Dipterocarpaceae (Flora of Sarawak and Sabah)

TREE FLORA
of
SABAH AND SARAWAK
Volume Five
edited by
E. Soepadmo, L.G. Saw and R.C.K. Chung
Government of Malaysia
TREE FLORA
of
SABAH AND SARAWAK
Produced with the financial support of
MINISTRY OF SCIENCE, TECHNOLOGY AND INNOVATION

GOVERNMENT OF MALAYSIA
Tree Flora of Sabah and Sarawak Editorial Committee ¾ Chairman:

E. Soepadmo (Forest Research Institute Malaysia). Members: A. Latiff
(Universiti Kebangsaan Malaysia), Ruth Kiew (Singapore Botanic
Gardens), L.G. Saw (Forest Research Institute Malaysia), R.C.K. Chung
(Forest Research Institute Malaysia). Editorial assistants: W.K. Soh
(Forest Research Institute Malaysia), S. Julia (Forest Research Centre,
Kuching). Advisors: P.S. Ashton (Harvard University, U.S.A.), J.
Dransfield (Royal Botanic Gardens, Kew, U.K.), W.J.J.O. de Wilde
(National Herbarium of the Netherlands, Leiden Branch, the
Netherlands).
TREE FLORA
of
SABAH AND SARAWAK
Volume Five
edited by
E. Soepadmo, L.G. Saw and R.C.K. Chung
A joint publication of
Sabah Forestry Forest Research Sarawak Forestry

Department, Malaysia Institute Malaysia Department, Malaysia
2004
electronic version prepared by

L.G. Saw & B. Aslina (2008)
Copyright Ó Forest Research Institute Malaysia (FRIM)
Sabah Forestry Department, Malaysia
Sarawak Forestry Department, Malaysia
Correspondence:
Forest Research Institute Malaysia (FRIM), Kepong, 52109 Kuala Lumpur, Malaysia
First published 30 September 2004
Perpustakaan Negara Malaysia CataloguinginPublication Data
Tree Flora of Sabah and Sarawak / edited by E. Soepadmo, L.G. Saw and
R.C.K. Chung.
Includes bibliographical references and index
ISBN 9832181593
1. Botany–Sabah. 2. Botany–Sarawak. 3. Botany–Classification.
I. E. Soepadmo. II. Saw, L.G. III. Chung, R.C.K.
581.959521
Front cover: Lowland and hill forests on ultramafic soil, Bt. Tawai FR, Sabah. (Photograph
by E. Soepadmo.)
Back cover: Koompassia excelsa (Becc.) Taub. with new growth. (Photograph by E.
Soepadmo.)
CONTENTS
Page
Foreword vii
Acknowledgements ix
FAMILIES:
1. Apocynaceae (David J. Middleton) 1

Alstonia R.Br. (Kade Sidiyasa) 13
2. Dipterocarpaceae (P.S. Ashton) 61
3. Symplocaceae (K.G. Pearce) 383
4. Thymelaeaceae (C.S. Tawan) 427
Appendix: New Dipterocarp Species from Sabah and Sarawak 479
Abbreviations of Frequently Cited References 481
Commonly Used Abbreviations for Localities 485
Plates 487
Index to Trade Groups of Shorea (Dipterocarpaceae) 501
Index to Scientific Names 503
Index to Vernacular Names 517

TREE FLORA OF SABAH AND SARAWAK
DIPTEROCARPACEAE
P.S. Ashton
Arnold Arboretum, Harvard University Herbaria, Cambridge, Massachusetts, U.S.A. and
Royal Botanic Gardens, Kew, England, U.K.
Blume, Bijdr. Fl. Ned. Ind. (1825) 223; F. Heim, Rech. Dipt. (1892) 1; King, J. As. Soc. Beng. 62, 2
(1893) 87; Brandis, J. Linn. Soc. Bot. 31 (1895) 1; Merrill, EB (1921) 397; Ridley, FMP 1 (1922) 209;
Gilg in Engler & Prantl, Nat. Pflanzenfam. edition 2, 21 (1925) 237; Foxworthy, Philip. J. Sci. 6
(1911) Bot. 231, Philip. J. Sci. 13 (1918) Bot. 163, Malay. For. Rec. 10 (1932) 4, Philip. J. Sci. 67
(1938) 241; Masamune, EPB (1942) 482; Symington, Malay. For. Rec. 16 (1943) 1; Brown, FTSB
(1955) 88; Backer & Bakhuizen f., FJ 1 (1964) 328; Ashton, MDB (1964) 1, MDBS (1968) 1, FM 1, 9
(1982) 237, in Kubitzki et al. (eds.), Fam. Gen. Fl. Pl. 5 (2000) 182; Meijer & Wood, Sabah For. Rec.
5 (1964) 9; Burgess, TBS (1966) 93; Anderson, CLTS (1980) 110; Kessler & Sidiyasa, TBSAEK
(1994) 86; Coode et al. (eds.), CLBD (1996) 66; Newman et al., MDFBLHW (1996) 55, MDFB
MHHW (1998) 57.
Mostly large canopy or emergent, resinous trees; usually buttressed. Bark smooth, fissured
or flaky. Indumentum comprising fascicled hairs, sometimes also simple hairs, unicellular
or multicellular glandular hairs or short or longlobed peltate scales. Shoot growth
rhythmic. Twigs with small or large, sometimes speciesspecific (Dipterocarpus), shoot
buds. Stipules speciesspecific in some taxa (especially in Dipterocarpus), generally
caducous leaving distinct scars. Leaves alternate; blade simple, generally penninerved, base
generally terminating abruptly at the frequently geniculate petiole, margin entire though
sometimes sinuate, often narrowly revolute, apex generally acuminate; lateral veins very
slender (densely parallel in Cotylelobium, Dryobalanops and some Hopea), often with
axillary domatia; intercostal venation scalariform or reticulate. Inflorescences generally
paniculate, sometimes racemose or rarely subcymose, terminal, axillary or sometimes
ramiflorous, furnished with paired modified stipules at nodes, generally regularly 1–2(–3)
branched, the branchlets bearing distichous or secund flowers. Flowers bisexual, radially
symmetrical, scented, nodding; calyx persistent, 5merous, sepals free and quincuncial
imbricate or united at base with valvate lobes, with 2–3(–5) (outer if imbricate) lobes
enclosing those remaining and generally greatly expanding and spatulate and lightly
twisted in fruit; corolla 5merous, petals contorted, weakly connate at base or free, almost
always partially unicellularhairy outside; stamens 5 to more than 100, 1–3verticillate or
irregular, hypogynous or subperigynous, centrifugal, filaments generally compressed at
base, tapering and filiform distally, or filiform throughout, frequently connate to petals,
anthers erect, 2loculed with (2–)4 pollen sacs, introrse or latrorse, connective with more or
less prominent appendage, variously short or long, aristate or stout, erect or reflexed; ovary
generally superior, sometimes semiinferior, (2–)3(–5)loculed, style generally columnar,
often thickened at base into a stylopodium, stigma generally obscure, more or less distinctly
trifid; ovules 2(–many) per locule, axile, pendulous or laterally anatropous, bitegmatic with
ventral raphe and superior micropyle. Fruits indehiscent 1(–many)seeded nuts with woody
pericarp and persistent calyx appressed to the base; calyx lobes more or less expanding into
spatulate wings; cotyledons generally fleshy, sometimes laminar, entire or variously lobed,
frequently unequal, one more or less enclosing the other, both enclosing the radicle.
Germination epigeal or hypogeal, the pericarp splitting loculicidally or irregularly.
63
TREE FLORA OF SABAH AND SARAWAK VOL. 5 (2004)
Distribution. Pantropical, with about 500 species in 3 subfamilies and 17 genera. In Sabah
and Sarawak the family is represented by 9 genera with 267 species.
Ecology. In Sabah and Sarawak, occurring especially in mixed and upper dipterocarp
forests but also in other nonsaline habitats, at altitudes to 1800 m. Rare in secondary forest
following clearing.
Notes. The resinous smell of fresh wood and bark, the twig with stipular scars, and the
stipulate leaf with simple entiremargined blade terminating abruptly at the frequently
geniculate petiole, serves to distinguish a dipterocarp at any stage in the field.
The Dipterocarpaceae have been regarded as allied to Guttiferae, Theaceae and Order
Malvales, especially the Tiliaceae. Ashton (1982) considered the family to be allied to
Sarcolaenaceae and Tiliaceae, and to be within Order Malvales. Molecular evidence
(Alverson et al., Am. J. Bot. 85 (1998) 876) indicates Dipterocarpaceae to share a clade
allied to but not within core Malvales, with Sarcolaenaceae (trees of Madagascar and
formerly Africa) and Cistaceae which are temperate shrubs. Molecular evidence (Alverson
et al., op. cit.; Dayanandan et al., Am. J. Bot. 86 (1999) 1182) supports the close affinity of
the three putative subfamilies of the Dipterocarpaceae.
Maguire & Ashton (Taxon 26 (1977) 341) subdivided the Dipterocarpaceae into three
subfamilies, which can be distinguished as follows:
1. Anthers basifixed; sepals, if amplicate, then generally conspicuously unequal with 2–3
becoming winglike in fruit; ovary (2–)3loculed, each locule with 2 ovules. Wood,
leaves and ovary with resin or secretory ducts; woodrays multiseriate. Tropical Asia
and Malesia.……subfam. Dipterocarpoideae
Anthers basiversatile; sepals equally accrescent, papyraceous; ovary (2–)3–4(–5)
loculed, each locule with 2–4 ovules. Wood, ovary and, commonly, leaves without
resin or secretory ducts……2
2. Petals longer than sepals; anthers little or deeply basiversatile, connective little or
moderately projected as an apical appendage; ovary 3(–5)loculed, each locule with 2
ovules. Woodrays predominantly uniseriate. Tropical Africa and Madagascar…………
subfam. Monotoideae
Petals shorter than sepals; anthers deeply basiversatile, connective conspicuously
projected as an apical appendage; ovary 4(–5)loculed, each locule with 4 ovules.
Woodrays predominantly biseriate. Tropical America………subfam. Pakaraimoideae
Phytogeography. The subfam. Dipterocarpoideae comprise 13 genera, with c. 475 species,

