/hectare, 10.29 pole/hectare and 1.86 tree
stage/hectare.
Centre of diversity : Mendoza (1941) concluded that the centre of diversity
of the genus was Borneo. About 20 of the approximately 30 recognized species
are found in Borneo, 15 of which are endemic. There are 11 species which
are found in Peninsular Malaysia and 5 of it are endemic. Only two recognized
species are endemic to Myanmar. Thus, although the genus most probably
originated in Borneo, it does seem to have spread up the Malay peninsula
before all contemporary species had evolved. The centre of diversity of the
genus is not in dispute, however, opinions abound on whether D. zibethinus
is native or introduced to regions outside Borneo. Furthermore, there is some
debate as to whether this species actually exists anywhere in the wild, or
whether it is the descendant of some wild species.
Wild form of Durio zibethinus : Popeno (1920) stated that J.D. Hooker did
not think that the natural distribution of D. zibethinus extended to the Malay
peninsula, and furthermore that Hooker suspected that D. malaccensis might
be the wild form. Although both these statements appear in ‘The Flora of
British India’ by J.D. Hooker, they appear in chapter 26 on the Malvaceae
written by M.T. Masters. Regardless of who made these claims, Kostermans
(1958b) states, without explanation, that ‘Hooker’s suggestion that D. malaccensis
is the wild form of D. zibethinus is, of course, entirely wrong.’ It should be
mentioned, however, that there has been tremendous confusion as to what
exactly constitutes D. malaccensis.
Van Steenis (1949) suggested that a species he discovered in Southern
Sumatra (D. spontaneus Bakh.) is the closest wild ally of D. zibethinus.
Kostermans (1958b), however, does not recognize D. spontaneus Bakh. as a
valid species, rather he includes this specimen under D. lowianus Scort. et
King. Although this species bears fruits similar to D. zibethinus, Kostermans
does not speculate on a close affinity between the two species; rather, he
opines that his newly erected species D. wyatt-smithii Kosterm. is perhaps the
wild ancestor of the cultivated durian (Kostermans 1958b). Soegeng-
Reksodihardjo (1962) states that the ‘wild form’ of D. zibethinus need not be
one of the extant wild species.
Some debate exists over the question of whether D. zibethinus is

native or introduced to the Malay peninsula. Both Masters (1874a) and Ridley
(1922) thought it unlikely that this species was actually native to the peninsula.
As stated previously, the centre of diversity of the genus Durio is definitely
Borneo. However, many wild species of Durio are, in fact, found on the
Malay peninsula, some of them exclusively.
There has also been some debate as to whether durians exist naturally
in the Philippines, or have been introduced. Wester and Barrett (1912), and
Wester (1916b, 1921) recorded D. zibethinus as confined to Mindanao and the
Sulu archipelago in the Philippines. Merrill (1926) listed Durio as a genus
represented only by introduced species in the Philippines, and Van Steenis
(1933) listed Durio as a non-native of the Philippines. Mendoza (1941)
thoroughly examined this issue, and claimed that D. zibethinus occurs naturally
on Mindanao and the Sulu archipelago as well as Palawan. Mendoza further
concluded that durian occurs only as an introduced species in the Visayas and
Luzon. Additionally, Mendoza recorded the discovery of a wild species (D.
testudinarum) growing naturally on the island of Palawan, further strengthening
his claim of endemism. Mendoza (1941) hypothesized that durians became
indigenous due to a land bridge that connected Palawan with Borneo during the
pleistocene era. Vendivil and Reynoso (1983) recently reported another wild
species, D. graveolens Becc. growing along a river in a secondary forest in
Palawan.
Durio zibethinus in the wild : Van Steenis (1949) believed that D. zibethinus has
never actually been found in the wild, and Whitmore (1990) regarded D.
zibethinus as completely unknown in the wild. Kostermans (1958b), on the other
hand, stated that D. zibethinus is probably wild in Sumatra and Borneo. This and
the question of its being native or introduced to certain regions share one property
in common, i.e., both questions are undoubtedly impossible to answer. Aboriginal
peoples have aided in the propagation of durian (knowingly or unknowingly) by
eating, trading and discarding the seeds of the fruit, probably for millennia. As
it can never be ascertained whether a tree growing in a forest grows there because
of the actions of these people or not, such arguments are purely semantic, and can
provide little on any useful information or insight.
Attempts at introduction : Durian trees can tolerate temperatures up to 46oC in

parts of Thailand and India (Watson 1984), but growth becomes limited below
22oC, and temperatures below 10oC cause premature leaf abscission (Watson
1984). Although found in Borneo, much of Indonesia, Peninsular Malaysia,
Thailand, Southern Myanmar and a few islands in the Philippines, durians were
never introduced into New Guinea, probably because Malay and Indonesian
traders never settled in New Guinea (Knight 1980). In more recent times, durian
has been grown at the Lowlands Experimental Station in Papua New Guinea
(Bettencourt et al. 1992), and successfully grown in the Solomon Islands at the
Dala Experiment Station (8.5o South latitude) (Anon. 1968). Durian has also been
grown on the Island of Ponape in the Kolonia Botanic Gardens (7o N) (Kanehira
1935). In the Philippines, it is found mostly in Mindanao and the Sulu Archipelago
(5-10o N), but it has fruited in Laguna and Quezon provinces (Galang 1955).
Recently, durian has been introduced and successfully cultivated in Northern
Australia, however, it can only be grown in this country north of 17 o (Watson
1993).
There are two species from Myanmar of Durio, however, D. zibethinus
is probably found there only through introduction. Gamble (1881) states that it
is wild in South Tenasserim, but it is cultivated as far North as Moulmein.
According to Knight (1980), it can only grow in Tenasserim (the southern most
part of Myanmar), but it cannot grow above 16oN in Myanmar. Durian apparently
formed forests in Lower Tenasserim from 14oN southwards (Kurz 1877; Gamble
1972). Further, in the ‘Report on the Settlement Operations in the Amherst
District 1891-1892’, the durian is believed to have been introduced into the
Amherst district of Myanmar from seeds planted from a fruit washed up on the
shore from a wrecked cargo ship in the 1700s (Anon. 1893).
Durian was introduced into Britain as a greenhouse curiosity in 1825
(Anon. 1849), but has never flowered or fruited in Europe. MacMillan (1908)
stated that a few young trees were once grown in Syon House Gardens near
London, and were apparently presented as a gift to Queen Victoria. The 29th
edition of the Official Guide to the Royal Botanic Gardens and Arboretum, Kew
Gardens (1885) lists durian as part of the collection.
According to Soegeng-Reksodihardjo (1962), the first trees to fruit
outside Asia were grown in Dominica from seedlings shipped from Kew Gardens
in 1884. A tree introduced to the botanic gardens at St. Aroment in Dominica (15o
N) grew vigorously and fruited after about 10 years (Anon. 1894).
Durian has been grown in several locations in and about India. Firminger
and Burns (1918) gave the following diagnosis of attempts to grow durian in
Calcutta: ‘they have never risen to more than 1 metre in height, when they have
uniformly died off, the climate of that latitude being quite unsuited to them’.
However, Firminger also recorded that trees grown from seed had reached
heights of 2 metres after three years at the Burliar Experimental Gardens at the
base of Nilgiri mountains at an elevation of 760 m in southern India (11o N). Some
research on vegetative reproduction by grafting has since been carried out at
Burliar (Khan and Sambashiva Rao 1952). Durian has also been introduced into
Port Blair in the Andaman Islands (12o N) (Parkinson 1923).
In Sri Lanka, the Peradeniya Botanic Gardens has/had trees over 40 m

tall (MacMillan 1909). Additionally, Durio ceylanicus Gardn. ‘is found in
Wooded hills near Galle in the Southern Province, but little above the sea level,
and very common in forests in the Central province at an elevation of about 3000
feet (910 m). Flowers in May’—Gardner (1847).7
There have been some attempts to introduce durian into the West
Indies and the Americas. Durian has been planted at the Botanical Gardens in
Trinidad (Pascoe 1882); however, ‘only one plant exists in the St. Clair
Experiment Station...It does not seem readily amenable to the Trinidad conditions
of climate’ (Freeman and Williams 1928). Bailey (1914) stated that durian has
been successfully introduced into Jamaica, however, I have been unable to
locate any other information or confirmation regarding this. Baker (1969)
reported a single mature tree growing in the botanical gardens at Lancetilla,
in the Honduras.
Durian was described as ‘at home’ in the Canal Zone Experiment

Gardens (9o N) in Panama (Allen 1941). Attempts have been made to introduce
durian to southern Florida (Green and Koopman 1978). According to Knight
(1980), the USDA has made four attempts to introduce durian into southern
Florida, all of which failed. It is not just the temperature extremes of southern
Florida that prevent its introduction, but the peculiar [coralline] high pH
limestone soils are not suitable for its cultivation (Lee 1985). An interesting
exception is an account of frost damage suffered by various ultra-tropical
trees in Florida during the 1989 winter (Whitman 1990) (I mention this
study since information on the effects of freezing on durian trees is very
scarce). In December 1989, the night temperature dropped to 0oC on two
consecutive nights. A single specimen of D. graveolens, covered with 63
shade cloth, survived the frost and lost only 45% of its foliage. A very lucky
tree indeed.
A few trees are found in Hawaii, and many bearing trees in Zanzibar
(Morton 1987). Durian is mentioned in Neal’s ‘In Gardens of Hawaii’ (1965)
and a small germplasm collection exists in Hilo (Bettencourt et al. 1992).
Finally, in this respect, I relate the following amusing anecdote regarding

the introduction of durian to South America: ‘They say people in the South
Pacific Islands will almost kill each other, or will divorce their wives (one at
least) to get a durian; but when I took one from Lancetilla and left it in the
bedroom of a friend, he rushed across the hall that night and said, ‘Come over
7
The species to which Gardner refers is probably what is now termed Cullenia ceylanica
(Gardn.) K. Schum. (see Table 1).
here and help me hunt; there must be a dead rat in my room but I can’t find
it’ — Popenoe (1956).
Clones
Clonal selection and hybridization : To date, well over 100 different Malaysian
clones of durian have been registered (Lim 1990), which are distinguished
mainly by fruit characteristics; probably more than 200 Thai durian cultivars
are recognized (Hiranpradit et al. 1987, 1992) (300 according to Malo and
Martin 1980a,b). Polchart (1952) tabulated 129 local varieties from the Dhonburi
area of Thailand alone. Fruits of the more desirable clones are sought after
and fetch a higher price in the market place (Fig. 3). Polprasid (1981) described
the use of local durian contests in Thailand to make known local cultivars and
maximize accessions of good durian germplasm. Thai durians have been divided
into six groups based on fruit morphology (Hiranpradit et al. 1987); a detailed
analysis of leaf, aril and spine morphology and fruit shape were also provided.
An itemized list of criteria for assessment of varietal desirability has been
established to aid the identification and collection of new clones (Hiranpradit
et al. 1992).
Despite the numerous Thai cultivars that have been identified, only
four (Chanee, Kradum, Mon Thong, Kan Yao) are grown on a commercial
scale (Subhadrabandhu 1993). Clones or varieties of other durian species
have not been produced or characterized. Clonal selection of durian began as
early as 1922 (Hasan and Yaacob 1986). Unfortunately, clones with very high
quality fruits (e.g. D2) are often not high yielding, while those which are
prolific yielders (e.g. D24) have lower quality fruits (Hassan, n.d.). Hassan
(n.d.) has suggested a mixture of clones is good practice in a durian orchard
to compensate for good clones being sparse bearers, and for the fact that
many durian clones are self-incompatible. The following clonal mixture is
suggested: 60% D24, 25% D16, 5% each of D10, D8, D2 (Hassan n.d.).
Kwok-Kong (1974) suggests essentially the same ratio with the substitution of
5% D7 for 5% D10. Two planting systems described by Zainal Abidin (1991b)
make use of a clonal mixture of 50% D24, 30% D99, 20% D98/D114. A row
planting system and some data on the growth and productivity of several
different durian clones grown at the experimental orchard at the Universiti
Pertanian Malaysia are given by Yaacob et al. (1978). Efficiency indices of
several individuals of three clones have been published (Hasan and Yaacob
1986).
Natural hybrids between D. zibethinus and D. graveolens are known

(Soegeng-Reksodihardjo 1962). Soepadmo and Eow (1977) suggested that
red-tinged flowers of D. malaccensis reported by Heaslett (1972) may have

belonged to a hybrid between D. malaccensis (normally white flowered) and
D. lowianus or D. pinangianus (pink or red flowered).
Artificial hybridization of D. zibethinus with several wild species (D.

graveolens, D. oxleyanus, D. kutejensis) has been attempted to improve fruit
quality (Hambali et al. 1989). These authors reported no fruit set in crosses
between D. zibethinus, and D. oxleyanus, D. oxleyanus and D. graveolens o r
between D. zibethinus and D. kutejensis. Interspecific crosses between D.
zibethinus and D. graveolens yielded viable seeds. Subhadrabandhu et al.
(1991) have recommended that attempts be made to hybridize D. zibethinus
with D. acutifolius and D. griffithii as both these wild species flower more
reliably than D. zibethinus.
A durian hybridization program was started in Malaysia in 1967.
Hybrids of 11 clones of D. zibethinus have been made and are being evaluated
(Chan 1992). In 1992, Zainal Abidin et al. reported on the success of the first
commercially-produced clonal hybrid trees which show, among other traits,
improved disease resistance.
Clonal identification : The clonal propagation of durian has become widely
Figure 3. Durian fruits of clone D2 and D24 for sale by the street side in
Kuala Lumpur. Durians of known and desirable clonal origin fetch
a much higher price in the marketplace.
Figure 4. The development of a durian fruit (Clone D8). The ovary undergoes
little expansion in the first month of development.
A) A fruit at 56 DAP (days after pollination), approximately 9 cm
in length. The bases of the peltate scales of the ovary have developed
into spines.
B) A fruit at 63 DAP. The fruit is approximately 10 cm in length
and shows little change from 56 DAP.
C) A fruit at 88 DAP. The fruit is now 19 cm in length and the
spines are more fully developed.
D) A fruit at 95 DAP. The fruit is now 25 cm in length and little
change occurs before the fruit abscinds at approximately 100
DAP.
established; in Malaysia between 1986 and 1989 over 800 000 clonally propagated
durian trees were produced for planting (Ali 1993). This increase has led to
a need to positively identify clonal material as mistakes have expensive and
very long-term consequences. Watson (1993), for example, reported that problems
in the introduction of durian as a crop in Australia were due to the importation
of misidentified clones.
Durian clones and varieties have been selected mainly for desirable
fruit characteristics. Thus, durian varieties can often be identified by differences
in fruit morphology. As mentioned previously, the shape of the spines has
been described for several Thai varieties (Hiranpradit et al. 1992). Hiranpradit
et al. (1987) claim that fruit shape and spine morphology are useful for
characterizing durians into groups, and both are highly heritable characteristics.
However, given the vast number of clones/varieties now recognized, it seems
rather unlikely that all fruits could be sufficiently identified from the shape of
their spines.
As mentioned above, durian clones have mainly been selected for

desirable fruit characteristics, but consistent differences in the morphology
of the flowers seem to exist between clones. Some of these differences have
been used to distinguish between several clones (Lye 1980). Lye’s study
relied mainly on stylar characteristics (5 different classes of stylar form
were identified) to classify the floral buds of several durian clones. Although
differences in stylar form do exist (e.g. width, degree of crookedness, etc.),
the natural variance within a clone is quite large (personal observation). No
indication of the variance of floral bud characteristics within a clone is
given by Lye (1980). Thus, how easily clones can be distinguished in practice
by floral characteristics remains to be shown. As there are clonal differences
in stigmatic shape and colour, number of filaments per stamen, etc. (personal
observations), the identification of clones by floral characteristics may be
possible.
The use of flower characters seems more promising than spine