confined to the Asian tropics between the Seychelles Is. and S Asia to New Guinea but
absent in the Lesser Sunda Is. east of Sumbawa. Most genera and species occur in the W
Malesia: Sumatra, Peninsular Malaysia, Singapore, Java, Borneo, the Philippines and
islands in between. Sabah and Sarawak, which have 267 species in 9 genera, is the richest
area in the world for dipterocarp species.
64
DIPTEROCARPACEAE (ASHTON)
Within NW Borneo, dipterocarps manifest rather distinct geographical provinces, partially

irrespective of the geographical range of their specific habitats. One distinct province is
north of a line from Pontianak following the Sg. Kapuas to the Kapuas Lakes in Kalimantan,
then across the Sarawak lowlands to Sipitang District in SW Sabah, with a few entities
occurring north to Papar. Species confined to this province, yet not endemic in Borneo,
indicate this province to be the Borneo extension of Corner’s Riau Pocket. Within it, there is
a distinct barrier at the Lupar valley, including several Peninsular Malaysian/Sumatran
elements west of it, as well as some endemics. It is noteworthy that the boundary of this
province coincides with the extent of major areas of peat swamp, kerangas, and mixed
dipterocarp forest on deep yellow humult sandy soils; several of these soils specialists
extend to E Sabah, mainly on ultrabasic substrates. A second mainly lowland province
occurs from Sandakan district in E Sabah southwards, with no distinct southern boundary
but gradual attrition of species southwards to Balikpapan and beyond though many only
reach Tidung and Berau in NE Kalimantan. This province includes some Philippine species,
some of which extend westwards to NE Sarawak and Brunei. Most species confined to
Borneo to this province occur in mixed dipterocarp forest on clay soils. A third province,
overlapping with the other two, contains species of the central Borneo uplands. Those
species of low altitudes are centred, in Malesia, around the Rajang hinterland, extending
across to Ulu Barito, Kayan, and sometimes the poorly explored Kapuas hinterland. Those
of the upper dipterocarp forest are mostly as yet unrecorded from Kalimantan and there is
increasing endemism towards the northeast, peaking around Mt. Kinabalu.
We have relied, for our distribution and ecological records, in this account on the extensive
data cited in the unpublished report of Brunig (1966)* the identifications for which I was
responsible; Anderson’s (1963) Checklist for the peat swamps, and my own data from 105
sample plots, and the 52 ha plot in Lambir Hills NP. In mixed dipterocarp from Miri to
Kuching districts.
Ecology. Members of the subfamily Dipterocarpoideae are overwhelmingly concentrated in