morphology as flowers possess more characteristics that can be easily quantified
than do spines. Furthermore, young durian trees will often start to flower
before they are old enough to actually produce fruit (Lye 1980), so flower
morphology may allow earlier identification of clones. Both the aforementioned
methods could be useful in positively identifying fairly mature trees which
are intended for use in the production of budwood, but would not be of use
in identifying or verifying the clonal origin of batches of young grafted trees.
Recently, preliminary work on clonal identification using isozymes

isolated from leaf samples, and separated by starch gel electrophoresis, has
been published (Salma 1993). This preliminary work showed that 5 durian
clones used in the study could be distinguished by this technique; the isozyme
patterns studied were those of acid phosphatase, alkaline phosphatase and
peroxidase.
Techniques such as isozyme assays and perhaps RFLP mapping would

obviously be useful and very powerful tools in the positive identification of
durian clones, and should be further investigated.
Nursery Care and Cultivation
Seeds : Direct sowing of seeds in the field is not recommended due to damage
from rodents (Anon. 1953a). Shaharuddin (1979) has also reported large losses
of germinated seeds due to rodents which eat the epicotyls. Although seedlings
usually cannot survive such injuries, De Vogel recorded an instance of a
seedling of D. zibethinus in which a ring of buds developed from a region
between the cortex and the stele on a seedling whose epicotyl (including the
top of the hypocotyl) had been bitten off (De Vogel 1980). If planted in a
seedbed, Coronel et al. (1983) recommended that seeds be planted 1 cm deep
with a spacing of 4-6 cm between them. Coronel et al. (1983) also recommended
the coating of the seeds with fungicide.
Branch pruning : Some of the earliest literature on this subject recommended

little or no pruning of durian trees (Parsons 1935; Mohamad Idris 1987).
Coleman (1959) stated that, when growing in a jungle, the lower branches of
durian trees are ‘self-pruning’, whereas orchard grown trees are pyramidal in
shape with low branches. Some authors have recommended pruning of young
trees to reduce wind resistance in particularly windy areas (Polprasid 1961b).
The pruning of the lower branches (especially in young budded trees) to
prevent infection of P. palmivora has also been recommended (Lee and Loh
1966; Navaratnam 1966; Tidbury 1976; Namuco 1988). Coronel et al. (1983)
suggested that all branches lower than 2 m from the ground be removed.
Zainal Abidin et al. (1991) have published detailed pruning instructions for
durian trees. Setiadi (1991) gave considerable advice on pruning, shaping and
training durian trees. In Thailand, trees are sometimes topped at 10 m
(Subhadrabandhu et al. 1991). Flower buds and small fruitlets are also
occasionally thinned to leave 1-2 fruits per inflorescence and 50-150 fruits
per tree (Subhadrabandhu et al. 1991).
Root pruning : Very little information on the roots of durian was available
until very recently. ‘The tree is very impatient of disturbance of its roots’,
(Anon. 1935) however, another early report stated that cutting the roots of
durian seedlings with a spade promotes the production of a fibrous root system
reducing loss during transplanting (Feilden and Garner 1936). More recently,
Chong (1985) suggested that the roots of durian should not be pruned to
reduce circling or kinking (as commonly occurs when young trees are grown
in polybags), as the trees are very sensitive to such treatment. This conflict
has very recently been resolved by Ghani (1992a,b). This empirical study
demonstrated that root pruning increased the growth of a fibrous root system,
contributed to increased overall growth and stem enlargement, and increased
survival rate of transplanted trees.
Application of fertilizer : According to Kanapathy (1976), durians do not

require much fertilizer to be successfully cultivated. However, problems associated
with low nutrient levels likely exist. Ding (1988) reported that up to 15% of
durian trees growing in polybags in some nurseries show nutrient deficiency
symptoms. Magnesium, manganese and copper deficiencies are the most common
deficiencies detected in the field (Zawadha et al. 1993).
Chiow (1976) recorded little difference in girth measurements between

manured and non-manured control plots, especially in early tree growth.
However, manured trees of four different clones yielded well over 4 times
as much weight in mature fruit as did controls (Chiow 1976). A manuring
schedule for durian trees is presented by Hassan (n.d.). Watson (1984) stated
that organic manures should actually not be used, as they are conducive to
P. palmivora infection. In the last 15 years, numerous different fertilizer
recommendations have been published for durian, most of them varying
widely in their advice from completely balanced fertilizers to high nitrogen
or high potassium fertilizers. For example, monthly application of 5g 6:6:6
(N:P:K) fertilizer for durian seedlings is recommended by Morton (1987).
Kanapathy (1976) suggested the use of 18:11:5:2.5 for the first 5 years and
13:9:15:3 or 12:6:22:3 afterwards; a fertilizing schedule is also given in this
work. Three applications annually of 15:15:15 until 5 or 6 years of age
followed by a higher potash fertilizer should be used according to Mohamad
Idris (1987). Woller and Idsava (1981) also recommended a balanced fertilizer
for the first few years (1st year 13:13:13 at 0.5 kg/tree, 2-3 years 13:13:13
at 1.5 kg/tree), but then recommend a high phosphorous fertilizer (4-5 years
12:24:12 at 2 kg/tree). In Thailand, fertilizers are added by drip lines; 1:1:1
as flowers develop, 2:1:2 at harvest times and 1:1:1 at 4 months after harvest
(Watson 1984).
Perhaps the most useful fertilizer recommendations come from various

nutrient removal studies. Ng and Thamboo (1967) performed nutrient removal
studies on durian fruits. They provided the following estimate of nutrients
removed from the soil to produce fruits (assuming a yield of 6720 kg/ha): N
Table 9. Nutrient content of D8 durians on a percent dry matter basis

(Jamil 1966)
Plant part Nutrient content

N P K Ca Mg
Pericarp 0.66 0.09 1.70 0.23 0.23
Seed 1.14 0.18 1.17 0.06 0.21
Aril 1.05 0.81 1.13 0.04 0.09
16.1; P 2.72; K 27.9; Ca 1.99; Mg 3.26 (all amounts in kg/ha). Ng and

Thamboo (1967) also provided estimates of the amounts of each of these
nutrients in the seed, aril and pericarp of 4 different clones, on a percent dry-
matter basis. Jamil (1968) reported the following results of nutrient removal
studies on durian (in pounds of nutrient removed per 1000 lb of fruit): N 2.4;
P 0.35; K 4.0; Ca 0.30; Mg 0.47. The nutrient content of durian fruits has also
been measured (Table 9).
Results of nutrient removal studies of entire trees of different ages

grown in an experimental orchard indicate that young durian trees do not
need excessive amounts of fertilizer, their root system is probably not elaborate
enough to take up excess nutrients, if provided (Yaacob 1983). Furthermore,
after trees have become established, nutrient removal data indicated they also
do not require heavy fertilization. Nutrient removal by fruits in the first two
years of bearing was found to be very low (Yaacob 1983). Based on these
results, Yaacob (1983) recommended the use of 2-4 kg/tree of 16:6:22:3 fertilizer
per year.
What all nutrient removal studies have shown is that K is much more
important than N for durians, being the major element removed by the fruits
and by the tree itself, the amount of P being small in comparison. This is
supported by the study of Jamil (1992c) who examined the effects of N, P and
K on young durian trees. Increasing N was found to have no visible effect on
plant form, increased P increased the tree height, while increased K greatly
affected tree form. Thus, of the former cited fertilizer suggestions, those
recommending high potassium formulations are the most valid.
Complete fertilizing schedules (and probably the most empirically

derived) for durian trees from 1 to 10+ years of age are presented by Zainal
Abidin et al. (1991) and Zabedah et al. (1993). Zabedah et al. (1993) also
presented recommendations to overcome magnesium, manganese and copper
deficiency.
Foliar sprays of KNO3 and other substances during fruit development

increased the overall size of fruits, the edible portion (aril) of the fruit, and
seed abortion (Punnachit et al. 1992). These effects were presumably achieved
by the reduction of competition for nutrients by inhibition of leaf flushing
(Punnachit et al. 1992).
According to a study by Jamil (1992a), N, P, K, Cu and Zn occur as

a decreasing percentage of leaf composition as leaves mature, whereas Ca,
Mn and B increase in mature leaves. The study of Anuar et al. (1992) suggested
that leaf nutrients of trees grown in wet zone area do not reflect tree phenology.
Unfortunately, this study suffered from sampling problems and hence the
results are limited in scope; actual data was, in fact, not presented in their
paper.
A very informative investigation was carried out by Lian (1981). In

this study, durian seedlings were grown in sand and selectively deprived of
various nutrients. Although no fertilizer recommendations are given, a description
and figures of leaves exhibiting different nutrient deficiencies are presented,
which is valuable in the diagnosis of possible nutrient deficiencies in durians
growing in poor soils.
Soil conditions : D. zibethinus is apparently suited to a wide variety of soil

types (Hassan n.d.), although peat, soils with poor drainage and very sandy
soils are to be avoided. Durians are suited to lateritic soils (Parsons 1932b)
and have grown well on granite derived soils. Trees do better in less fertile
upland soils than in more fertile marine or alluvial soils (Hassan n.d.; Kanapathy
1976). Trees do best in deep, well drained, loamy soils with a high content
of organic matter (Hassan n.d.; Coronel 1986). Durian is suited to low
country wet zones from sea level up to 460 m (Parsons 1932a). Although
durian is tolerant of poor soils (Anon. 1953a), on stiff clays and poor soils
the trees are stunted and often unproductive (Anon. 1935). D. zibethinus has
been described as having low to moderate tolerance to flooding/waterlogging,
however, durians can tolerate infrequent prolonged flooding (Maas et al.
1979). Durians have a low tolerance of shallow soil (depths greater than 75
cm are recommended (Maas et al. 1979)), a moderate tolerance of drought,
moderate to high tolerance of infertile and acid soils, and may be suitable
for growth on podzols (Butt and Sia 1982). Some notes on the suitability of
durians to several soil types and ecological regimes are presented by Terra
(1952). A ground-water salinity of less than 1000 µmhos/cm is recommended
for durian (Maas et al. 1979).
Durian is suitable for planting on undulated or sloped land with

inclines up to 35o (Hassan n.d.). Maas et al. (1979) stated that durian is
suitable for growth on slopes up to 25o. A study of moisture requirements
of young trees on steep (30 o) and low (19o) slopes demonstrated that trees
on steeper slopes fared less well, even with identical rainfall, to those on
less steep slopes, due to the lower soil moisture reserve (Yaacob 1992).
Poorer growth on steep slopes can be alleviated by mulch and irrigation
(Yaacob 1992).
Water relations : Some information regarding the water relations of durian

orchards in Thailand has been published. The average initial water infiltration
rate of soil in durian orchards from 6 sites is estimated at 813.46 mm/h, the
average constant infiltration rate at 416.67 mm/h (Wittawatchutikul and
Rouysongnern 1982a). Through-fall of water through a closed canopy of durian
trees in Rayong Thailand was measured to be 81.13% (Wittawatchutikul and
Rouysongnern 1982c). Canopy interception of rainfall of a stand of durian
trees at Tapong Nai village in Rayong is given at 47.94% (Wittawatchutikul
and Rouysongnern 1983). Measurements of evapotranspiration as a percentage
of rainfall for 12 continuous months is given for a durian orchard, the average
for the year was 61.66% of rainfall (Wittawatchutikul and Jirasuktaveekul
1992).
Dollah et al. (1993) presented estimates of the daily water requirements

of durian trees of different ages and in three different climatic zones. Mature
durian trees require up to 360 litres of water per day. Mohd. Razi (1993)
measured the rates of leaf and stem growth, and the rate of photosynthesis in
greenhouse grown seedlings of clone D24 subjected to different levels of
water stress. In areas with a pronounced dry season, proper irrigation of
durian orchards is necessary (Subhadrabandhu et al. 1991).
Transplanting : Newly germinated seedlings are most successfully transplanted

before the first set of leaves open (Coronel et al. 1983). Mulching is necessary
or weeds will overcome newly planted seedlings (Mohamad Idris 1987). Seedlings
are often transplanted into black perforated polyethylene bags at 3-4 weeks
(Tidbury 1976). These bagged seedlings are then used as rootstocks for grafting.
Durian trees are very sensitive to transplanting in the field. Survival is negatively
correlated with moisture stress. Some clones are better able to withstand drought
stress than others, durian clone D99, for example, was shown to withstand
moisture stress substantially better than D24 (Masri 1992). Seedlings are normally
planted one month before the start of the rainy season (Subhadrabandhu et al.
1991).
The minimum dimensions of the transplant hole should be a cube

with sides of 75 cm, but larger holes are more advantageous (Hassan n.d.).
The removed soil should be mixed with 12 kg of manure and replaced. The
young trees can be planted one week after the hole has been prepared (Hassan
n.d.). It has been suggested that organic manures may facilitate infection by
P. palmivora (Watson 1984).
A study on the most suitable size material for planting in the field
revealed that budded material of clone D24 with a stem circumference of
3 cm had 66-77% survival after one year. Saplings with smaller stem
circumferences had survival rates as low as 29% (Ghani 1988). Some clones
(D24 in particular) are apparently difficult to establish in orchards (Ghani
1988).
Lee and Loh (1966) recommended a spacing of 12-15 m between

trees to maximize yield. A spacing of 12×12 m is common (Hassan n.d.),
however, such a large spacing reduces potential yield on a per hectare basis:
he recommended 10.5×10.5 m resulting in a density of 86 trees per hectare.
After a few years of fruit-bearing, the trees can be thinned gradually to about
67 trees per hectare to increase yield. According to Hassan (n.d.), the four
most common planting systems for durian are : square, quincunx, triangular
and contour planting. Planting systems are also described by Zainal Abidin et
al. (1991). Planting distances and other technical information for the growing
of durians for budwood is given by Chong and Raziah (1993).
Intercropping : Young durian trees are sensitive to strong sunlight and should,
therefore, be intercropped with other plants to provide shade (Hassan n.d.).
The major concern for intercropping is that the intercrop does not harbour
Phytophthora (Coronel 1986). It is, of course, an additional benefit if the
intercrop is itself economically valuable. Bananas have been recommended
for this purpose (Hassan n.d.; Hashim et al. 1985; Mohamad Idris 1987).
Coronel et al. (1983) stated that newly planted trees are shaded with bananas
for the first three or four years. Pineapples are also used for intercropping
(Coronel et al. 1983). Gliricidia and guava have also been recommended for
intercropping (Mohamad Idris 1987). Osman and Basri (1987) recommended
intercropping with cocoa. Conversely, durian trees are used as shade trees for
young cocoa trees (Anon. 1991a). A planting scheme for intercropping durian,
cocoa and Gliricidia is presented by Jelani et al. (1992). It has been suggested
that intercropping durian with cocoa can lead to increased incidence of serious
pathogens such as Phytophthora and Rhizoctonia. Nawi and Mohd. (1991)
have studied this claim and conclude that it is largely unfounded, any small
increase in outbreaks of disease that may be associated with this intercropping
can be readily controlled by improving crop management practices. Coronel
et al. (1983) recommended that papaya and coconut should not be used to
intercrop durian for reasons of disease control.
Polchart (1952) recommended intercropping with Erythrina which he

claimed serves the dual purpose of increasing the nutrient content of the soil
and aerating the soil. Conversely, durian trees have been tested for use as
shade trees for coffee plantations (Sulaiman and Anuar 1987). The growth
and survival of D. zibethinus in mixtures with seedlings of 5 other tree species
on several soil types is described by Sastrapradja et al. (1982). A study of the
economics of establishing and maintaining a durian orchard is presented by
Kwok-Kong (1974).
Post-harvest Technology
Information regarding the harvesting and post-harvest processing of
durian fruits is of great importance as the two factors that seriously limit the
durian fruit development as a crop are its smell and its short shelf-life; both
of these factors can be controlled or affected by harvest and post-harvest
practices. Most of the post-harvest technology research has originated in Thailand.
Although Thailand has made great strides in this area, much of the published
research is difficult to obtain and is written in the Thai language, which
undoubtedly limits its exploitation elsewhere. Thai research has produced
valuable information in numerous areas, such as harvesting, local transport
and preparation of fruits for market, methods to overcome the difficulties of
export peculiar to this fruit, and detailed examination of the physiology of
ripening. All of this research has provided insight into how durian's two most
limiting properties can be overcome. Experiments in the physiology of fruit
ripening, in particular, suggest methods to manipulate ripening which offer
the prospects of greatly extending the shelf-life and limiting the undesirable
nature of its smell.
Durian fruits generally fall from the trees at night (Teo 1991). In
Malaysia, fruits are normally collected after they fall (Watson 1984; Mohamed
1990) as it is believed that harvesting the fruits before they are mature affects
the flavour (Nanthachai et al. 1994). In Thailand, fruits are detached from the
tree just before maturity (Watson 1984) and then allowed to ripen. A recent
study by Pauziah et al. (1992) demonstrated that Malaysian D24 durians
harvested at 105 and 110 days after anthesis and allowed to ripen had the
same organoleptic properties as mature fallen fruits; however, fruits harvested
too early (100 DAP) did not ripen properly. Hand harvesting immature D24
fruits increased the shelf-life to 9-11 days from that of only 3-4 days for
fallen fruits (Pauziah et al. 1990,1992). Similar conclusions were reached in
a study in the Philippines (Pascua and Cantila 1991). Thus, knowledge of the
ripening process, and the exact time at which to harvest immature fruits of
each particular clone or variety can be of great value in extending the shelf-
life of the fruit, and yet not affect fruit quality.
Grading : Chattavongsin and Siriphanich (1987) studied the anatomy of fruit

pedicels during development of several Thai cultivars of durian. The firmness
of the fruit stem increases with maturity owing to a proliferation of phloem
fibres during late development; hence, pedicel firmness can be used as an
indicator of fruit maturity (Chattavongsin and Siriphanich 1990a,b). Experiments
in estimating fruit maturity with an ‘Effigi’ firmness tester resulted in 85%
accuracy. The degree of firmness (abundance of phloem fibres) was not found
to vary between fruits of different sizes, but may be affected by the age of the
tree, fruits of younger trees having stiffer stems (Chattavongsin and Siriphanich
1990a,b). This fact complicates the use of pedicel firmness measurements as
practical estimates of maturity.
A comprehensive set of quality standards for three Thai cultivars has

been established as well as a classification scheme for fruit shapes within
these cultivars (Hiranpradit et al. 1992). Quality standards are also presented
by Jitjumnong (1988).
Shipping and cold storage of fruit : The earliest experiments on cold

storage of durian fruit are those of Cumming and Hodges (1920). Durians
kept below freezing were found to be good after 4 months, but not up to the
standards of fresh ones. Many of the durians had unfortunately become
impregnated with the brine, which affected the flavour and preservation.
The earliest recorded attempt at the cold storage and shipping of durian
fruits seems to be that of Kopp (1929). A complete English translation of
this French article is available (Anon. 1929). Interestingly enough, Kopp
did not believe that the shipping of chilled durians to Europe held any
commercial promise. Neighbouring items are at risk of being impregnated
by the smell, and ‘at +3°C, the smell remains weak during all the journey,
to return to all its virulence during the maturation which follows the reheating’
(Anon. 1929; Wardlaw 1937).
Mathur and Srivastava (1954) reported that 3.9-5.6oC and 80-90%

RH were the optimum cold storage conditions for durian fruits. Weight loss
and decrease in ascorbic acid content and acidity as well as increase in reducing
sugar content and total soluble solids were also tabulated for various cold
storage regimes.
Bauchau (1972) concluded that freezing of whole fruits is unfeasible,

but that frozen arils keep their organoleptic quality for 3 months when frozen
at -23oC. After longer storage, arils started losing flavour (Anon. 1960). It is
reported that preliminary experiments in freezing durian arils in polyethylene
bags at -22oC succeeded fairly well. Praditdoung (1986) reported that the
shelf-life of intact durian arils (containing seeds) could be extended up to 30
days when wrapped in low density polyethylene and stored at 4oC. Moleeratanond
et al. (1990) demonstrated that the quality of durian arils wrapped in plastic
and kept at 2oC could be maintained for 48 days for Chanee durians, and 30
days for Mon Thong. Numerous physical data (weight loss, CO2 buildup, etc.)
were tabulated for different treatments. Romphophak and Palakul (1990), who
studied cold storage of whole fruits, provided results somewhat contrary to
those just described. Using Chanee durians kept at 5oC, either untreated or
treated with calcium carbide, they found the pericarp of cold stored durians
showed signs of chilling injury after just one week. Soluble solids in cold
stored fruits remained low and did not rise even 3 days after removal to room
temperature. Durian pulp remained firm while in cold storage, and became
soft upon removal to room temperature. Thus, aril softening and increase in
soluble solids are apparently governed by separate processes. More importantly,
they reported that the palatability, scored by a panel of 6 members, was much
lower for fruits stored for even 1 week than for unrefrigerated controls. No
increase in palatability of refrigerated fruits occurred even after returning the
fruits to room temperature for up to 3 days.
Abdullah et al. (1988) stated that durians can be satisfactorily stored

at 10oC and 90% RH for 1-3 weeks. Pawinakan and Hiranpradit (1989) have
also published some work on freezing and storage of durian arils.
Packaging : Some of the problems associated with packaging and exporting

durian in Thailand (costs, containers and pests) have been examined (Anon.
1985). Surface coating of the fruit slowed ripening and helped reduce the
odour (Anon. 1990). Coating material containing gibberellic acid significantly
reduced internal ethylene concentrations in the fruit, improved the colour and
delayed ripening. Tongdee, Suwanagul, Neamprem and Bunruengsri (1990)
demonstrated that the coating of durian fruits with wax preparations extended
the storage life of fruits by delaying over-ripening and reducing odour. They
recommended a 1:4 dilution of SF 320 or SF 7055 coating for optimal storage.
The use of surface coating extends the shelf life of durian to 2 weeks (Tongdee
1992).
The smell of durians can be successfully contained for at least 46

hours by packing them inside a double-walled corrugated cardboard box
which is then shrink-wrapped with PVC film (40 mm thickness) (Swatditat
and Pathomyothin 1979). Paklamjeak et al. (1986) investigated the best
packaging method for intact ripe durian fruits. Corrugated fibreboard boxes
(5.24 mm thick) with an area of ventilation of 2.5% were selected as the
best type of export shipping container (Paklamjeak et al. 1986). Furthermore,
Paklamjeak et al. (1986) showed that the belief that inclusion of a few basil
leaves in the packing container will eliminate or absorb the odour was false,
however, no other consideration was given in their study to smell containment
properties of a desirable container. The design and structural features of
durian export packaging containers is further elaborated by Paklamjeak et
al. (1988, 1989).
Mohamed (1990) investigated the effects of shrink wrapping, sawdust

packing and tying shut of the fruit stored at ambient temperature of 8oC.
Tying the fruit shut to prevent dehiscence (a traditional method) was found
to be completely ineffective at prolonging shelf-life. Shrink wrapping of durian
was found to keep them fresh for approximately 4-5 weeks. It was hypothesized
that shrink wrapping inhibits dehiscence by the 4 mechanisms: (1) maintains
a high internal CO2 level and hence inhibits respiration; (2) greatly slows
moisture loss; (3) mechanically holds the valves together; and (4) inhibits
microbial action due to the high CO2 and low O2 internal atmosphere.
Ripening of fruits : Measurements of CO2 and ethylene production in ripening

durian fruits have revealed that durian is a climacteric fruit (Tongdee et al.
1989a; Booncherm and Siriphanich 1991). The respiratory climacteric peak is
higher for fruits harvested at a more mature stage (Tongdee et al. 1989b) and
occurs later after harvest for more immature harvested fruits. In Chanee durians,
the appearance of a distinct odour indicating that the fruit is ready to eat
occurs one day before the respiratory peak (Tongdee et al. 1989b). The pH
of durian fruit drops as the fruit is left to ripen (Jenie 1978); that is to say that
the total acidity increases. During ripening, the aril softens and starch hydrolysis
gradually occurs, this is accompanied by a rapid increase in soluble sugars
(Ketsa and Pangkool 1994). Aril softening and increase in soluble solids are
thought to be governed by separate processes (see section on shipping and
cold storage of the fruit). As the fruit ripens, the colour of the pulp also
changes. This phenomenon has not been fully explored, however, Ketsa and
Pangkool (1994) suggest it may be due to the synthesis of ß-carotene. The
green colouration of the husk also disappears when the fruits are ripened at
low humidities (Ketsa and Pangkool 1994). Differences in the speed of ripening
between clones may be due to differences in enzyme activity, notably ACC
synthase and ACC oxidase (Siriphanich et al. 1994).
Durian fruits typically split open as they ripen. This splitting is due
to the development of an abscission zone. Very little is known about structure
of the abscission zone, however, the abscission zone cells are known to have
tannin deposits (Sriyook et al. 1994). Low humidity is known to stimulate
dehiscence (Ketsa and Pangkool 1994). Durians loose a lot of moisture after
harvest. In one study, moisture content was shown to decrease by 21% after
9 days of storage at 30oC and 70% relative humidity, although moisture loss
was lower when fruits were stored at higher humidities (Ketsa and Pangkool
1994). The majority of the moisture lost originates from the pericarp, no
significant drop in moisture content of the arils was found even after 5 days
at 75% RH (Ketsa and Pangkool 1994).
Effects of atmosphere : Low oxygen (O2) concentrations affect the ripening

of durian fruits. Fruits (Mon Tong variety) held in air have a typical climacteric
pattern of respiration with an initial carbon dioxide (CO 2) production of 60
ml/kg/hour and a peak of 145 ml/kg/hour (Tongdee and Suwanagul 1989);
ethylene production peaked at a rate of 10 µl/kg/hour. Fruits held at 10% O2
had a peak respiration rate of only 85 ml/kg/hour with a peak ethylene production
of only 3 µl/kg/hour; however, this did not delay or affect ripening. Fruits
held at 2, 5, 7.5% O2 had respiratory rates and ethylene levels that remained
constant over time; the fruits did not ripen. The quality of fruit stored at 7.5%
or less O2 was slightly affected, but remained acceptable. Fruit stored at 2%
O 2 failed to ripen even when returned to fresh air (Tongdee and Suwanagul
1989; Tongdee et al. 1990). Reduction of atmospheric O2 levels (5 to 7.5%)
was shown to cause a reduction in CO2 and ethylene production, and delayed
ripening (Tongdee et al. 1990); ripening resumed upon removal of the fruits
to air, although this rise was non-climacteric. Storage of fruits at 10 or 20%
CO2 caused no change in ripening but the later treatment slightly lowered
internal ethylene levels (Tongdee et al. 1990).
Plant growth regulators : CO2 and ethylene show a parallel increase during
ripening of durian fruits as shown by Tongdee et al. (1990) in their study on
the effect of O2 and CO2 on ripening. Furthermore, CO2 and ethylene production
remain high after the climacteric (Tongdee et al. 1989a). A high ethylene
level at the time of harvest (1.4 ppm) in Chanee durians indicates that ethylene
actually accumulates before fruit abscission, at least in fruits of this variety
(Tongdee et al. 1989a).
In experiments where seeds with arils were separated and isolated

from the pericarp of the fruit, the bulk of respiration and ethylene production
was found to occur in the pericarp (Booncherm and Siriphanich 1991). The
peak of ethylene and respiration in durian arils occurred before that of the
pericarp, and aril ripening still proceeds normally after its removal from the
pericarp (Booncherm and Siriphanich 1991). It is, therefore, possible that the
ripening of the arils stimulates the ripening of the husk in intact fruits (Booncherm
and Siriphanich 1991).
Chanee durian fruit picked at 75% maturity ripened unevenly or failed

to ripen. Fruits that failed to ripen only had very slight increase in internal
ethylene and CO2 concentrations (Tongdee et al. 1989b); the arils of these
fruits remained hard after 8 days. Fruits harvested at 75% maturity, that did
ripen but ripened unevenly, did show a climacteric rise in CO2 and ethylene.
The most commonly harvested stage of Chanee durians is 85% maturity. By
this stage, the ripening process appears to have already commenced (Tongdee
et al. 1989b).
In a somewhat contradictory study, Cheyglinted (1993) reported that

Chanee durian fruits harvested at 75% maturity did undergo a climacteric rise
in ethylene production and respiration even though they either failed to ripen
or ripened abnormally. Normal ripening could be induced in these fruits upon
treatment with 1000 or 2000 ppm ethephon (Cheyglinted 1993). These
experiments with ethephon suggest that the failure of harvested immature
fruits to ripen is due not to lower endogenous ethylene levels per se, but to
a lower sensitivity to ethylene. Immature fruits require a level of ethylene
higher than that found endogenously within the fruit to elicit proper ripening.
This is supported by Ketsa and Pangkool (1994) who stated that ethylene
application promoted the dehiscence of mature durian fruits more than that of
immature ones.
Ketsa and Pankool (1994) showed that internal ethylene concentrations

were higher in durians stored at low humidity, they speculated that this may
be caused by water stress. The increased ethylene may induce cell wall breakdown
in the abscission zones and thus be responsible for fruit dehiscence; ethylene
has been shown to have a greater effect than weight loss on fruit abscission,
and has also been shown to accelerate fruit dehiscence (Sriyook et al. 1994).
The application of GA3 can delay fruit dehiscence, although the mechanism
is unknown (Sriyook et al. 1994). This finding has led to the suggestion that
a GA3 and wax surface coating may be useful in delaying dehiscence during
shipping of fruits (Sriyook et al. 1994).
Post-harvest technology : Several machines have been developed specifically

to aid with the post-harvest problems associated with durian fruit. Most of this
post-harvest technology has been developed in Thailand. For example, a
mechanical durian cleaning machine was described by Jarimopas et al. (1990).
This machine can wash up to 1350 kg of durian per hour.
The accurate grading and sizing of durian fruits presents problems.