aseasonally wet lowland evergreen forests on welldrained soils, at altitudes below 800 m
and with declining numbers above 400 m. There, the subfamily is characterised by supra
annual mass flowering especially among canopy species, mast fruiting synchronised at
subfamily level, and high survival of germinants: forests most dense in dipterocarp species
have abundant saplings at all times. Dipterocarp species are either late successional, their
juveniles with relatively high growth rates and low shade tolerance; or climax, their
juveniles having relatively low maximum growth rates but high shade tolerance.
Dipterocarps are most rich in species in forests with high canopy heterogeneity imparted by
frequent small to moderatenot largegap formation. Dipterocarps are ectotrophic
mycorrhizal: they are most speciesrich on yellow soils with shallow acid raw humus, and
their juveniles are easily eliminated by soil surface disturbance; dipterocarps return to
abandoned ladang slowly, in part owing to their limited dispersal; but they may not return
for centuries on humic soils whose surface has been removed or destroyed by machinery,
burning or cultivation. Nevertheless, the fast growth rates and superior timber quality of
many species, combined with their abundant regeneration, makes them the easiest major
rain forest timber family, worldwide, to be sustainably managed for timber. Most
dipterocarps are highly habitat specific: local conservation planning for dipterocarp species
must first conserve adequate area of each of the main habitats in which they occur.
*Brunig, E.F. 1966. Der Heidewald von Sarawak und Brunei—eine Studie seiner
Vegetation und Ökologie. 2 Beschreibung der Arten. Pp. 41–73.
65
Particularly, species are generally highly soilspecific, and appropriate choice of species is
essential for plantation. Many species appear to be able to occur, occasionally, to the
altitudinal limits of lowland tropical soils, which is generally at 700–1200 m altitude; those
that are confined to low altitudes appear to be on soils similarly constrained. The subfamily
as a whole reaches its highest biomass, and stand density on yellow and red lowland soils,
where the major division in forest species composition correlates with the presence or
absence of surface acid raw humus. On these yellow/red soils, dipterocarps comprise more
than half the aboveground tree biomass, and more than 70% of the emergent individuals;
they are less well represented by species which reach maturity in the main canopy or
subcanopy though these ‘mixed dipterocarp forests’ support many juvenile dipterocarps of
all sizes, especially saplings. In this treatment, commonness is estimated in relation to
ecology, for forest prior to logging.
Dipterocarps are insect pollinated; species within each major taxon share the same insect
pollinators. Flowering within a major taxon, especially in the sections of Shorea, is
sequential within one habitat and species association, with overlap in flowering between
successive species. Dipterocarps are selfcompatible, though available evidence suggests
that outcrossing is high in nature, and that progeny from outcrossing have higher
survivorship than those originating from selfing. Apomixis through adventive embryony
appears to be important in some species. Dipterocarp seeds lack dormancy, the species
populations therefore being dependant on an abundant population of juveniles. Fruits are
primarily dispersed by gyration occasioned by the twisted winglike fruit sepals, though
many appear to have no means of dispersal. Terrestrial mammals, especially rodents, may
inadvertently disperse seed for short distances, and scatterhoarding occurs. A few species
are waterdispersed. Fruit dispersal is, therefore, mostly very limited in the aseasonal wet
tropics except during exceptional wind storms. Dipterocarps may take centuries, or longer,
to disperse back into extensive cleared areas.
Dipterocarps are, generally, a poor source of food for larger animals; their resinous parts are
astringent with tannin and difficult to digest, while the lipidrich fruits are only produced at
long intervals when their superabundance far exceeds the capacity of animals to consume
them; this thereby permits their copious regeneration. Supraannual mast fruiting appears to
explain both the success of dipterocarps in western Malesia, and the relatively low animal
carrying capacity of vertebrates there compared with evergreen lowland tropical forests in
Africa and America. Recently, it has been discovered that wild pigs (Sus scrofa) in Pasoh
FR, Peninsular Malaysia, preferentially harvest dipterocarp saplings to build their nests; the
increase in food availability for pigs, provided by fallen fruit in oil palm plantations adjacent
to mixed dipterocarp forest, is thereby leading to dramatic declines in dipterocarp saplings,
and may lead to a declining role of the subfamily at least near the edge of the forest.
Uses. Dipterocarps have had a wealth of traditional uses (PROSEA 5, 1 (1993) 94, 139, 166,
186, 238, 299, 325, 384, 458 & 461), few of which have survived rural economic change.
The earliest recorded use was for camphor, from Dryobalanops aromatica, which was
exported in medaeval times to the Middle East, where it has been said it was worth its
weight in gold as a base for incense. Iban girls formerly used camphor resin in a similar
way, melting the camphor then infusing it with other scents; it was then used as a solid
perfume and hidden in their pillows. There would appear to be opportunities for commercial
development of this attractive fashion for a wider market.
66
The viscid resin of Dipterocarpus was used, eventually commercially, as a basis for paints
and for caulking wooden boats, particularly in seasonal continental Asia where several
species produce copious resin and are tapped with the aid of fire. This practise continues in
the poorer rural economies. The crystalline resin of some species of Hopea, known as mata
kucing, was used as a base for medicines, and for varnishes. A charcoal, prepared from resin
of Shorea sect. Richetioides, was formerly used for blackening teeth, a custom then among
Lun Dayuh and Brunei women.
The abundance of astringent compounds in dipterocarp tissues, specifically the inner bark,
was availed of as laru: macerated bark introduced to rice and coconut beer to arrest further
fermentation; it was also used to prevent frothing during boiling of palm syrup during sugar
manufacture.
The fatrich cotyledons have formerly been an important source of famine food throughout
Asia, and nuts of certain Shorea are still collected in Sri Lanka for preparation of
sweetmeats, and in Borneo as a source of cooking fat with a high melting point. The large
nuts of some species of Shorea sect. Pachycarpae and also sect. Brachypterae, sect. Mutica
and sect. Shorea are particularly favoured, and are exported during mast years for use in
chocolates, lipstick and other industrial products.
But since the early seventies, dipterocarps have provided the bulk of hardwoods traded on
international markets; the light hardwood Shorea timbers being particularly in demand
though the heavy hardwood selangan batu is marketed for decking, garden furniture and
other purposes where a high quality weather resistant wood is required. Although timber
harvesting can be managed sustainably (but almost certainly not also sustaining
biodiversity), this has not been done because it involves additional costs and reduces profits.
In particular, careless and inappropriate use of machinery, and especially reentry for further
logging while the residual stand is developing, has led to soils disturbance and damage to
the original stand on such a scale that it is unlikely that economically merchantable stands of
dipterocarps will return for at least one, and possibly several centuries. Although this
practise is logical financially to the concession holder, it has economically impoverished the
nation by degrading its most valuable sustainable resource.
By virtue of their commanding role in the structure of the lowland and hill forests of Sabah
and Sarawak, dipterocarps possess unique economic service values, esthetically including
for tourism, and in the conservation of the overall biodiversity of these ecosystems which
are the most speciesrich forests in the Old World, being matched only by some in the
headwaters of the Amazon River. Their value in sequestration of atmospheric carbon is
almost certainly greater than any other tropical plant family. The harvesting of the
dipterocarp trees for timber dramatically affects the demography, hence interdependences,
of the other plants and animals of the forest, and extinctions can be predicted. This are
major reasons why the forests of the national parks must remain sacrosanct.
Conservation status. I have endeavoured, on available evidence and personal experience,