Manual sizing of fruits is subject to large errors (43%) (Jarimopas et al.
1992). Recently, a durian sizing machine has been developed which can size
1.35 tonnes of fruit per hour with an error rate of only 17% (Jarimopas et al.
1992).
Jarimopas and Srihawong (1991) have developed a two-cylinder, 24

HP diesel durian transporter suitable for use within orchards. The transporter
is capable of carrying up to 900 kg of durians at a speed of 15.4 km/h. Several
articles written in the Thai language by Jarimopas et al. (1987 1990) describe
a similar vehicle.
In Malaysia, MARDI has developed a hand-held durian opener, known

as the MAAY2 durian opener. Its design and performance are described by
Sukra (1990, 1991) and Ahmad Tarmizi et al. (1991).
Processed food products and their packaging : Because of the extremely

short shelf-life of durian fruits and their seasonality, some investigations have
centred on developing and improving processed durian products. In 1972,
Bauchau presented the results of some experiments with drying durian arils
and improvement of packaging; evaluation and storage of durian cake has
been discussed by Paweenakarn et al. (1992). Long term storage of durian
cake at room temperature is not possible, but cake can be stored at -20oC for
up to 3 years (Paweenakarn et al. 1992). Technical information on the preparation
of durian paste from Chanee durians is presented by Sisawad et al. (1988).
This paste has a shelf life of over six months when stored at room temperature.
A process for the preparation of dried durian flakes is presented by Sinthavali
and Harutaitanasan (1987). Durian flakes apparently retain the ‘good characteristic
smell’ and have a long shelf life (Sinthavali and Harutaitanasan 1987).
Experiments on suitable packaging material for durian powder have

been performed in order to optimize the factors of cost and protection of the
delicate flavour of durian as well as containment of its more objectionable
qualities. The recommended packaging is laminated aluminum foil which renders
a shelf life of at least 6 months (Hanousek 1971) and possibly up to a year
(Huruthaithanasan 1985). Polypropylene bags (0.1 mm thickness) are less
expensive, but are only suitable for short term storage (1.5 months) (Hanousek
1971). Polyethylene is not suitable for packaging as oil from the durian can
penetrate it (Hanousek 1971).
Forestry Aspects
Timber characteristics of Durio species and close relatives : What is commonly

referred to as ‘durian’ in the timber industry includes wood from more than
just D. zibethinus. In fact, it often includes other related genera. For example,
the species listed as ‘durian’ are by Keith (1947): Boschia griffithii Mast.
[=Durio griffithii (Mast.) Bakh. sensu Kostermans 1958b], Durio zibethinus
Murr. and Durio spp. Menon (1959) also listed Coelostegia griffithii Benth.,
D. lowianus Scort. ex King, D. malaccensis Planch. [=D. malaccensis Planch.
ex Mast. sensu Kostermans 1958b], D. oblongus Mast., D. oxleyanus Griff.,
D. testudinarum Becc. var. macrophyllus King [=D. macrophyllus Ridl. sensu
Kostermans 1958b], D. wrayii King [=D. testudinarum Becc. sensu Kostermans
1958b] and Neesia altissima Bl. Martawijaya and Kartasujana (1981) listed D.
carinatus Mast., D. oxleyanus Griff. and D. zibethinus Murr. as species under
the Indonesian timber term ‘durian’. Normally, wood properties listed for
‘durian’ pertain to a mixture of the above and perhaps other species. Some
information on individual species is given.
Boschia griffithii Mast. [=D. griffithii (Mast.) Bakh.] : This wood is of a

yellowish red colour, yields a smooth grain after machining, but is prone to
attack by insects (Howard 1948). According to Keith (1947), the wood shrinks
badly upon drying. The wood has a weight of 553-832 kg/m3 (Ridley 1901,
1903); 30 pounds per cubic foot (air dry) (480 kg/m3) (Keith 1947); and 42
lb per cubic foot (dry) (673 kg/m3) (Howard 1948). Two values of 47 and
51.1 pounds per cubic foot at 15% moisture (753 and 819 kg/m3) are listed
by Burgess (1966). This wood is used for house building and making beams
(Ridley 1901), and sometimes furniture (Keith 1947).
Coelostegia griffithii Mast. : The weight of the wood has been recorded as
713 kg/m3 by Ridley (1901, 1903), and 705 kg/m3 at 15% moisture by Burgess
(1966). The bark of this tree is used to tan nets. The wood is hard, flexible
and durable (Ridley 1901, 1903).
Cullenia excelsa Wight : [=D. ceylanicus Gardn.; = in part C. ceylanica

(Gardn.) K. Schum. sensu Kostermans 1958b] : Numerous measurements on
the strength of this wood are enumerated by Tisseverasinghe (1963). At
12% moisture, the wood weight is 513-625 kg/m3 (Pearson and Brown 1932).
The weight is recorded as 625 kg/m3 by Tisseverasinghe (1963). The physical
and mechanical properties from Pearson and Brown (1932) are: specific
gravity = approximately 0.50; 18.4% moisture in untreated timber; spike
pulling elastic limit = 4849 kg/cm2; spike pulling maximum load = 9133 kg/
cm2; compression perpendicular to the grain = 1838 kg/cm2; and side hardness
= 1724 kg/cm2.
Durio carinatus Mast. : The bark of this species is apparently used for roofing
(Uphof 1968; Usher 1974). An examination of charcoal, charcoal briquets and
alcohol production from different types of wood waste of D. carinatus is
presented by Syachri (1983).
The descriptive measurements from Chu (1969) are: air dry density-
average 630 kg/m2 range 551-700 kg/m3; fibre length - average 1.65 mm,
range 1.55-1.79mm; fibre average maximal tangential diameter ave. 32.63
µm, range 30.45-34.10 µm; fibre average maximal tangential lumen ave. 25.71
µm, range 24.26-28.32 µm; fibre average maximal tangential wall thickness
ave. 7.25 µm, range 5.89-9.08 µm.
Durio dulcis Becc. : An analysis of wood from this species is given by Sudradjat
(1980). The wood is listed as having a specific gravity of 0.73. Cellulose
accounts for 50.9% of the oven dry weight of the wood, 36.7% is accounted
for by lignin, 14.6% pentosan and 1% ash. Cockrell (1942) listed the specific
gravity of the wood of D. conicus Becc. [=D. dulcis Becc. sensu Kostermans
and Reksodihardjo 1958] as 0.60.
Durio kutejensis (Hassk.) Becc. : The only published information on wood of

this species comes from Burgess (1966) who listed the weight of the wood as
599 kg/m3 at 15% moisture.
Durio ?lowianus Scort. ex King : The weight of wood from this species was
listed as 657 kg/m3 air dry (Desch 1941).
Durio malaccensis Planch. ex Mast. : The weight of wood from this species
was listed as 705 kg/m3 air dry (Desch 1941).
Durio ?oblongus Mast. : The weight of wood from this species was listed as
657 kg/m3 air dry (Desch 1941).
Durio oxleyanus Griff. : According to Kumarasamy and Burgess (1956), D.

oxleyanus rated extremely well with respect to resistance to splitting when
nailed. The wood shrunk 3.0% radially and 4.0% tangentially when dried, and
was very susceptible to attack by powder-post beetles (Burgess 1966). The
specific gravity of the wood is 0.57 (Cockrell 1942). ‘Green’ wood at 87%
moisture had a specific gravity of 0.50, while wood dried to 16% moisture
had a specific gravity is 0.53 (Shukla and Rajput 1988).
The weight of the wood was listed at 753 kg/m3 air dry by Desch
(1941) and 755 kg/m3 by Ridley (1901, 1903). A value of 610 kg/m3 at 15%
moisture was listed as the mean weight of 10 specimens by Burgess (1966).
The physio-mechanical properties of this wood presented by Shukla and Rajput

(1988) were: weight at 15% moisture = 610 kg/m2; modulus of rupture = 751
kg/cm2; modulus of elasticity = 119500 kg/cm2; maximum height of drop
impact bending = 161.0 cm; maximum crushing strength in compression parallel
to grain = 398 kg/cm2; fibre stress at elastic limit in compression perpendicular
to grain = 42 kg/cm2; and side hardness = 363 kg.
Durio testudinarum Becc. : The wood of this species is recorded as having an

air dry weight of 692-710 kg/m3 (Desch 1941). According to Burgess (1966),
the weight is 660 kg/m3 at 15% moisture.
Durio zibethinus L. : The wood of this species is a dull red brown colour with
a hard grain, yielding an uneven rough surface. It is liable to warp and unsuitable
for export (Howard 1948). The wood is straight grained and moderately heavy
(Keith 1947). The wood has a specific gravity of 0.42 (Cockrell 1942). The
weight is listed as 481 kg/m3 (air-dried) by Keith (1947); 545 kg/m3 air dry
by Desch (1941) and Howard (1948); 570 kg/m3 at 15% moisture by Burgess
(1966); and 645 kg/m3 by Ridley (1901, 1903). The wood is suitable for
general construction (Keith 1947).
Durability of ‘Durian’ timber : Durian wood is not very durable (Jackson

1957). Durability tests have been conducted on the wood. In burying experiments,
durian wood pieces were all destroyed after 1.5 years, the first pieces destroyed
after only 6 months (Foxworthy and Woolley 1930). The wood of D. zibethinus
is known to be highly susceptible to damage by termites (Martawijaya and
Sumarni 1978) and moderately susceptible to attack by powder-post beetles
(Menon 1957; Anon. 1964). Durian wood is also susceptible to marine borers
(Burgess 1966). Fortunately, durian wood readily absorbs preservatives, which
helps compensate for its natural lack of durability; absorption levels of up to
96 kg/m3 was obtained with a mixture of creosote and diesel oil (Anon. 1964;
Burgess 1966). Durian wood is normally free from defects except for sponginess
of the pith (Thomas 1952, 1979).
Uses of ‘Durian’ timber : Durian wood is used for making clogs in Sarawak
(Foxworthy 1921; Thomas 1952, 1979; Anon. 1964). It is also used for light
construction, cheaper grades of furniture (Thomas 1952, 1979; Anon. 1964)
and temporary construction work (Foxworthy 1909). Durian wood is
recommended for non-impact tool handles (Lim 1988) and cigar boxes
(Martawijaya and Kartasujana 1981). Durian wood makes satisfactory plywood
(Thomas 1952, 1979). Studies of the gluability of durian plywood have been
conducted by Tsai (1975); the wettability of wood veneers of durian with
different adhesives has also been studied (Wang 1975), as has the bonding
Figure 5. The inside of mature fruit of D. zibethinus. Two of the five locules
are visible. The seeds are covered within the fleshy yellow arils
which surround them and fill the locules.
strengths of different adhesives (Wang 1977; Taki 1986). Tensile tests have
been conducted on plywood made from durian (Kitamura et al. 1982).
The properties of wood wool board manufactured from D. zibethinus

have been described (Sulastiningsih et al. 1987). Durian wood has been
investigated for its suitability as a substrate for the growth of oyster mushrooms
(Pleurotus ostreatus) by Suprapti (1987). Out of 11 woods tested, durian was
found to rate second only to Hevea wood in terms of yield of oyster mushrooms
per kilogram dry weight of substrate.
Studies of pitching problems of tropical woods including durian were

examined by Tachibana et al. (1976). Durian wood pulp causes additional
pitching problems after bleaching as large amounts of residual resin in the
fibres cause discolouration.
Properties of ‘Durian’ timber : Durian wood has approximately 100% moisture

when felled. To dry it from green to air dry under cover in Malaysia takes 2
to 2.5 months for 1.25 cm boards and 3.5 to 4 months for 3.8 cm boards
(Thomas 1952, 1979). Durian wood shrinks badly when dried (Foxworthy
1921). The shrinkage of the wood is estimated at 3.0% radially and 4.0%
tangentially (Anon. 1964).
The weight of wood labelled as ‘durian’ varies considerably as it

consists of several species. Some representative published values for ‘durian’
are 577 kg/m3 (Foxworthy 1916); 465-625 kg/m3 (Foxworthy 1921); 481 kg/
m 3 air dry (Keith 1947); and 527-753 kg/m3 air dry (Menon 1959).
The density of wood of different durian species also differs, some

are sinkers (Anon. 1953a). Nevertheless, estimates of the specific gravity of
‘durian’ wood have been published. The air dry specific gravity of durian
is 0.57+/-0.02, the oven-dry specific gravity is 0.63+/-0.02 (Ma and Chen
1981). Numerous physical properties of ‘durian’ at 12% moisture have been
measured by Ma and Chen (1981). They are fibre stress at elastic limit =
789+/-71 kg/cm2, modulus of rupture = 1372+/-61 kg/cm2, modulus of elasticity
= 167 200+/-18 200 kg/cm2, absorbed energy in toughness = 0.82+/-0.06
kg+m/cm2, compression parallel to the grain = 716+/-13 kg/cm2, shear strength
on tangential surface = 118+/-23 kg/cm2, shear strength on radial surface =
113+/-15 kg/cm 2, hardness of end surface = 6.44+/-0.82 kg/mm2, hardness
of tangential surface = 2.07+/-0.27 kg/mm2, hardness of radial surface =
1.91+/-0.23 kg/mm2, at 17.93% moisture cleavage in radial direction = 37+/
-2 kg/cm, and at 18.02% moisture cleavage in tangential direction = 29+/-
2 kg/cm
Wood anatomy : The wood anatomy of Durio has been examined in some
detail by several authors (Bargagli-Petrucci 1904; Moll and Janssonius 1906;
Cockrell 1942; Menon 1959). The wood anatomy of D. lowianus is described
in a Japanese paper (Anon. 1966). According to Menon (1959), there is silica
in the wood parenchyma of B. griffithii Mast. [=D. griffithii (Mast.) Bakh.
sensu Kostermans 1958b] and C. griffithii Benth.; D. grandiflorus is also
siliceous (Burgess 1966). No silica is present in D. oxleyanus, D. testudinarum,
D. zibethinus or Neesia altissima. There are crystals in chambered parenchyma
strands in all ‘durian’ save B. griffithii Mast. [=D. griffithii (Mast.) Bakh.
sensu Kostermans 1958b] (Menon 1959). Vertical traumatic intercellular canals
are present (Metcalfe and Chalk 1957). Mucilage cavities are present in the
pith of Durio, Coelostegia and Boschia (Solereder 1908).
The wood contains diffuse vessels with no definite arrangement. About

10% to less than 50% of the vessels are solitary, the remainder are found in
radial pairs or groups of up to 10 (Menon 1959). There are normally less than
4 vessels per mm2. The vessel members are 300-1200 µm in length (Cockrell
1942). The maximal vessel diameter (tangential) is 220-370 µm (Menon 1959).
Vessels are further characterized by the absence of tyloses and simple perforation
plates (Menon 1959). The vessel walls are 3-9 µm in diameter (Cockrell
1942)
There is no storied parenchyma in Durio, and the fibres have simple

or indistinctly bordered pits, equally numerous in radial and tangential walls
(Metcalfe and Chalke 1957). Furthermore, the fibres are in radial rows, non-
libriform, occasionally semi-libriform, 20-40 µm in diameter, 900-3000 µm
long, the walls are 3-8 µm thick, and pits are restricted to the radial walls
(Cockrell 1942). Chu (1969) described the fibres of D. carinatus Mast; he
listed the average maximal tangential diameter of these fibres as 30.45-34.10
µm, the average maximal tangential lumen as 24.26-28.32 µm and the average
maximal tangential cell wall thickness as 5.89-9.08 µm.
Growth rings are distinct, and are delineated by denser fibrous tissue
at the outer margin (Cockrell 1942). The sapwood of durian is white or pale
yellow-brown, while the heartwood is brown to deep brown (Anon. 1964).
Both uni- and multi-seriate rays are present in the wood. Measurements
of medullary ray elements from several durian species are tabulated by Bargagli-
Petrucci (1904). The xylem rays of Durio are characterised by the presence
of tile cells (rows of narrow upright cells that resemble tile work when viewed
in radial section). Tile cells typically occur in certain members of the Malvales.
These tile cells are grouped into two types. The first type are the Durio type
tile cells, these are about the same height as the procumbent ray initials, as
is typical in the genus Durio and 17 other genera in the Malvales (Manchester
and Miller 1978). Additionally, there exists the Pterospermum type of tile
cell, in which the tile cells are two to several times higher than the procumbent
initial cells. This type of tile cell is found in approximately 30 Malvalceous
genera (Manchester and Miller 1978). The development of Durio type tile
cells has been described by Chattaway (1933). Resin is abundant in procumbent
cells of durian, but rare in tile cells. The tile cells often contain crystals
(Chattaway 1933). Durio type tile cells are found in all members of the
Durioneae except Camptostemon and Maxwellia (Metcalfe and Chalk 1957).
A study of fossilized eocene wood revealed the earliest known existence of
tile cells, these being of the Pterospermum, or an intermediate Pterospermum-
Durio type. No known fossil woods with Durio type tile cells are known,
however, based on the mode of development, Manchester and Miller (1978)
speculated that the Pterospermum type may have arisen from the Durio type.
Forsaith (1915) claimed from a study of anatomical features of the