to predict the degree of endangerment of the dipterocarp species in our region. My criteria
require explanation. The International Union for the Conservation of Nature (IUCN) has
drawn up criteria for making standard comparable assessments of conservation status for
organisms in general, plants as well as animals. Their criteria are based, however, mainly on
animal population characteristics. Most animals are less physicalhabitatspecific than
plants, and their populations therefore generally extend more continuously over large areas.
Animals also have shorter lifespans than trees. IUCN has placed high emphasis, therefore,
67
on the amount by which habitat has been reduced in area over the last three generations of a
species. Were this criterion literally applied to dipterocarps, all our species would be judged
to be at some level of endangerment!. Dipterocarp populations, by being highly habitat
specific, are already often fragmented in nature into an archipelago of habitatislands. It is,
therefore, the number of these islands that survive forest harvesting or conversion that
matters to them, rather than the absolute decline in primary (or logged) forest area. A further
important rule of thumb, which can reasonably applied to both plants and animals, is the
minimum acceptable size of populations of mature reproductive individuals that is needed to
sustain the genetic diversity found in nature through outbreeding. It is generally assumed
that a minimum of 200 mature individuals, in a given area/habitat, is needed, that is a
minimum of 200 within the foraging range of the pollinating insect in the dipterocarp case.
It is thus obvious that logging has a major impact on the distribution and frequency of
adequate dipterocarp breeding populations. It can reasonably be argued that the regeneration
which survived from the primary forest through the logging process will in due course grow
to become new populations of acceptable density. We hope that this may be the case, but
fear that soil disturbance, uniformly low light conditions beneath the later regrowth, and the
slow growth rates of climax species in relation to the length of the felling cycle, may all
have adverse effects, particularly on the soil specialists and other less common species. I
have therefore placed high emphasis on the accessibility and suitability of species’ habitats
for conversion to agriculture, and their occurrence in the national parks system, when
assigning a level of threat to a species’ survival within Sabah and Sarawak. The generally
lower level of endangerement of species in Sabah and Sarawak when compared to their
global distribution (cf. Oldfield et al. 1998. The World List of Threatened Trees, WCMC &
IUCN) testifies largerly to their generally good conservation in the legislated Sabah and
Sarawak park systems, which have been largerly protected from logging. In logged forests,
surviving juveniles of the more lightdemanding species may be expected to survive to
maturity, as may those of understorey species; but the shadetolerant, generally heavy
hardwood species have an uncertain future there.
Key to genera
(based on flowering/fruiting specimens)
1. Fruit calyx lobes valvate, not incrassate, at base. Chromosome number x = 11 (tribe
Dipterocarpeae)……………………………………………………………………...…2
Fruit calyx lobes at least slightly imbricate at base (often becoming fully valvate in
Dryobalanops and Parashorea), with a distinct incrassate expanded spoonshaped base
(united into a cup in Dryobalanops). Chromosome number x = 7 (tribe Shoreae)….…6
2. Base of fruit calyx united into a tube enclosing at least the basal half of the nut…….…3
Fruit sepals free to the base, or united into a shallow cup enclosing less than one third
of the nut……………………………………………………………………………...…4
3. Connectival appendage of stamen stout, tapering, at most 1½x the length of anthers.
Nut free within calyx tube. Epidermis lacking peltate scales. Leaf bud prominent;
stipules large, amplexicaul; leaf aestivation plicate; lateral veins not uniting into an
intramarginal vein………………………………………………….…3. Dipterocarpus
Connectival appendage of stamen slender, at least twice the length of anther. Nut
adnate to calyx tube. Epidermis in part furnished with peltate scales. Leaf bud small;
68
stipules linear, with short scars; leaf aestivation not plicate; lateral veins joining distally
into a prominent looped intramarginal vein.…………………………..…1. Anisoptera
4. Stamens at least 25; connectival appendage at least twice the length of anther. Nut
triangular in crosssection………………………………………………………8. Upuna
Stamens at most 15; connectival appendage at most 1½x the length of anther. Nut round
in crosssection……………………………………………………………………….…5
5. Anthers linearoblong, setose along lateral margins; style at least twice the length of
ovary; stigma small, trifid. Leaf lateral veins densely parallel, unequal, slender and
hardly raised, main veins joining distally uniting into a looped intramarginal
vein………………………………………………………………..…... 2. Cotylelobium
Anthers broadly oblong to subglobose, glabrous; style less than twice the length of
ovary; stigma conical, prominent. Leaf lateral veins wellspaced, usually distinctly
raised below, with short or no intermediate veins, not or obscurely distally joined into
an intramarginal vein………………………………………………………..…9. Vatica
6. Fruit calyx lobes all equal, all spatulate, winglike, or all short. Leaf lateral veins very
slender, dense, parallel, connected by a vein just within the margin…..4. Dryobalanops
Fruit calyx lobes unequal, spatulate (except in Shorea isoptera), or all short. Leaf lateral
veins, if slender and dense, not joining into an intramarginal vein…………………..…7
7. Fruit calyx lobe base narrow, hardly imbricate; nut verrucoselenticellate, ripening
globose to ellipsoid…………………………………………………….…6. Parashorea
Fruit calyx lobe base broad, distinctly imbricate; nut not verrucoselenticellate, ripening
not so………………………………………………………………………………….…8
8. Flower calyx lobes with 2 outer ones thicker, more acuminate than the 3 inner ones, 2
becoming winglike, spatulate in fruit, or all 5 short………………………..…5. Hopea
Flower calyx lobes with 3 outer ones thicker, more prominently acuminate than the 2
inner ones, 3 becoming winglike, spatulate in fruit or all 5 short………….…7. Shorea
Field key to the genera and main field groups