Malvales that the primitive condition in this order included diffuse parenchyma
and aggregate rays; characters that are only possessed currently by the genus
Durio. Thus, from an anatomical point of view, Forsaith (1915) claimed that
Durio has the most primitive wood structure in the Malvales. After a study of
wood anatomy of the Bombacaceae, Metcalfe and Chalk (1957) concluded
that the Matisieae are more closely related to the Durioneae than is the
Adansonieae.
Major Diseases, Parasitism and Associated Organisms

The major diseases of durian have been extensively covered in an
excellent book by T.K. Lim (1990), which is a valuable source of information
on durian diseases. Attempt is made here to list all recorded pests, and pathogens
and organisms found associated with the tree or its products (fruit and timber),
and all known literature pertaining to them. Such a list is not presented by
Lim (1990), and could be of value to researchers in this area. As many of
these organisms are discussed by Lim (1990), this work has only been listed
as a reference in cases where it is the only published mention of the organisms
relationship with durian.
Bacteria
Enterobacter sp. -stem canker Liu (1977b); Singh (1980) Flavobacterium sp.
-stem canker Liu (1977b); Singh (1980); Pseudomonas sp. -stem canker Liu
(1977b); Singh (1980).
Fungi
Aschersonia -leaf fungus Singh (1980); Turner (1964, 1971); Aschersonia

placentae B. & Br. [see also the telomorphic state Hypocrella raciborskii
Zimm.] -probably beneficial fungus, Lim (1990).
Botryobasidium salmonicolor (Berkeley & Broome)Venkatarayan, Weber (1973);

Botryodiplodia theobromae Pat. [see Lasiodiplodia theobromae (Pat.) Griff.&
Maubl.].
Calonectria rigidiuscula (Berkeley & Broome)Saccardo (the perfect state of

Fusarium decemcellulare) -stem rot, Jamil (1966); Johnston (1960); Singh
(1973, 1980); Weber (1973); Wiltshire (1956a,b).
Capnodium moniliforme Fraser -sooty mould, Singh (1980); Turner (1964,

1971).
Cercospora sp. -leaf spot, Chandrasrikul (1962), Giatgong (1980), Singh (1980),
Williams and Liu (1976),
Chaetomium trilaterale Chiv. -associated with seedling dieback, Singh (1980);

Turner (1971).
Cladosporium fulvum Cooke, Tigvattnanont and Pramual (1990).
Colletotrichum sp. (Hassan n.d.) anthracnose, root rot, Johnston (1960),

Singh (1973, 1980); Subhadrabandhu et al. (1991); Thompson (1939); Tidbury
(1976).
Colletotrichum durionis Koorders, Coronel et al. (1983), Soegeng-

Reksodihardjo (1962); Colletotrichum gloeosporioides (Penz.) Penz. & Sacc.
[the anamorphic state of Glomerella cingulata (Stonem.)Spauld & Schrenk.]
-anthracnose; Alahakoon et al. (1994); Alahakoon and Brown (1994); Lim
(1980); Nik Masdek et al. (1991); Shaji et al. (1993); Colletotrichum zibethinum
(Sacc.) Petrak -leaf damage, Coronel et al. (1983), Soegeng-Reksodihardjo
(1962).
Coriolus vesicolor (L. ex Fr.) Quel., Yamamoto and Hong (1989).
Corticium salmonicolor Berkeley & Broome -Pink disease, Anon. (1967);

Chandrasrikul (1962); Coronel et al. (1983), Giatgong (1980), Nik Masdek et
al. (1991); Shaji et al. (1993); Singh (1973, 1980); Soegeng-Reksodihardjo
(1962); Turner (1966, 1971); Vichirananda (1983); Watson (1984).

Williams and Liu (1976).
Corticium solani (Prill & Delacr.) Bourd. & Galz. -leaf spot and dieback,
Singh (1973, 1980); Turner (1963; 1971).
Corynespora cassiicola (Berk. & Curt.) Wei. - associated with leaf blotch,
Anon. (1967); Singh (1980); Williams and Liu (1976).
Curvularia sp., Ganapathi and Chinnathambi (1993); Curvularia affinis Boedijn

-leaf spot, Singh (1973, 1980); Curvularia eragrostidis (Henn.)J.A. Meyer
[anamorphic state of Cochliobolus eragrostidis (Tsuda & Ueyama) Sivan.] -
fruit rot, Lim (1990).
Diplodia sp. -scion dieback, Singh (1973, 1980); Soegeng-Reksodihardjo (1962);

Thompson (1938, 1939); Diplodia durionis Sacc. & Sydow. -die back, Baker
(1914; 1931); Chandrasrikul (1962); Giatgong (1980); Reinking (1919).
Fomes lignosus [synonym of Rigidoporus lignosus (Klotzsch)Imazeki] -wood

rot, San Juan (1976).
Fusarium sp. -root disease, Singh (1973, 1980); Soegeng-Reksodihardjo (1962);

Thompson (1938); Fusarium solani (Martius)Sacc., Anon. (1980b); Griffiths
and Lim (1966); Jamil (1966); Weber (1973); Fusarium decemcellulare Brick.
[see also the perfect state Calonectria rigidiuscula] -stem rot, Heath (1956)
Note:Nectria ochroleuca and N. haematococca were also found to be associated,
but probably secondarily, Jamil (1966), Johnston (1960); Fusarium oxysporum
Schlecht. ex Fr., Singh (1973; 1980).
Fusicoccum sp. -twig blight, Chandrasrikul (1962), Giatgong (1980).
Ganoderma pseudoferreus (Wakefield) Overeen & Steinmann, Van Overeen

(1925), Weber (1973).
Gloeosporium sp. -leaf spot, Singh (1973, 1980); Gloeosporium zibethinum

Sacc. Anon. (1928), Chigg (1918, 1921); -anthracnose, Chandrasrikul (1962);
Giatgong (1980).
Glomerella cingulata (Stoneman)Spauld & Schrenk. [see also Colletotrichum

gloeosporioides the anamorphic state of this fungus] -leaf anthracnose and
leaf spot, Anon. (1962); Anon. (1967); Jamil (1965); Singh (1973, 1980);
Turner (1964, 1971); Weber (1973); Williams and Liu (1976).
Helminthosporium capense Thumen. [see Spiropes capensis (Thumen.) M.B.

Ellis.].
Homostegia durionis Rac. -black and brown leaf spots, Coronel et al. (1983);
Soegeng-Reksodihardjo (1962); Subhadrabandhu et al. (1991).
Hypocrella raciborskii Zimm. [the telomorphic form of Aschersonia placentae

B. & Br.] -probably beneficial fungus, Lim (1990).
Lasiodiplodia sp., Ganapathi and Chinnathambi (1993); Lasiodiplodia theobromae

(Pat.) Griff.& Maubl. (=Botryodiplodia theobromae Pat.), Johnston (1960);
Liu (1977b); Singh (1973, 1980); Turner (1963, 1971).
Lentinus subnudus Berk., Chipp (1921).
Leptoxyphium sp. -black crusty growths on leaves and twigs, Lim (1989).
Macrophomina phaseoli Maubl., Watson (1984).
Marasmiellus scandens (Mass.) Dennis & Reid -thread blight, Singh (1980),
Turner (1971).
Meliola durionis Hansf. -black mildew, sooty mould, Hansford (1956); Johnston
(1960); Lim (1989); Nik Masdek et al. (1991); Singh (1973, 1980); Turner
(1961, 1971).
Metacapnodium dennisii S.J. Hughes, Hughes (1976).
Mucor sp. -fruit rot, Lim (1990).
Myrothecium verrucaria Ditmar ex Fr. -leaf spot, Singh (1973, 1980).
Nectria sp. -associated with bark rot, Anon. (1954); Nectria haematococca
Berk. & Br. -stem rot, Johnston (1960); Singh (1973, 1980); Nectria ochroleuca
(Schw.) Berk. -stem rot, Johnston (1960); Singh (1973, 1980).
Oidium sp. -powdery mildew, Chandrasrikul (1962); Vichirananda (1983);

Oidium nephelii Hadiwidjaja -powdery mildew, Giatgong (1980)
Perisporium sp. -seedling dieback, Singh (1980); Turner (1964, 1971)
Pestalotia sp. -leaf spot, Chandrasrikul (1962); Giatgong (1980)
Phomopsis sp. -leaf spot, Chandrasrikul (1962); Giatgong (1980); Singh (1973,
1980); Soegeng-Reksodihardjo (1962); Thompson and Lim (1965); Phomopsis
durionis Syd. -leaf spot of seedlings, branch and stem necrosis, Liu (1977b);
Nik Masdek et al. (1991); Singh (1973; 1980); Weber (1973).
Phragmocapnias betle (Sydow & Butler) Theissen & Sydow emend. Reynolds.
-sooty mould of leaves, twigs and fruits., Lim (1989)
Phyllachora makrospora Zimm., Stevens (1921)
Phyllosticta sp. -leaf spot, seedling rim blight, Anon. (1967); Liu (1977b);
Singh (1973; 1980); Thompson (1939); Tidbury (1976); Williams and Liu
(1976); Phyllosticta durionis A. Zimmermann -leaf spot, rim blight, Chandrasrikul
(1962); Coronel et al. (1983); Giatgong (1980); Hassan (n.d.); Soegeng-
Reksodihardjo (1962); Subhadrabandhu et al. (1991); Weber (1973).
Phytophthora sp., Anon. (1980a); Jamil (1965); Kwok-Kong (1974); Thaveechai

et al. (1982); Phytophthora botryosa, Chee and Hashim (1971); Suzui et al.
(1979b); Tai (1971); Zentmyer and Mitchell (1986); Phytophthora nicotianae
B. de Haan var. nicotianae -leaf blight, Anon. (1983); Liu (1977a,b); Pawakul
and Chittanawasarn (1977a, 1977b); Singh (1980); Phytophthora nicotianae
var. parasitica (d’Haan)Tucker, Suzui et al. (1976, 1979b), Weber (1973);
Phytophthora palmivora Butler. -root rot, bark rot, fruit rot, hypocotyl rot,
patch canker, Anon. (1986b); Butt and Sia (1982); Chan and Lim (1986;
1987); Chee (1969); Chee and Hashim (1971); Coronel (1986); Coronel et al.
(1983); Ding (1988); Ganapathi and Chinnathambi (1993); Giatgong (1980);
Hamilton (1975); Hassan (n.d.); Holliday (1980); Jamil (1966, 1969); Johnson
(1993), Johnston, (1959), Khew (1990, 1991); Khoo et al. (1983);
Kueprakone et al. (1977, 1979); Lee (1991), Lee et al. (1993, 1994), Lee and
Loh (1966); Lee and Varghese (1974); Lim (1991); Lim et al. (1992); Lim
and Chan (1986 a,b); Lim and Yassin (1985); Ling (1991); Meon and Varghese
(1986); Navaratnam (1966); Ng and Choong (1991); Nik Masdek et al. (1991);
Pawakul et al. (1977); Pawakul and Chittanawasarn (1977a; 1977b); Pupipat
(1984); Roger (1951); Shaji et al. (1993); Singh (1973, 1980); Snowdon (1990);
Soegeng-Reksodihardjo (1962); Subhadrabandhu et al. (1991); Suzui et al.
(1976, 1978, 1979a, 1979b); Tai (1970, 1971, 1973); Thompson (1934, 1938);
Thompson and Lim (1965); Tongdee Neamprem and Chayasombat (1989);
Tidbury (1976); Turner (1960); Vichirananda (1983); Vichitranond et al. (1981);
Watson (1984); Weber (1973); Zainal Abidin et al. (1992); Zentmyer et al.
(1973); Zentmyer and Mitchell (1986); Phytophthora parasitica (Butler) Dastur.,
Kueprakone et al. (1977).
Placosphaeria durionis Sacc. -leaf parasite, Baker (1914); Reinking (1919).
Plokamidomyces colensoi Bat., Costa & Cif., Lim (1989).
Polychaeton sp. -leaf mould and black crust of fruit, Lim (1989).
Pyrenochaeta sp. -leaf spot, Johnston (1960); Singh (1973, 1980); Williams
and Liu (1976).
Pythium sp. -root rot, Coronel (1986); Lee et al. (1993); Ng and Choong
(1991); Nik Masdek et al. (1991); Pythium complectans Braun [synonym of
P. vexans de Bary] -root disease, patch canker., Belgrave (1939); Coronel et
al. (1983); Johnston (1959); Meon and Varghese (1986); Pawakul and
Chittanawasarn (1977a; 1977b); Roger (1951); Singh (1973, 1980); Soegeng-
Reksodihardjo (1962); Thompson (1938; 1939); Watson (1984); Weber (1973);
Pythium vexans de Bary [see also P complectans which is a synonym], Ling
(1991); Shaji et al. (1993)
Rhizoctonia sp. -leaf fall, leaf blight, Chandrasrikul (1962); Giatgong (1980);
Vichirananda (1983); Rhizoctonia solani Kuhn. -leaf blight, Anon. (1986b);
Lim et al. (1987); Ng et al. (1986); Ng and Choong (1991); Nik Masdek et
al. (1991); Shaji et al. (1993); Singh (1973; 1980).
Rhizopus sp., Subhadrabandhu et al. (1991); Rhizopus artocarpi Racib. -fruit

rot, Johnston (1960); Singh (1973; 1980); Rhizopus stolonifer (Ehrenb. ex
Fr.) Lind., Lim (1990)
Rigidoporus lignosus (Klotzsch) Imazeki [see also the synonym Fomes lignosus]
-white root disease, Ng and Choong (1991); Nik Masdek et al. (1991); Shaji
et al. (1993); Singh (1980); Williams and Liu (1976)
Sclerotium sp., Ganapathi and Chinnathambi (1993); Sclerotium rolfsii Sacc.

(the sclerotial state of Corticium rolfsii Curzi) -fruit rot., Lim and Sijam
(1989a).
Scorias spongiosa (von Schweinitz) Fries emend. Reynolds., Lim (1989)
Septobasidium sp. -felt fungus, velvet fungus, Chandrasrikul (1962); Giatgong

(1980).
Spiropes capensis (Thumen) M.B. Ellis (=Helminthosporium capensis) -leaf

mould, Anon. (1980b); Singh (1980); Turner (1964, 1971)
Thanatephorus cucumeris (Frank) Donk. [telomorphic state of Rhizoctonia

solani Kuhn.] -leaf rot, Singh (1980); Weber (1973); Williams and Liu (1976).
Trametes persoonii Mont. -grows on bark of living trees, Reinking (1919),

(1920).
Trichomerium grandisporum (Ellis & Martin) Bat. & Cif. -sooty mould of
leaves, petioles and twigs, Lim (1989).
Trichopeltheca asiatica Bat., Costa & Cif. -black film on leaves, Lim (1989).
Tripospermum sp. -black colonies on leaves and fruits, Lim (1989).
Ustulina sp. -scion dieback, Singh (1973, 1980).
Verticillium sp., Ganapathi and Chinnathambi (1993).
Lichens
Buellia canescens (Dicks.) DeNot. -epiphyte, Anon. (1980b), Parmelia sp. -

epiphyte, Lim (1990)
Algae
Cephaleuros virescens Kunze. -red rust, Nik Masdek et al. (1991); Singh
(1980); Vichirananda (1983); Williams and Liu (1976); Phycopeltis sp. -largely
harmless epiphyte, Lim (1990); Pleurococcus nagelii -largely harmless epiphyte,
Lim (1990); Trebouxia sp. -largely harmless epiphyte, Lim (1990); Trentepohlia
sp., Long et al. (1991); Trentepohlia aurea (L.) Martius, Lim and Sijam
(1989b); Trentepohlia monile De Wildeman [=Physolinum monilis (De Wildeman)
Prinz. =T. moniliformis Karsten], Lim and Sijam (1989b); Trentepohlia arborum
(C.Ag.) Hariot [=T. bisporangiata Karsten.], Lim and Sijam (1989b).
Ferns
Drymoglossum pilloselloides (L.)Presl. -problematic epiphyte, Lim (1990);

Drynaria quercifolia (L.)J. Sm., Anderson (1966)
Angiosperms
Elytranthe barnesii -mistletoe, Sands (1924); Ficus benjamina L. -strangling

fig, Anderson (1966); Loranthus pentandrus L. -mistletoe, Sands (1924).
Insects
Ambrosia beetles -damage to timber, Thomas (1952); longicorn beetle larvae,

Chevalier (1934); Thomas (1952, 1979); Termites -damage to lumber of D.
carinatus, Isamanto and Sumarni (1992); Scale insects -damage to leaves,
Chiow (1976).
Acrocercops sp. -pest of leaves, Yunus and Hua (1980).
Adoretus sp. -pest of leaves, Shaji et al. (1993).