of Hopea and Shorea
1. Leaf lateral veins curving round towards the margin and anastomosing to form a
distinctly looped intramarginal vein………………………………………………….…2
Leaf lateral veins either without or with indistinct intramarginal vein, or intramarginal
vein straight…………………………………………………………………………..…3
2. Bark surface remaining smooth at first, eventually irregularly flaky and becoming
scrollmarked; inner bark homogeneous pale brown. Leaf intercostal venation obscure;
main lateral veins very slender, hardly raised below, with many variably shorter
intermediates; petiole not geniculate; leaf blade not lepidote below…...2. Cotylelobium
Bark surface vertically cracked, oblongflaky; inner bark distinctly tangentially
laminated. Leaf intercostal venation distinct, wellspaced, scalariform or reticulate;
main lateral veins distinctly raised below; petiole geniculate; leaf blade peltate lepidote
below………………………………………………………………………1. Anisoptera
69
3. Leaf lateral veins densely parallel, ending abruptly in a more or less straight intra
marginal vein close within margin………………………………….…4. Dryobalanops
Leaf lateral veins pinnate or, if densely parallel, then not ending in intramarginal
vein………………………………………………………………………………………4
4. Bark with large pale lenticels. Leaf lateral veins straight, hardly arched except towards
the margin; leaf blade usually distinctly corrugated with distinct ridges between the
lateral veins…………………………………………………………………………...…5
Bark with small lenticels hardly paler than bark surface. Leaf lateral veins arched; leaf
blade flat or bullate but not corrugated with distinct ridges………………………….…6
5. Bark surface yellowish to orange or rustbrown, rarely pinkish brown, becoming more
or less irregularly flaky; lenticels scattered or in clusters, pale yellowish brown.
Terminal (leaf) buds prominent; stipules large, scars amplexicaul; leaf blade coriaceous,
never silvery below; petiole geniculate…………………………….…3. Dipterocarpus
Bark surface mauve to dark purple or tawnybrown, closely shallowly fissured and
narrowly oblongflaky; lenticels initially in vertical lines, eventually following the
bottom of fissures and ridges of buttresses, whitish. Terminal buds and stipules
generally small (except in Parashorea macrophylla), stipular scars short (except in P.
macrophylla and P. malaanonan); leaf blade usually silvery below, particularly in
juveniles……………………………………………………………….…6. Parashorea
6. Lower leaf surface, twig, lateral veins and petiole densely whitish felty with brown
tomentum; bark dark chocolatebrown, closely vertically cracked and flaky; buttresses
stout, in groups…………………………………………………………………8. Upuna
Leaf lower surface not whitish felty or, if so, bark tawnybrown, and buttresses not
grouped…………………………………………………………………………………7
7. Buttresses low, stout; bark surface smooth except in very large trees; inner bark
homogeneous, pale creambrown; wood dense, fine textured, ray ends not glistening on
tangential surface. Leaf intercostal venation generally reticulate but not drying darker
than the leaf blade…………………………………………………………..…9. Vatica
Bark surface, if smooth, then tree with prominent buttresses; inner bark fibrous; wood
dense, ray ends glistening on tangential surface. Leaf intercostal venation scalariform or
if reticulate, drying much darker than the leaf blade……………………………………8
8. Leaf lateral veins densely parallel, variously unequal to subequal...........................…

………………………………………………………...5. Hopea (in part; many species)
Leaf lateral veins strictly pinnate, sometimes with short intermediates only………...…9
9. Wood dense, hard, ray ends not glistening on tangential surface……………………...10

Wood not dense, not hard, ray ends glistening on tangential surface…………………11
10. Inner bark yellowish brown or, if reddish, closely eventextured, not fibrous…………...
………………7. Shorea (in part; selangan batu) and 5. Hopea (in part; some species)
Inner bark fibrous, reddish orange, pale at cambium………………………………...…
………………………………….7. Shorea (in part; S. inaequilateralis and S. kunstleri)
11. Inner bark distinctly tangentially laminated; leaf midrib obscure, sunken above………..
………………………………………………………..7. Shorea (in part; white meranti)
70
Inner bark not distinctly laminated; leaf midrib usually evident, if furrowed,
occasionally obscure above……………………………………………………………12
12. Inner bark greenish yellow; dammar blackish; bark tawny flaky (except in S.
acuminatissima). Leaf intercostal venation subscalariform, or reticulate and drying
darker than the blade……………………………..…7. Shorea (in part; yellow meranti)
Inner bark yellowish to reddishbrown; dammar whitish or yellow or, if blackish, then
inner bark pinkish; bark otherwise. Leaf intercostal venation scalariform……………13
13. Bark becoming blackish brown, flaky, with prominent white dammar incrustations;
inner bark and sapwood homogeneous, cream…….…5. Hopea (in part; e.g., H. sangal)
Bark, if blackish, with yellow dammar incrustations, inner bark and sapwood not cream
or homogeneous…………………………………………………………………….…14
14. Buttresses narrow, frequently with flying buttresses and stilt roots; bark surface
chocolatebrown and greymottled, remaining smooth or patchily cracked and thinly
flaky………………………………………………………………...…5. Hopea (in part)
Buttresses stout, rarely stilted; bark, if remaining smooth, pale not chocolatebrown…...
…………………………………………………….7. Shorea (in part; most red meranti)
1. ANISOPTERA Korth.
(Greek, anisos = unequal; pteron = wing; the unequal fruit calyx lobes)
kelapok (Iban), mersawa (Malay), pengiran (preferred name in Sabah)
Kruidk. (1841) 65; King, J. As. Soc. Beng. 62, 2 (1893) 100; Ridley, FMP 1 (1922) 218; Merrill, EB
(1921) 400; Slooten, Bull. Jard. Bot. Buitenz. 3, 8 (1926) 3; Symington, Gard. Bull. S. S. 8 (1934) 1,
Malay. For. Rec. 16 (1943) 199; Masamune, EPB (1942) 482; Browne, FTSB (1955) 92; Ashton,
Gard. Bull. Sing. 20 (1963) 230, MDB (1964) 9, MDBS (1968) 3, FM 1, 9 (1982) 327; Backer &
Bakhuizen f., FJ 1 (1964) 329; Meijer & Wood, Sabah For. Rec. 5 (1964) 292; Burgess, TBS (1966)
93; Anderson, CLTS (1980) 110; PROSEA 5, 1 (1993) 94; Kessler & Sidiyasa, TBSAEK (1994) 89;
Coode et al. (eds.), CLBD (1996) 66; Newman et al., MDFBLHW (1996) 57. Synonyms:
Antherotriche Turcz., Bull. Soc. Nat. Mosc. 2 (1846) 505; Scaphula Parker in Fedde, Rep. 30 (1932)
326.
Emergent trees with stout, short or tall and straight buttresses; crown irregularly
hemispherical, rather narrow, often diffuse, with a few large twisted ascending branches.
Bark becoming shallowly irregularsection fissured and thinly flaky; inner bark
prominently tangentially laminated, thick, fibrous. Sapwood pale yellowish white,
siliceous; dammar opaque, yellow or white, occurring as smearlike exudates on bole.
Indumentum on young parts and leaf undersurface in part consists of dense unicellular
peltate scales. Twigs more or less prominently ribbed on drying. Leaf buds ovoid, acute,
small, c. 5 × 3 mm, somewhat compressed. Stipules relatively large, lineartapering, falling
early and leaving short scars. Leaves aestivation not plicate; thinly to thickly coriaceous,
peltatelepidote below; blade oblongobovate, base usually obtuse, equal, apex shortly
acuminate; midrib sunken above; main lateral veins distinctly raised below, curving toward
margins and anastomosing to form a distinctly looped intramarginal vein; intercostal
venation distinct, laxly scalariform, or subreticulate, or reticulate, prominent and raised
below; petiole relatively long, prominently geniculate. Inflorescences racemose, terminal or
71
axillary, pendent; rachis lax, long, manyflowered; bracteoles small, caducous. Flowers
distichous, pedicellate; buds lanceolate or globose (in A. laevis); sepals imbricate to valvate
(in A. laevis), the 2 outer lobes obtuse, thicker than the 3 acute inner ones; petals linear;
stamens c. 15 (in A. laevis) or more than 30, glabrous, filaments short, tapering, connate at
base, anthers latrorse, connectival appendage slender, at least twice the length of anther;
ovary semiinferior, with distinct stylopodium, style long or short, stigma distinctly trifid,
rarely obscure (in A. laevis). Fruits: calyx lobes valvate, with a subglobose base enclosing
and adnate to the nut, with 2 long narrowly spatulate untwisted 3veined lobes and 3 short
lobes. Nuts imbedded in the calyx tube but for the apex, with broad discoid (in A. laevis) or
cylindrical stylopodium and persistent columnar (in A. laevis) or short style; cotyledons
unequal; first leaves paired with interpetiolar stipules, or in a whorl of 4 without stipules;
germination epigeal, the pericarp breaking open irregularly.
Distribution. Ten species, ranging from Chittagong (Bangladesh) to S IndoChina, east and
southeastwards to New Guinea. Five species in Borneo, all of which occur in Sabah and
Sarawak; 2 endemic in Borneo of which one endemic to Sabah, Sarawak and Brunei.
Ecology. In mixed dipterocarp, mixed peat swamp and kerangas forests, rarely at altitudes
above 1000 m. In our area generally as scattered individuals, but occasionally semi
gregarious in the seasonal tropics.
Uses. The timber is a light hardwood; it is siliceous, but makes excellent peeler for veneer.
Key to Anisoptera species