Allocaridara malayensis Crawford -pest of leaves, Lee et al. (1994).
Anomala sp. -pest of leaves, Shaji et al. (1993).
Apogonia sp. -pest of leaves, Shaji et al. (1993).
Aprosterna pallida Fabr., Boonyong (1983).
Asterolecanium sp. -leaf scales, Anon. (1959a); Chuan et al. (1981); McIntosh
(1951); Yunus and Hua (1980); Asterolecanium ungulata Russel, Lim et al.
(1991); Long et al. (1991); Yunus and Hua (1980).
Batocera gultata Vollenh. -stem borer, Nik Masdek et al. (1991); Shaji et al.
(1993); Yunus and Hua (1980).
Cacoecia machlopis Meyr. -pest of leaves, Yunus and Hua (1980).
Canopia sp. -borers, Coronel et al. (1983).
Carpophilus sp. -sap beetle, Lee et al. (1994); Carpophilus floveicollis -sap
beetle, Lee et al. (1994).
Cephonodes ?higlas L., Chiow (1976).
Chalcoscelis albiguttata Snell. -pest of leaves, Yunus and Hua (1980).
Conogethes punctiferalis Guen. -fruit borer, Mohd. Shamsudin (1992); Nik

Masdek et al. (1991); Shaji et al. (1993); Zainal Abidin et al. (1992).
Conopia sp. -trunk borers, secondary infection after P. palmivora, Navaratnam

(1966); Yunus and Hua (1980).
Cremastopsyche pendulla -leaf eating bagworms, Shaji et al. (1993).
Cryptotermes cynocephalus Light. -destructive wood pest, Ismanto and Sumarni

(1992); Martawijaya and Sumarni (1978).
Daphnusa ocellaris Walk. -Hawk moth which eats durian leaves, Ramasamy
(1980).
Dichocrocis punctiferalis Walk. [=Monogatus punctiferalis] -fruit borer,

Boonyong (1983); Coronel et al. (1983); Mohd. Shamsudin (1991); Mohd.
Shamsudin and Norsiah (1990); Yunus and Hua (1980).
Drosophila punctpennis v.d. Weele., Frannsen (1936).

Erizada lichenaria Wlk. -pest of leaves, Yunus and Hua (1980).
Eutetranychus africanus -pest of leaves, Lee et al. (1994).
Hypomeces squamosus F., Shaji et al. (1993); Yunus and Hua (1980).
Hypoperigea leprosticta Hamps. (=Plagideicta leprosticta) -durian fruit borer;

Frannsen (1936); Kalshoven (1981); Subhadrabandhu et al. (1991).
Idiophantis chiridota Meyr., Yunus and Hua (1980).
Indarbela flavina Mell., Yunus and Hua (1980).
Lacifer javanus Chamb., Yunus and Hua (1980).
Lyctus sp., Menon (1957).
Microtermes pallidus (Haviland), Nik Masdek et al. (1991); Yunus and Hua
(1980).
Minthea rugicollis Walk. -powder-post beetle, Browne (1938) [recorded on

D. lowianus and D. oxleyanus].
Monogatus puntiferalis Guen. (=Conogethes punctiferalis Guen.) -durian husk

borer, Lee et al. (1994).
Monolepta bifasciata Hornst., Shaji et al. (1993); Yunus and Hua (1980).
Mudaria luteileprosa Holloway -seed borer, Zainal Abidin et al. (1992);

Mudaria magniplaga Wlk. -seed borer, Lee et al. (1994); Mohd. Shamsudin
(1992); Nik Masdek et al. (1991); Shaji et al. (1993); Zainal Abidin et al.
(1992).
Oligonychus biharensis -pest of leaves, Lee et al. (1994).
Orgyia postica Walk., Yunus and Hua (1980); Orgyia turbata Butl., Yunus
and Hua (1980).
Orthaga semialba Meyr., Yunus and Hua (1980);
Paralecanium expansum Green, Yunus and Hua (1980); Paralecanium vacuum

Morr., Chuan et al. (1981); Takahashi (1939); Yunus and Hua (1980).
Phenacaspis hedyoticlis Green, Yunus and Hua (1980).
Phostria nicoalis Wlk., Yunus and Hua (1980); Phostria xipharesalis Wlk.,
Yunus and Hua (1980).
Plagideicta magniplaga Walk. [=Mudaria magniplaga] -durian seed and fruit

borer, Anon. (1986b); Butt (1982); Hassan (n.d.); McIntosh (1951); Mohd.
Shamsudin (1991); Mohd. Shamsudin and Norsiah (1990); Ng and Choong
(1991); Yunus and Hua (1980).
Platypus cupulatus Chap., Nik Masdek et al. (1991); Yunus and Hua (1980).
Postelectrotermes militaris, Thirugnanasantharan (1987).
Pseudococcus sp., Lee et al. (1994).
Rhadinomerus sp., Yunus and Hua (1980).
Rhytidodera simulans -stem borer, South (1922).
Saissetia sp., Yunus and Hua (1980).
Scheloribates sp., Yunus and Hua (1980).
Seira jacobsoni (Borner), Yunus and Hua (1980).
Squamura sp. -bark feeder, Shaji et al. (1993).
Sylepta derogata F., Yunus and Hua (1980); Sylepta bipunctalis Warr., Yunus
and Hua (1980).
Synanthedon sp., Nik Masdek et al. (1991); Yunus and Hua (1980).
Tenaphalara malayensis Crawford., Boonyong (1983); Braza and Calilung

(1981); Gadug and Hussein (1987); Hodkinson (1983); Shaji et al. (1993);
Tigvattnanont and Pramual (1990); Yunus and Hua (1980).
Tetranychus fijiensis Hirst., Lee et al. (1994); Yunus and Hua (1980).
Tirathaba mundella Walk., Yunus and Hua (1980); Tirathaba ruptilinea Hamps.,
Frannsen (1936).
Tonica terrasella Wlk. -fruit borer, Lee et al. (1994); Mohd. Shamsudin (1991);
Mohd. Shamsudin and Norsiah (1990); Mohd. Shamsudin (1992); Nik Masdek
et al. (1991); Zainal Abidin et al. (1992).
Tyrophagus putrescentiae (Schrank); Yunus and Hua (1980).
Xyleborus cordatus Hag., Yunus and Hua (1980); Xyleborus declivigranulatus

Sch., Yunus and Hua (1980); Xyleborus ferrugineus (F.), Yunus and Hua
(1980); Xyleborus formicatus Eichh., Boonyong (1983); Xyleborus similis Ferr.,
Yunus and Hua (1980); Xyleborus testsceus Wlk [=X. perforans Wall.], Yunus
and Hua (1980).
Zeuzera sp. -borers., Polchart (1952); Zeuzera coffeae Nietn. -stem borer,
Coronel et al. (1983), Nik Masdek et al. (1991), Yunus and Hua (1980).
Nematodes
Helicotylenchus sp., Fong and Poh (1970); Sidam and Yuen (1992); Singh
(1973).
Macroposthonia sp., Sidam and Yuen (1992).
Meloidogyne sp., Sidam and Yuen (1992).
Pratylenchus sp., Sidam and Yuen (1992); Pratylenchus coffeae
(Zimmerman)Filipjev & Schuurmans Stekhoven -hypocotyl rot, Giatgong (1980).
Radopholus sp., Fong and Poh (1970); Singh (1973).
Rotylenchulus reniformis Lin. & Ol., Sidam and Yuen (1992).
Tylenchorhynchus sp., Sidam and Yuen (1992).
Xiphinema sp. -attack roots, Ng and Choong (1991); Sidam and Yuen (1992).
Other animals
Squirrels can be a nuisance in durian plantations (Butt 1982). For instance,

a fruit loss of 15.6% was attributed to squirrels in one season (Chiow 1976).
Ithnin (1992) has examined the distribution of several rodents in mono- and
inter-cropped durian orchards. These include three species of squirrel
(Callosciurus canisep, Callosciurus nigrovittatus, Callosciurus notatus) and
one species of rat (Rattus tiomanicus). Squirrel populations were found to be
higher in durian plantations intercropped with cocoa. Rat populations were
unaffected by intercropping. The three squirrel pests of durian are also mentioned
by Nik Masdek et al. (1991). Shaji et al. (1993) remark that C. notatus can
be controlled in durian orchards by placing metal shields around the base of
the tree, or by the use of poisoned bananas. Oei-Dharma (1969) states that
rats, boars and bears are problem animals in Indonesia. Rats are also known
to cause damage to stored fruit awaiting shipment (Lee et al. 1994). Snails
have been listed as a pest of durian leaves by Shaji et al. (1993).
Hyperparasitism in durians : Several cases of hyperparasitism have been

reported on durian. In these instances, durian parasites are themselves parasitized.
One such case is that of the fungus Gliocladium roseum Banier., which is a
parasite of the durian parasite Phytophthora palmivora Butler. (Lim and Chan
1986b). P. palmivora is the causal organism of several durian diseases and is
probably its major pathogen. G. roseum penetrates and destroys the
chlamydospores of P. palmivora. Although this system immediately suggests
a form of biological control for P. palmivora related durian diseases, Lim and
Chan (1986b) stress that much comprehensive study is needed before this
system could be exploited as a method of control. Unfortunately, I have not
come across any further work in this area.
The scale insect Asterolecanium ungulata is an economic pest of

durians. Damage done by this insect often leads to secondary infection of the
tree. Lim et al. (1991) described the natural control of this pest by the fungi
Aschersonia placenta and Hypocrella raciborskii. More information on this
‘microbial pesticide’ is presented by Ibrahim et al. (1993).
A most spectacular case of hyperparasitism of D. zibethinus is related

by Sands (1924). Mistletoes are parasites of many tropical trees. They can
even be found parasitizing other mistletoes. Sands (1924) reported a specimen
of D. zibethinus upon which was growing Elytranthe barnesii. This mistletoe
was being parasitized by another mistletoe (Viscum articulatum), which, in
turn, was being parasitized by another (Loranthus ferrugineus).
Vegetative Propagation of Durians
Although durian seeds are viable and trees have historically been grown
from seed, durians are now preferably propagated asexually by grafting, seeds
being used for the production of rootstocks. Seeds of the cultivar Chanee are
used to produce rootstocks for grafting in Thailand (Subhadrabandhu et al.
1991). The major problems with direct sowing of seeds are that the genetic
makeup of the trees is unknown, and the emerging seedlings are susceptible
to predation by rodents. Seeds are now generally grown in a nursery in polybags
to produce material for grafting (Hassan n.d.).
Asexual reproduction of durian trees by grafting offers three main

advantages over trees produced directly from seeds. Firstly, the genetic nature
of the tree is known and can be controlled. This is desirable as fruit quality,
size, time of flowering, length of time to maturity and fruitfulness of trees
varies widely between clones. Secondly, grafted trees are generally smaller in
stature than are trees produced from seed and some clonal material (e.g. D99).
Consequently, more grafted trees can be planted per hectare of land and the
fruits are produced closer to the ground. Thirdly, grafted trees start bearing
fruits at an earlier age than do trees raised from seed.
The disadvantages of vegetatively propagated material are likewise
three-fold. Vegetative propagation is much more labour intensive, different
grafting methods have varying degrees of success, and durian trees are sensitive
to transplanting; budded durian trees frequently suffer 50% mortality (Feilden
and Garner 1936). Thus, it is preferable to keep trees in the nursery for
extended periods after grafting to increase the success of transplanting. Because
of the great advantages to be gained by asexual propagation, much research
has been conducted in this aspect of durian cultivation.
Etiolated shoot method : The etiolated shoot method is a type of layering in

which shoots are produced from partially buried branches. Some early work
on asexual propagation focused on the etiolated shoot method of propagating
durian (Lambourne 1935), which showed some success (Anon. 1935). Analysis
of results obtained using this method are published (Lambourne 1935), but
the technique does not appear to be very promising. The removal of some of
the bark from the shoot has been shown to promote faster root formation in
durian (Feilden and Garner 1936).
Double root system : ‘Three legged’ durian trees are created in Thailand
(Boonbongkarn 1960). Using this technique, the stems of a few seedlings are
grafted onto a more mature tree. This is done after the tree has reached
1-1.5 m in height. The resultant tree has more than one root system, is better
able to obtain food and water, and can withstand stronger winds (Boonbongkarn
1960). Old durian trees with declining productivity can apparently be revitalised
by grafting a young shoot onto the trunk, and then removing the original stem
(Boonbongkarn 1956). Ahmad and Ghani (1990) have noted that this technique
is occasionally used on D. zibethinus to induce rapid flowering.
Preliminary attempts have been made to graft durian scions onto

multiple rootstocks simultaneously (Zabedah et al. 1992; Zabedah 1993). These
experiments were carried out to investigate the possibility of creating more
vigorous planting material with a higher survival rate. Initial results indicated
that double and triple grafted material grew more vigorously and had higher
fruit set than grafted plants with single rootstocks (Zabedah et al. 1992; Zabedah
1993).
Approach grafting : Success with approach grafting of durian trees has been
reported using 6-8 months old seedlings (Chua and Teoh 1973). Up to 100%
success was reported by Chua and Yong (1978), however, success of graft
was higher using older (8-10 months) rootstocks, and an extended time before
cutback. Baga Kalie (1979) described a technique for simple splice approach
grafting of durian. Baga Kalie and Anwarudin (1980) compared results of
simple splice approach grafting and tongued approach grafting of durian;
these two techniques were found not to differ significantly in success rate.
The highest rate of success was achieved using 8.5 months old rootstocks. An
approach grafting technique for grafting seedlings onto branches of mature
trees is presented by Zainal Abidin et al. (1991). In Thailand, bagged rootstocks
are also grafted onto branches of mature trees by inarching (suckle grafting)
(Subhadrabandhu et al. 1991).
Inarching : Early experiments with inarching (a modified form of approach

grafting) of durians were carried out at the Burliar Research Station in India;
a success rate of 100% was obtained from inarches made in October, 80%
success in November, the success rate of inarching in other months varied
from 20 to 60% (Khan and Sambashiva Rao 1952). Furthermore, these authors
noted that the graft union took an inordinate amount of time to form. Although
inarching durians requires minimal skill, it is laborious, and is not recommended
for large scale propagation (Coronel et al. 1983). This method of grafting has
the disadvantage of requiring larger plants and thus is more expensive (Coronel
1986).
Top grafting : The effects of scion length for top grafting has been investigated
using four Thai cultivars of durian (Anwarudin and Sunarjono 1987). Scions
of 2, 3 and 4 nodes length were tested; no significant difference in graft take
was found. Some durian varieties were found to top graft more easily than
others. Rootstocks of different ages have also been tested to discover the most
suitable age. No significant difference in suitability was found in rootstocks
aged between 1.5 and 3 months (Anwarudin et al. 1987).
The wedge (cleft) grafting experiments of Chua and Yong (1978)