(base on flowering/fruiting specimens)
1. Flower buds subglobose; stamens c. 15; stylopodium in flower broad, discoid, glabrous;
style columnar, slender, glabrous, stigma obscure (sect. Glabrae)…………3. A. laevis
Flower buds spindleshaped or ovoidlanceolate; stamens 20–36; stylopodium in flower
as broad as ovary, ovoid, oblong to cylindrical, densely pubescent; style and stigma
short, trifid (sect. Anisoptera)……………………………………………………….…2
2. Leaf greyish brown or bright yellow lepidote below………………………………..…3

Leaf golden scaly or glabrous below.………………………………………………..…4
3. Leaf drying greyish brown above, greyish brown lepidote below; venation below and
petiole greyish brown tomentose; stylopodium cylindrical, not tapering at base………...
………………………………………………………………………………1. A. costata
Leaf drying mauvebrown above, bright yellow lepidote below; venation below densely
rufous scabridpubescent; stylopodium ovoid, tapering at both ends….2. A. grossivenia
4. Leaf golden scaly below, matt above; lateral veins slender, hardly raised below;
stamens c. 25; stylopodium truncate……………………………………4. A. marginata
Leaf not golden scaly below, shiny above; lateral veins stout, prominent below; stamens
c. 35; stylopodium tapering at apex…………………………………...…5. A. reticulata
72
Key to Anisoptera species

(based on field characters)
1. Leaf blade, at least on the veins below and petiole, variously hairy/lepidote………...…2
Leaf blade below and petiole glabrous or scaly…………………………………………4
2. Leaf blade drying mauvebrown above, rufous scabridpubescent on the veins below….
…………………………………………………………………………2. A. grossivenia
Leaf blade drying greyish brown above, shortly greyish brown lepidote/hairy below….3
3. Leaf blade sparsely scabridhairy and greyish brown lepidote below; buttresses tall,
straight……………………………………………………………………...1. A. costata
Leaf blade below sparsely tomentose on the veins only; buttresses short…..……………
…………………………………………………………………4. A. marginata (in part)
4. Leaf blade drying yellowish brown below, shiny above; lateral veins prominent below;
intercostal venation reticulate……………………………………………5. A. reticulata
Leaf blade more or less golden scaly below, not shiny above; lateral veins slender;
intercostal venation subscalariform…………………………………………………..…5
5. Leaf midrib sharp below, drying conspicuously darker than the leaf blade; twigs pale,
striated; bark surface patchily fissured and flaked………………………...…3. A. laevis
Leaf midrib rounded below, drying greyish brown; twigs uniform brown; bark closely
parallel fissured and flaked………………………………….…4. A. marginata (in part)
1. Anisoptera costata Korth.

(Latin, costatus = ribbed; the prominent venation of the leaf blade)
Kruidk. (1841) 67; Merrill op. cit. 400; Slooten op. cit. 7, Reinwardtia 2 (1952) 8; Symington op. cit.
(1934) 9, op. cit. (1943) 204; Masamune op. cit. 482; Ashton op. cit. (1964) 11, op. cit. (1968) 5,
Gard. Bull. Sing. 31 (1978) 13, op. cit. (1982) 330; Meijer & Wood op. cit. 293; Burgess op. cit. 93;
Anderson op. cit. (1980) 110; PROSEA op. cit. 99; Newman et al. op. cit. 60; Coode et al. (eds.) op.
cit. 66. Type: Korthals s.n. (= RHL Sheet No. 90214642), Borneo, S Kalimantan, G. Sakumbang,
Tanah Laut (holotype L). Synonyms: Anisoptera oblonga Dyer, Fl. Brit. Ind. 1 (1874) 301; A.
cochinchinensis Pierre in Lanessan, Pl. Util. Colon. Fr. (1886) 268; A. robusta Pierre, Fl. For. Coch. 3
(1889) t. 236; A. marginatoides F.Heim, Bot. Tidsskr. 25 (1902) 44; A. mindanensis Foxw., Philip. J.
Sci. 13 (1918) Bot. 181.
Huge tree, to 65 m tall, to 1.5 m diameter; buttresses massive, straight, to 4 m tall and
continuing as ribs up bole to 10 m. Bark greyish brown, vertically fissured and thinly
flaking; inner bark laminated cream and pale yellow; dammar secretions as greyish green
smears. Young parts more or less sparsely greyish brown hairy. Leaves thinly coriaceous,
frequently slightly bullate, sparsely scabridhairy and greyish brown lepidote below, drying
greyish brown above; blade oblong to obovate, 6–18 × 7–11 cm, base obtuse or broadly
cuneate, apex with an acumen to 0.5 cm long; lateral veins 8–22 pairs, prominent and
tomentose below; petiole 2–4 cm long, greyish brown shorthairy. Inflorescences to 20 cm
long, pubescent. Flowers: buds ovoidlanceolate, to 12 × 6 mm; corolla cream, petals
narrow; stamens c. 25, subequal, anthers oblong, tapering, connectival appendage c. twice
the length of anther, tapering; stylopodium cylindrical, somewhat tapering apically, not
tapering at base, densely pubescent; style and stigma short, trifid. Fruits: calyx tube to 1.2
73
cm diameter, puberulent; calyx lobes glabrous, 2 longer lobes spatulate, to 16 × 2 cm,

constricted to c. 5 mm at base, 3 shorter ones hastate, to 2 × 0.4 cm, hardly constricted at
base. Nut apex broadly conical, shortly pubescent; stylopodium cylindrical, c. 2.5 × 1.5mm.
Vernacular names. Sabah—pengiran kesat (preferred name). Sarawak—mersawa kesat

(preferred name).
Distribution. Coastal hills of Myanmar, S and C Thailand, Cambodia and CochinChina,

Sumatra, Peninsular Malaysia, W Java, Borneo, the Philippines (Mindanao), and Maluku. In
Borneo, scattered throughout Sabah and recorded from Keningau, Kota Marudu, Lahad
Datu, Sandakan, Semporna, and Tawau districts (e.g., FMS 41471, FMS 48764, SAN 7839,
SAN 15283, SAN 21511, and SAN 122039) and in Sarawak from Limbang district (e.g., S
32209). Also occurring in Brunei (e.g., BRUN 3387, BRUN 3389 and BRUN 5651) and
Kalimantan (e.g., bb. 31198, Kostermans 4384 and Kostermans 10577).
Ecology. Evergreen and semievergreen lowland forests of IndoBurma; in mixed

dipterocarp forests on clay ultisols in Sabah and Sarawak, at altitudes to 700 m. Uncommon,
vulnerable.
2. Anisoptera grossivenia Slooten

(Latin, grossus = an unripe fig, venius = veined; the purple lateral veins of the leaf blade)
Bull. Jard. Bot. Buitenz. 3, 16 (1940) 431; Symington op. cit. (1943) 13; Masamune op. cit. 483;
Browne op. cit. 93; Ashton op. cit. (1964) 12, op. cit. (1968) 55, op. cit. (1982) 331; Meijer & Wood
op. cit. 296; Burgess op. cit. 94; Anderson op. cit. (1980) 110; PROSEA op. cit. 100; Coode et al.
(eds.) op. cit. 66; Newman et al. op. cit. 61. Type: bb. 29708, Borneo, Kalimantan, Dusun Lands,
Pepas, Muara Teweh (holotype BO; isotypes KEP, L).
Tree to 60 m tall, to 1.5 m diameter; buttresses low, stout. Bark purplish brown, closely
fissured and flaking; inner bark pale yellow and rich orange laminated; dammar smears rich
yellow, long. Leaf venation below, petiole, stipule outside, buds, inflorescence, and twig
persistently rufous scabridpubescent. Leaves often slightly bullate, drying mauvebrown
above, brilliant yellow lepidote below (juveniles excepted); blade narrowly obovate to
oblong, 9–12 × 3–5 cm, base cuneate, margin slightly recurved, apex with narrow acumen
to 1 cm long; lateral veins 18–28 pairs, dense, arched; intercostal venation hardly raised
(juveniles excepted), subreticulate; petiole 2–2.5 cm long. Inflorescences to 20 cm long.
Flowers: buds lanceolate, to 5 × 2.5 mm long; corolla magenta in bud, yellow on opening,
petals narrow; stamens c. 36, anthers subglobose, connectival appendage c. thrice as long as
anther, slender; stylopodium ovoid, tapering at both ends, densely golden pubescent; style
and stigma short, trifid. Fruits: calyx sparsely puberulent; tube to 1.3 cm diameter; 2 longer
calyx lobes narrowly spatulate, to 20 × 2.5 cm, constricted to c. 5 mm at base, 3 shorter ones
linear, to 0.6 cm long. Nut apex obtuse, densely rufous puberulent; stylopodium oblong
ovoid, c. 8 × 2 mm.
Vernacular names. Sabah—pengiran kunyit (preferred name). Sarawak—mersawa kunyit

(preferred name).
74
Distribution. Endemic in Borneo. Widespread throughout the lowlands of Sabah and

recorded from Beaufort, Sandakan and Sipitang districts (e.g., KEP 80257, SAN 15058 and
SAN 15159) and in Sarawak from Belaga, Kuching, Lawas, Lundu, and Miri districts (e.g., S
8915, S 9617, S 10285, S 22312, and S 46495). Also occurring in Brunei (e.g., BRUN 3028,
BRUN 3055, BRUN 3286, S 1644, and S 5780) and C and NE Kalimantan (e.g., bb. 22874,
bb. 29605, Jarvie & Ruskandi 5896, and Kostermans 9113) where its habitat occurs.
Ecology. In mixed dipterocarp forest and its ecotone to kerangas forest, on shallow and
deep yellow sandy soils, at altitudes below 400 m. Well represented in Bako and Lambir
NPs and not currently endangered.
3. Anisoptera laevis Ridl. Fig. 1.

(Latin, levis = smooth; the leaf blade)
FMP 1 (1922) 219; Symington op. cit. (1943) 205; Ashton op. cit. (1964) 13, op. cit. (1968) 5, op. cit.
(1982) 335; Meijer & Wood op. cit. 298; Burgess op. cit. 94; Anderson op. cit. (1980) 100; PROSEA
op. cit. 101; Coode et al. (eds.) op. cit. 66; Newman et al. op. cit. 62. Type: Ridley SFN 6886,
Singapore Botanic Gardens (holotype SING; isotype KEP).
Vast emergent tree, to 65 m tall, to 2 m diameter; buttresses to 15 m tall, 4 m wide, stout.

Bark pale greyish buff, patchily fissured and flaked; inner bark pale brownish laminated,
rather thin; dammar in greyish brown smears, often not present. Young parts pale brown to
rust caducous lepidote, epilose (without hairs). Twigs pale, striated. Leaves flat, golden
scaly below (mature trees), not shiny above, drying pale greenish brown below; blade
oblongobovate, 7–11 × 3–4 cm, base obtuse or broadly cuneate, apex with an acumen to 1
cm long; midrib sharp below, drying conspicuously darker than the blade; lateral veins 10–
14 pairs, slender; intercostal venation subscalariform; petiole 1.5–2 cm long, minutely
scaly. Inflorescences to 12 cm long, pale brown caducous puberulent. Flowers: buds
subglobose; petals small, ellipticoblong, pale yellow; stamens c. 15, anther narrowly
oblong, connectival appendage short, erect; stylopodium discoid, glabrous; style columnar,
glabrous, stigma obscurely 3lobed. Fruits: calyx glabrous; tube to 1.5 cm diameter; 2
longer lobes spatulate, to 15 × 1.5 cm, 3 shorter ones hastate, linear, to 1.5 × 0.3 cm, slightly
recurved. Nut apex discoid, puberulent; style remnant to 8 mm long, prominent, filiform.
Vernacular names. Sabah—pengiran durian (preferred name). Sarawak—mersawa durian

(preferred name).
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo. In Borneo, known in

Sabah from Beaufort, Sipitang and Ranau districts (e.g., SAN 15083 and SAN 16944) and in
Sarawak from Bau, Kapit, Lawas, Limbang and Tatau districts (e.g., S 1109, S 17756, S
22279, S 22305, S 24810, and S 32297). Also occurring in Brunei (e.g., BRUN 3053, BRUN
3192, BRUN 5251, FMS 35455, and S 5780).
Ecology. Widespread but uncommon in mixed dipterocarp forest on clay loam soils derived
from shale and basic to intermediate volcanic rocks. Recorded from Mulu NP but
vulnerable.
75
4. Anisoptera marginata Korth. Fig. 2G–H.