were not very successful with a maximum of 39% success using 6 months old
rootstocks. However, Chong (1985, 1992b) also investigated the success of
cleft grafting in 5 clones of durian at different rootstock ages. Chong reported
high rates of success (up to 100%), even with 5 weeks old rootstocks in all
clones tested. The technique is depicted diagrammatically by Chong and Chai
(1986), and is also described in detail by Chong (1992a). It differs from the
original technique of Chua and Yong (1978) in that the shoot was bagged, and
the plant was well watered for several weeks to ensure the scion received high
humidity. Cleft grafting has apparently become a popular technique for
commercial propagation of durian in the Philippines (Coronel 1986) as it is
fast and requires comparatively little skill.
Bimantoro (1982) described a method of cleft grafting durian scions

onto one month old durian seedling hypocotyls, avoiding having to grow and
mature rootstocks to the one year old seedling stage. Using this method, parts
of the leaves of the scion are removed to reduce transpiration, and the grafted
plant is covered by a plastic bag to maintain high humidity. Sunarjono and
Jawal (1985), and Sunarjono (1988) described the grafting of durian scions to
the epicotylar and hypocotylar regions of seedling rootstocks. The highest
success rate was reported for epicotylar grafting (Sunarjono 1988).
Hasan and Inoue (1989) investigated inlay grafting of durians, a modified

form of cleft grafting, which allows the joining of a scion with a rootstock
having a much greater diameter. This technique was investigated using 2.5
years old bagged rootstocks, and scions of different clonal origins. The effects
of the number of leaves on the scions was also studied. The grafts were
covered in polyethylene bags as described in Bimantoro (1982). When all
leaves were removed from scions, inlaying had a 100% success rate, the best
rate of success being obtained from two leaved scions in which the leaves had
been cut in half. There was large difference in success rate between different
clones; D2 and D7 had adequate success rates but the technique is unfortunately
unsuitable for propagating D24 which is one of the most desirable clones
(Hasan and Inoue 1989).
Budding techniques : Wester (1914c) reported on the shield budding of

durian using the inverted-T method. A success rate of 100% occurred when
leaf blades of the budwood were removed from the petioles before grafting,
so that a good leaf scar had formed when the petioles had abscinded. If the
above treatment was not carried out, bud decay occurred.
Budding experiments using the modified Fokert method resulted in

90% success during the rainy season, but only 40% during the dry season
(Magielse and Ochse 1931). Feilden and Garner (1936) reported that the
modified Fokert method was used in Java employing one year old non-petioled
budwood.
Naik (1949) stated that inarching experiments met with 50% success,
but there was no success in shield budding; however, he did not describe his
shield budding technique. Wester (1918) provided the following instructions
regarding shield budding of durian: ‘Use non-petioled, fairly mature, but not
old and hard budwood; cut the bud 2.5 to 3.5 cm long; age of stock at point
of insertion of bud unimportant’.
Belgrave (1939) reported that bud patches from previously ring-barked

branches are superior for grafting than bud patches from non-ring-barked
branches. It is important to note that buds will fail to unite if they become wet
(Sankhariksha 1968; Coronel et al. 1983).
Sankhariksha (1968) reported low success using several budding

techniques, however, these results were attributed to the fact that the rootstocks
were not actively growing at the time of grafting. This report did, however,
demonstrate that plastic wrap yielded a higher percentage of ‘takes’ than did
traditional budding tape.
Bud grafting experiments with durian have showed that a high percentage
success can be obtained. Younger rootstocks are preferable (in this case 4
months) (Chua and Yong 1978). Coronel (1986) stated that patch budding is
the least popular method used in the Philippines due to the complexity of the
technique, and the need for more mature plant material. Commercial bud
grafting of durian has been practised in Malaysia since at least 1954 (Dickinson
1959); according to Hassan (n.d.), bud grafting is the most common form of
durian propagation practised in Malaysia.
Other asexual propagation techniques : A side grafting technique for durian

is described pictorially by Yong-Ho (1988); this technique requires the use of
5-6 months old bagged rootstocks. I have found no published information on
success rates using this technique. Very little information on marcotting of
durians is available (Coronel et al. 1983); marcotting can be accomplished but
it is not recommended. An investigation on the formation of roots on stem
cuttings of durian has been published (Hasan and Dodd 1989). Two years old
durian seedlings were found to readily form adventitious roots when the stems
were cut, dipped in IBA and a fungicide, and planted.
Very little work on the tissue culture of durian has been published.
Some success with shoot tip culture of D. lowianus has been reported. Although
durian tissue can be made to callus, no report of its regeneration has been
published.
Grafting to other species : Experiments have been carried out at the Burliar
Fruit Station to graft D. zibethinus scions onto Cullenia excelsa rootstocks
(Anon. 1953b). Grafting onto Cullenia excelsa rootstocks in India has been
reported by Sundararaj et al. (1970). Tidbury (1976) tabulated results of grafting
experiments onto rootstocks of this species at the Burliar Research Station.
These results showed that approach grafts to C. excelsa rootstocks averaged
much greater growth than did grafts to D. zibethinus rootstocks. Grafting to
C. excelsa also contributed to earlier flowering. Unfortunately, no record of
actual success rates of grafts to C. excelsa rootstocks were uncovered in the
available literature.
As early as 1934, Chevalier recommended grafting D. zibethinus scions

to rootstocks of one of several wild species including : D. graveolens, D.
dulcis, D. carinatus and D. gratissimus Becc. (now D. oxleyanus Griff. sensu
Kostermans 1958b). Hamilton (1975) and Ding (1988) suggested attempting
to graft D. zibethinus onto rootstocks of other durian species which may be
resistant to Phytophthora palmivora in order to provide resistance against this
important pathogen. In this regard, it should be noted that the work of Tai
(1973) showed that several clones of D. zibethinus also differ in their susceptibility
to infection by P. palmivora.
In their review, Malo and Martin (1979) stated that D. zibethinus
grafts well onto D. malaccensis. More recently, Pupipat (1984) has reported
that the resistance originally possessed by D. malaccensis roots to Phytophthora
has since broken down, and that grafting on this species is now useless as a
preventive measure.
Pawakul and Chittanawasan (1977a,b) reported that D. lowianus is
a suitable stock for grafting D. zibethinus. Furthermore, they demonstrated
that D. lowianus is more resistant to the pathogen P. palmivora than D.
zibethinus. Thus, these authors recommended grafting D. zibethinus scions
onto rootstocks of this species as a disease control measure. Watson (1984)
and Subhadrabandhu et al. (1991) stated that D. lowianus and D. mansoni
have resistance to P. palmivora when used as rootstocks. Tinggal (1993)
suggested that D. testudinarum may be tolerant to several soil-borne diseases
including Phytophthora, and that it would make excellent rootstocks for
grafts with D. zibethinus. However, no results of the success of grafts with
this species have been published.
Songpol (1987) described the results of grafting scions from 8 species
of durian to D. zibethinus rootstocks. Scions differed greatly in their ability
to take, D. lowianus and D. kutejensis had the highest percentage of takes
(approx. 75%) while intraspecific grafts of D. zibethinus cv. Chanee showed
only 33% success after 1.5 months. After one year, D. graveolens scions
showed the highest percentage of takes (approx 70%). Scions of different
species were also shown to vary in the increase in diameter of the graft union
stem diameter, stem length and number of nodes produced.
Grafting to rootstocks of D. mansoni may have a dwarfing effect on
the tree (Subhadrabandhu et al. 1991) as the rootstock and the scion thicken
at different rates, however, a low percentage of graft occurs.
Although frequent claims are made that grafts of D. zibethinus scions
to rootstocks of other species could be useful in overcoming diseases of
economic importance, it seems unlikely that this would hold true in practice,
especially since such grafted trees would need to be planted in large numbers
to significantly overcome the economic effects of the disease. The broken
resistance of D. malaccensis stands as an example of the futility of this simplistic
approach (Pupipat 1984). Grafting to other species could perhaps become a
useful tool to produce trees of smaller stature, trees which bear fruit at an
earlier age, and possibly grafted seedlings with a greater survival rate upon
transplanting. Future research on inter-specific grafts would do well to concentrate
on these objectives.
Advanced planting material : In order to overcome the problems of establishing

grafted seedlings in an orchard, especially when no or little irrigation is provided,
the use of ‘Advanced Planting Material’ is recommended (Zainal Abidin et al.
1992). Such material consists of grafted seedlings allowed to reach 15-18
months in the nursery before planting. This material significantly increases
the survival rate of trees. Kamariah and Mohammad Isa (1992) have presented
some evidence that planting material with a girth of less than 2 cm has a
significantly higher mortality rate. Another major advantage to using advanced
planting material is the shortening of the time trees take in the orchard before
they bear fruit. Normally, grafted durian trees take 6-7 years before they
flower, advanced planting materials can flower after 3.5-4 years of growth in
the orchard (Zainal Abidin et al. 1992).
‘Hybrid’ Durians : Durian hybrids (clonal) have become a valuable and

desirable new commodity. Three clones, MDUR 78, 79 and 88, have been
released by MARDI, and are the products of experimental crosses between D.
zibethinus clones D10 and D24 (Zainal Abidin et al. 1992; Zainal Abidin
1993). These ‘hybrid’ clones show combinations of desirable fruit and tree
characteristics including a natural increase in shelf-life of the mature fruits,
desirable shipping characteristics, and the trees bear fruits at a relatively early
age (Zainal Abidin et al. 1992).
Economics and Prospects for Development

A description how durian fruits were locally grown and marketed in
Malaysia in 1954 is provided by Wilson (1954). Today, this account is more
of historical value than of relevance to modern durian economics. More recent
information on strategies to develop durian as a domestic and export crop in
Malaysia has been published (Ali 1993; Hamid 1993; Lim 1993). Also, a
study of the preferences of Malaysian consumers for durian fruits has recently
been published by Syed Abdillah et al. (1993). An excellent description of the
marketing of durian fruits in different ASEAN nations is provided by Alim et
al. (1994).
There is little immediate possibility of successfully developing durian

as a crop outside Austral-Asia. The durian has no prospects for development
as a crop in Western Australia (Boughton et al. 1983; Hawson 1983), and
is unsuitable for growth in the Mediterranean (Galán Saúco 1994). Durian
is, however, suitable for growth in Northern Australia (35 hectares were
planted as of 1990), however, durian can only be grown north of 17o S
(Watson 1993). Watson (1993) stated that the demand for fruit is great enough
that the hectarage could easily be tripled. The major problem with its
introduction into Australia has been the planting of unsuitable clones due
to misidentification. Chadha and Pareek (1988) have suggested that durian
is worthy of consideration for development in India.
In 1987, Malaysia had almost 40 000 hectares under durian cultivation

(Hasan 1989), whereas Thailand had 84 500 hectares under cultivation with
a production of 222 000 tonnes (Subhadrabandhu 1992) (a figure
approximately double the recorded tonnage in 1983).
Although durian may be unsuitable for growth outside Austral-Asia,

the potential for valuable export markets certainly exists. Malaysian trade
statistics for 1988 showed that durian already accounts for over 40% of fruit
exports (Karim 1992). Although, currently most exported durians end up in
Hong Kong (Wurdack 1985), the possibility of developing other overseas
export markets remains open (Ito and Hamilton 1990). Japanese import
statistics (Kitagawa et al. 1990) show that imports of durians into Japan
almost tripled during 1986-1988. Although export markets for fresh durians
certainly exist, Nichols (1992) has suggested that durian might be better
promoted in the form of processed products.
The cultivation of durian is a rapidly growing enterprise as the

following statistics will show. Thailand is undoubtedly the worlds largest
producer of durian, thus several statistics on their production are available
(Watson 1984; Visetbhakdi 1988; Paweenakarn et al. 1992). The main durian
growing regions of Thailand are the provinces of Rayong and Chanthaburi
(Subhadrabandhu 1992; 1993). In 1985, Thailand produced 167 800 metric
tonnes of durian fruits, of which 9800 tonnes were produced for export
(Paweenakarn et al. 1992). In 1989, 89 927 ha were planted with durian
with a total production of 486 644 tons [494 430 tonnes] (Subhadrabandhu
1993). The export potential for Thai durians has been steadily growing; exports
almost tripled over the 6 years period from 1981 to 1987 (Visetbhakdi 1988);
7 million baht worth of durians were exported to Canada alone in 1986,
over twice the amount of longan and pomelo exports to Canada combined
in the same year.
According to Watson (1984), Indonesia is the second largest producer

of durian fruits with a total production of 157 000 tonnes (Circa 1975).
However, the second largest producer of durians today is more likely Malaysia.
In 1962, 14 025 ha were under durian cultivation in Malaysia (Jamil 1965).
However, in 1972, only 8296 ha were under cultivation. Durian cultivation
has been steadily increasing since the 1970s with 31 225 ha reported in
1985, 49 820 ha in 1989 (Ali 1993). The annual production of durians in
Malaysia in 1991 was 250 000 tonnes (Zainal Abidin et al. 1992). Johore
is the major durian growing state in Malaysia (Lim 1990), accounting for
32% of the total durian hectarage in the country (Alim et al. 1994).
Furthermore, durian currently occupies the largest area under cultivation in
Malaysia of any fruit tree species (Jamil 1992c).
A thorough description of the economics of production and marketing

of processed durian products in the Philippines is presented by Lisondra
(1984), but few published statistics on durian acreage in this country have
been encountered. Durian is a relatively minor crop in the Philippines, most
likely owing to its limited distribution. In 1977, only 1080 hectares of durian
were grown in the Philippines with a total production of 6048 tonnes (Coronel
et al. 1983), however, in 1985, 1900 hectares of durians existed with a total
production of 14 992 tonnes, which represents more than a doubling of
production in less than ten years.
Nichols and Christie (1993) state that development should concentrate

on germplasm conservation and development of efficient production and post-
harvest technology. Setefarzi et al. (1991) state that the biggest problem to
be overcome is the lack of profitability which springs from a lack of
technology, knowledge and high investment costs.
Significant genetic erosion within the genus Durio has already been
reported (Sastrapradja 1975), especially in Malaysia, Thailand and Indonesia,
where clonal varieties have become popular. Hasan (1989) stated that within
Malaysia, durians should be the first priority in development. Karim (1992)
suggested that different flowering seasons of different clones should be
exploited on a national plantation scheme to spread the durian season. This
idea is thoroughly discussed by Zainal Abidin et al. (1986) (see section
on manipulating seasonality). Karim (1992) also suggested that research
is needed to reduce the gestation time of the fruit. Pietrzyk (1992) has
stated the need for the creation of cDNA libraries for durian and other
Malaysian fruits.
Genetic Resources and Conservation

The issue of plant genetic resources conservation is one that hardly
needs introduction. In recent years, concern has arisen that the genetic variability
of many of the world's most widely grown crops (i.e. wheat, corn, rice) have
succumbed to some degree of genetic erosion. Mere popularity of a crop does
not ensure the maintenance of genetic diversity necessary for the creation of
future varieties, in fact it often produces the opposite effect: those who grow
crops are usually interested in increasing their productivity by replacing old
and perhaps poorer yielding varieties with new ones.
Although we have very little first hand knowledge of the degree to

which durian is threatened by an erosion of genetic resources, it is quite
reasonable to suppose that genetic erosion is a very real problem. The majority
of durian species are endemic to Borneo which is suffering from intense
deforestation. Very little, if any, information is available on the natural genetic
variability that exists within durian. For D. zibethinus, it is tempting to guess
that genetic variation is high, based on the large number of clones now catalogued,
however, the amount of variability that exists in any of the wild species of
durian is simply unknown. RFLP analysis and isozyme studies are methods
which could very easily be applied to answering this queston. Research efforts
in this direction would not be prohibitively expensive and would certainly be
worth the investment. Such knowledge would aid in estimating the extent, if
any, of the problem and provide empirical evidence that remedial action is
requried.
Conservation of genetic resources, especially for plant such as durian,

needs to be approached holistically in order to achieve desired results. For
this reason, this section appears at the end of this work, so that it may draw
upon the wealth of knowledge that does exist on durians that has been discussed
so far. There are several unique problems that surround the genetic conservation
of Durio. Firstly, durian cannot be grown with any success outside the humid
tropics, limiting sites for the collection of varieties. The durian is not held in
great esteem in most of the world (excepting South East Asia), nor is it
terribly well suited for growth in other tropical areas of the world. The genus
is unsuited for conservation in traditional seed banks as the seeds have a very
short period of viability. Some studies have been conducted on cryopreservation
of seeds (see section on viability) but no success has yet been achieved.
Growing plants in a 'living germplasm' collection has advantages over
conventional seed bank storage, but as all members of the genus are large
trees, maintaining large numbers of accessions is costly and not always practical.
A diverse range of genetic resources to draw upon is crucial for this genus,
as it is a crop that has undergone very little domestication and one which has
a large potential for improvement. The genetic resources of the common
durian are threatened, to some extent, apart from deforestation, due to the new
'improved' clones themselves. The proliferation of popular varieties has led to
considerable genetic erosion as less popular non-clonal trees are replaced
(Sastrapradja 1975). This is a particular problem in Indonesia, Malaysia, Thailand
and Vietnam (Smith et al. 1992).
The wild species of Durio present another and perhaps larger challenge
for genetic conservation. D. zibethinus faces the dilution of its gene pool by
the large scale planting of genetically identical clones. By comparison, all
of the wild species are quite rare, some are in danger of extinction and a few
may already be extinct. It is claimed that a D. macrantha tree growing in
Bogor (a newly described species with edible fruit) may be one of the last
living specimens (Anon 1992). Smith et al. (1992) stated that several wild
species are threatened by logging. Although admittedly not much is known
about the genetic diversity of most of the wild species, erosion has been
documented for D. kutejensis (Sastrapradja 1973b). The genetic resources of
the wild species are equally, if not more, valuable than those of D. zibethinus
for three reasons. Firstly, some of these species do have potential as fruit
tree crops themselves. Although not grown anywhere near the scale of D.
zibethinus, several other species are cultivated. D. kutejensis is cultivated in
the eastern part of Kalimantan and D. oxleyanus is in rare instances cultivated
in some Bornean villages (Soegeng-Reksodihardjo 1962). At least a dozen
of the known species have edible fruits (see section on edibility), although
the majority of these have never been cultivated. Some of the these wild
fruits may actually be superior in quality to those of many of the clones of
D. zibethinus. Secondly, there is evidence that some of the wild species may
have higher resistance to some of the major durian pathogens (Phytophthora
in particular). Hamilton (1975) has already called for the investigation of
resistant rootstocks to P. palmivora from other durian species. There is
interest in not only using wild Durio species as rootstocks for D. zibethinus
(as Phytophthora is a root disease), but in producing true hybrid trees.
Thirdly, the possibility of producing hybrid trees opens the door to improvement
possibilities far beyond resistance to pathogens. Several Durio species are
so closely related to D. zibethinus, they most likely do, or have already been
shown to hybridize with it. Possibilities thus exist for altering the size of the
tree, time of flowering, reducing the age to first flowering, affecting seasonality,
increasing yield, improving the flavour, smell, colour, texture and size of
the fruits.
Wild rootstocks were rather recently held to be the solution to

Phytophthora infections, however, attempts to implement this as a solution by

mass planting of grafted trees have not only failed, but destroyed the resistance
that originally existed in D.malaccensis (Pupipat 1984). The key in maintaining
diversity in Durio and successfully using it, while avoiding similar misfortunes,
lies in a multifaceted approach to durian improvement. Perhaps a three pronged
approach such as the following is a solution.
There is a role that can be played by seed bank type institutions.

Although the seeds of durian cannot be stored conventionally, successful
cryopreservation may, with research, be achieved. Even better would be the
production and cryopreservation of callus. In my experience, durian callus is
relatively easy to produce, and it seems a more likely candidate for surviving
cryopreservation than do the seeds. What is so far lacking is not a method for
callus production, but rather a method of regenerating plants from callus.
Attempts to regenerate durian callus have so far been unsuccessful, but very
few attempts have been made. Developing methods for the successful preservation
of durian germplasm may simply be a matter of informing researchers that
this is a problem that needs attention. Much research on durian has been
carried out in several ASEAN nations, however, much repetition exists. A
combined effort of the ASEAN nations is a key to surmounting this problem
and perhaps to increasing awarenss of genetic resource erosion of durian as
well as identifying those areas of research most in need of attention.
Germplasm collections of trees (not necessarily the most popular

clones) are of great value. Collection of durians in this manner has already
begun. Chang and Lee (1975) stated that Singapore has carried out some
collection and conservation of D. zibethinus (although the extent of this remains
unknown). Attempts to collect and preserve genetic variability in 3 species of
Durio in Indonesia have been reported (Anon. 1979b).
Several collections of considerable size already exist (Table 10). The

MARDI germplasm collection, one of the largest collections of Durio in the
world, consisted of about 40 trees in 1975 (Arasu 1975) but now contains
almost 400 accessions (Chan 1992). However, such collections are not in
themselves the entire answer. These collections need to be supplemented by
the preservation of large tracts of forest in areas where wild species are naturally
found (Lamoureux 1975). Reforestation could even be exploited to these ends
as durian trees have been recommended for such purposes (Djober 1953).
Furthermore, some consideration needs to be given as to which materials need
to be collected, especially in the case of the wild species. Hopefully, this
manuscript has provided some insight into this matter.
In utilizing genetic resources for the improvement of durian, we need

Table 10. Durian germplasm holdings at different locations
Country Location Accessions No. Accessions No. Source

(D. zibethinus) (other Durio
species)
Australia Darwin, N.T. 10 1
Guadeloupe, CIRAD 2 Thai, 1 Indonesian 2

France
Indonesia Bogor (Agricul- 86 advanced 2

tural University) Indonesian cultivars
Bogor (National ? ? 2
Biological Institute)
Lembang 20 advanced 1,2

Indonesian cultivars
Malaysia Kuala Lumpur 310 (including other 1

species)
Penang (Relau Two collections: 4

Agricultural 25 varieties (mainly
Centre) from Penang)
and 40 varieties (from
other Malay states)
Sabah 75 clones D. dulcis (1) 6

(Department of D. graveolens (3)
Agriculture) D. kutejensis (4)
D. oxleyanus (5)
D. testudinarum (2)
Durio sp. (1)
MARDI 122 accessions in the 3

breeders’ collection
and 394 accessions in
the MARDI-IBPGR
germplasm collection
(Contd....)
Table 10. Contd.
Papua New Lowlands Agri. 1 2

Guinea Experimental
Station
Philippines Los Banos 97 advanced cultivars 1,2

+others
Location 200 accessions 5

unknown
Taiwan Chiayi Agri. 1 2

Experiment
Station
Thailand Bangkok 68 Thai 1,2

Chanthaburi 504 (500 advanced D. graveolens (102) 1,2
Thai, 4 old D. kutejensis (14)
Philippines) D. malaccensis (5)
D. mansoni (158)
D. oxleyanus (8)
Durio spp. (36)
USA Miami, Florida 16 1
National Clonal 6 advanced Thai 1,2

Germplasm lines
Repository Hilo,
Hawaii

1
Anon. (1986a).
2
Bettencourt et al. (1992).
3
Chan (1992).
4
Murad et al. (1993).
5
Smith et al. (1992).
6
W.W.W. Wong (1993).
to ask the question of exactly what is meant by improvement. If improvement

entails increased resistance to pathogens and diseases, increased productivity,
decreased seasonality and increased number of cultivars, then improvement
means diversity, not homogeneity. Thus, plans for ‘improvement’ that involve
the use of genetic resources for the mass production of new clones/hybrids/
grafts are not actually improvement, but the opposite, and may exacerbate the
problems they allegedly are trying to solve. This hardly means that the genetic
diversity of a genus such as Durio cannot be used for improving durians in
various ways, indeed it should, but actual improvement requires some foresight
as to the consequences of actions.
The ability to improve durians by providing new flavours for those

who wish to eat them, spreading out the season and coping with pathogens
lies in diversification of the materials that are planted. This is perhaps more
crucial for durians than for annual crops as the risks of monoculture have
more long-term effects (5-7 years to start up a new orchard).
As discussed above, durians are generally outcrossed, and the nature

of the parent trees may affect the degree of fruitfulness. Clones vary in their
susceptibility to pathogens as do the varied species. Durians also vary greatly
in their flavour, size and desirability. For these reasons, the most intelligent
approach to designing orchards should be carefully planned diversity; to their
credit, this appears to be the approach taken by MARDI (Chan 1992). A
diversity of rootstocks, a diversity of clones, a diversity of hybrids, and a
diversity of species, and in this way the numerous and varied advantages of
different trees can all be exploited and the varied disadvantages diluted. This
approach also offers the real possibility of extending the durian season and
thus increasing availability and profitability. It is easy to see how this approach
could limit the effect of pathogen outbreaks. The exploration of as yet uncultivated
(or barely cultivated) species offers the potential for entire new markets.
In conclusion, much has been discovered and written about the

mysterious durian fruit, but so much more remains to be achieved. I hope this
review has helped to identify and polarize those areas which are in need of
further clarification. While we strive to accomplish new goals through our
many varied researches, I would remind the reader of the poignant words of
Alfred Russel Wallace who, in my estimation, has written the most profound
words about man’s relationship with this noble fruit.
“Poets and moralists, judging from our English trees and fruits, have
thought that there existed an inverse proportion between the size of the one
and the other, so that their fall should be harmless to man. Two of the most
formidable fruits known, however, the Brazil Nut (Bertholletia) and the Durian,
grow on lofty trees, from which they both fall as soon as they are ripe, and
often wound or kill those who seek to obtain them. From this, we may learn
two things: first, not to draw conclusions from a very partial view of Nature;
and secondly, that the trees and fruits and all the varied productions of the
animal kingdoms, have not been created solely for the use and convenience
of man” — Wallace (1856)
Bibliography
The following bibliography consists of over 1000 references (largely

of a scientific nature) pertaining to durian. The bibliography consists only
of items of which an original or a facsimile has been examined by the
author. Much of the research conducted on durian has been published in
extremely obscure and difficult-to-obtain books, journals, conference
proceedings and reports. As these materials have often never been abstracted,
the need for a comprehensive bibliography is even more pressing. This
bibliography attempts to present detailed (non-abbreviated) citations for all
items, to aid others in identifying and obtaining copies of these works. The
bibliography also contains parenthetic notes [square brackets] which have
been added to provide additional information, such as the language of
publication, what portions of the work (if any) are written in English, where
possible an English translation of non-English titles and/or information on
published translations of works. The date(s) following the author’s name(s)
represent the actual date of publication (as far as is known). This occasionally
deviates from the dates printed on the work, especially for older works,
annual reports, etc. Explanatory notes are given as appropriate. To
accommodate to the realities and limitations of obtaining copies of scholarly
works, and the severe limits imposed on research by modern copyright
legislation, the pages pertaining specifically to durian in long works and/or
works in which durian forms only a small and discreet portion of the whole
are noted at the end of references, when appropriate. Attempts are made to
clarify the identity of species where the current taxonomic identity has changed
from that used in the publication. Although exhaustive attempts were made
to fully identify and give credit to all authors responsible for each publication,
this is often difficult or impossible to accomplish. All works, to which no
actual person could be attributed authorship (including publications of
institutes, associations, etc.), are listed chronologically as anonymous. I
apologize in advance to the authors of what I estimate to be approximately
150 research articles on durian not included in this bibliography. Despite
exhaustive efforts, copies proved impossible to locate or examine.

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Durio - A Bibliographic Review

INTERNATIONAL PLANT GENETIC RESOURCES INSTITUTE

© International Plant Genetic Resources Institute

The International Plant Genetic Resources Institute (IPGRI) is an autonomous international

ISBN 92-9043-3 18-3

Copies can be had from:

IPGRI Regional Office for Asia, the Pacific and Oceania

IPGRI Office for East Asia

Under the IPGRI Project on ‘Promoting Conservation and Use of

Dr. Kenneth W. Riley

Despite the preponderance of information that exists on durian,

Currently, no comprehensive bibliography on durian literature exists

The research conducted on some of the world’s major crops (rice,

In Malaysia, the value of durian exports alone accounted for over

Studies, which have been conducted on durian, have been scattered

importance. For these reasons, a comprehensive bibliography for durian is

De Clercq (1909) enumerated many vernacular names for durians

In this book, the word ‘durian’ will be used to refer only to D.

Historical works : The durian appeared in pre-Linnaean literature as early as

The earliest European description of the durian is perhaps that of

one form or another in several later publications. An English translation of

many of the taxonomic problems. The genus Durio, as stated by Chevalier

The type species Durio zibethinus is attributed to (Murr.); J.A. Murray,

The authority L. appeared several times in early taxonomic literature.

As stated previously, the earliest valid publication on the species is

As Durio is not mentioned in the 12th or earlier editions of ‘Systema

Formation of the modern concept of Bombacaceae : In early times, there

De Jussieu (1789) placed durian in the Capparideae, largely due to

What is now the family Bombacaceae was originally treated as a

the three subtribes of the former Bombaceae (Adansonieae, Matisieae, Durioneae)

Wild species of durian : The taxonomy of the common durian is convoluted

In 1889, Beccari produced volume three of ‘Malesia’ which included

King (1891) described only 6 species of Durio (D. zibethinus, D.

The genus was thoroughly reworked by Bakhuizen Van Den Brink in

new species enumerated by Beccari, 5 were dropped in Bakhuizen Van Den

Corner (1939) offered a tentative scheme to untangle some of the

Kostermans' (1958b) book stands as the most recent and useful

Post-Kostermans (1958b) treatment : Since the publication of his monograph,

I am not convinced that further subjective analysis of herbarium material

Soegeng-Reksodihardjo (1965) described a new species, D. burmanicus,

Kochummen et al. (1970) proposed that the species D. macrolepis

An incomplete specimen collected from Johore, Malaysia (FRI 8677)

Kostermans has recently published descriptions of two new species,

Table 1. Published species and synonyms of Durio

Boschia acutifolia Mast. (1874b)

Species Fruit Flower Seed Tree Leaf

Cullenia ceylanica LN5LN13 LN5BW13 LN13BW17 BW10 (BW,LN)13

Cullenia exarillata LN25LN33* LN25LN33* LN25 LN25LN33*

Cullenia rosayroana LN33* LN33* LN13LN33*

Durio acutifolius LN6 LN4c LN4c BW15 LN6

Durio affinis LN6LN15,16 LN4LN15,16 LN4LN6

Durio beccarianus LN15,16 LN15,16

Durio bukitrayaensis LN18 LN18 LN18

Durio burmanicus LN28 LN28

Durio carinatus LN4d LN4d LN4d LN4d

Durio excelsus LN12* LN2LN12 LN7* LN12*