(Latin, marginatus = bordered; the distinct intramarginal vein of the leaf blade)
Kruidk. (1841) 66; Merrill op. cit. 401; Slooten op. cit. 5; Masamune op. cit. 483; Symington op. cit.
(1943) 206; Anderson, Gard. Bull. Sing. 20 (1963) 157, op. cit. (1980) 110; Ashton op. cit. (1964) 14,
op. cit. (1968) 5, op. cit. (1982) 328; Meijer & Wood op. cit. 298; Burgess op. cit. 93; PROSEA op.
cit. 101; Coode et al. (eds.) op. cit. 66; Newman et al. op. cit. 63. Type: Korthals s.n. (= RHL Sheet
No. 90214648), Borneo, Kalimantan, G. Pamaton (holotype L). Synonym: Anisoptera grandiflora
Brandis, J. Linn. Soc. Bot. 31 (1895) 43.
Tree to 45 m tall, to 1.5 m diameter; buttresses low, rounded. Bark greyish brown,
shallowly fissured and flaking; inner bark brown and cream laminated, thick; dammar as
greyish cream smears. Twigs uniformly brown. Young parts greyish brown pubescent.
Leaves matt above, golden to greyish brown lepidote below or sparsely tomentose on the
lateral veins only, drying greyish to yellowish brown, shallowly bullate in juveniles; blade
oblongobovate, 7–10 × 3.5–4.5 cm, base obtuse or broadly cuneate, apex with an acumen
to 0.6 cm long; midrib round below, drying the same colour as the blade; lateral veins 10–
14 pairs, slender, hardly raised below; intercostal venation subscalariform; petiole 1.5–2
cm long. Inflorescences to 14 cm long. Flowers: buds lanceolate, to 8 × 3.5 mm; petals
broadly elliptic, pale yellow; stamens c. 25, anther oblong, connectival appendage c. twice
the length of anther, slender; ovary and stylopodium puberulent, stylopodium cylindrical,
truncate, style and stigma short, trifid. Fruits: calyx tube to 1.3 cm diameter; 2 longer calyx
lobes spatulate, to 12 × 2 cm, 3 shorter ones linear, to 2 × 0.3 cm. Nut apex almost flat;
stylopodium oblong, c. 3 × 1.5 mm.
Vernacular names. Sabah—pengiran kerangas (preferred name). Sarawak—mersawa paya

(preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo (widespread). In Sabah known

from Beaufort, Sandakan and Tawau districts (e.g., D.D. Wood 2386, SAN 18461 and SAN
31588) and in Sarawak from Belaga, Betong, Kapit, Lawas, Limbang, Lundu, Marudi, Miri,
Sibu, Sri Aman, and Tatau districts (e.g., S 8038, S 18862, S 21177, S 26550, and S 69174).
Also occurring in Brunei (e.g., FMS 30418, Niga NN 153 and Wong s.n.) and Kalimantan
(e.g., bb. 2624, bb. 9431, Endert 5056, Kostermans 7144, and Kostermans 8896).
Ecology. In mixed peat swamp forest both in the coastal swamps and, rarely, inland valleys;
and in poorly drained kerangas forest, sandy hill crests, and in Sabah on ultrabasic rocks, at
altitudes to 850 m, as scattered trees or locally frequent. Recorded from Mulu NP;
vulnerable, possibly endangered.
5. Anisoptera reticulata P.S.Ashton Fig. 2A–F, Plates 2A–B.

(Latin, reticulatus = netted; the intercostal venation of leaf blade)
Gard. Bull. Sing. 22 (1967) 260, op. cit. (1968) 5, op. cit. (1982) 333; Coode et al. (eds.) op. cit. 67;
Newman et al. op. cit. 64. Type: G.H.S. Wood SAN 15172, Borneo, Sabah, Ulu Sipitang (holotype K;
isotypes KEP, L, SAN).
Large tree, to 65 m tall, to 2 m diameter, with short rounded buttresses. Bark pale yellowish
brown, vertically fissured and flaky. Young parts at first, inflorescence, flower bud, ovary,
stylopodium, and fruit calyx golden brown puberulent; parts otherwise glabrous. Leaves
thickly coriaceous, drying yellowish brown below, shiny above, glabrous; blade elliptic
76
Fig. 1. Anisoptera laevis. A, flowering leafy twig; B, detail of venation and indumentum on
lower leaf surface; C, flower bud; D, abaxial view of larger sepal; E, abaxial view of smaller
sepal; F, adaxial view of petal; G, flower bud with petals and stamens removed; H, flower
bud with sepals, petals removed; I, adaxial view of stamen; J, abaxial view of stamen; K,
longitudinal section of gynoecium; L, fruit; M, longitudinal section of fruit. (A–K from SAN
15083, L–M from Ridley 6886.)
77
Fig. 2. Anisoptera reticulata (A–F) and A. marginata (G–H). A, fruiting leafy twig; B,
flower bud; C, open flower; D, longitudinal section of open flower; E, adaxial view of
stamen; F, abaxial view of stamen; G, mature fruit; H, longitudinal section of mature fruit.
(A from SAN 92981, B–F from FRI 41363, G–H from Niga NN 153.)
78
obovate, 4.5–13 × 2.2–5 cm, base broadly cuneate, apex shortly abruptly acuminate; midrib
flat or slightly sunken above, prominent below; lateral veins 9–14 pairs, stout and elevated
below; intercostal venation reticulate; petiole 1.5–3.5 cm long, drying black. Inflorescences
to 6 cm long. Flowers: buds spindleshaped; corolla cream; stamens c. 35, subequal, anthers
oblong, tapering, connectival appendage filiform; stylopodium oblong, tapering at apex,
densely puberulent; style and stigma short, trifid. Fruits: calyx tube to 2 cm diameter; 2
longer calyx lobes loratespatulate, to 13 × 3 cm, constricted to c. 9 mm at base, 3 shorter
ones linearlanceolate, to 2 × 0.3 cm; stylopodium remnant oblong, to 4 × 3 mm, obtuse.
Vernacular name. Sabah—pengiran gajah (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from Kinabatangan, Labuk Sugut,

Ranau, Sandakan, and Sipitang districts (e.g., FRI 41363, SAN 15172, SAN 92981, SAN
75357, and SAN 100178) and in Sarawak from Belaga, Lawas and Miri districts (e.g., S
1503, S 66963, S 68906, and S 68908). Also occurring in Brunei (e.g., Wong WKM 1654).
Ecology. Rare, in mixed dipterocarp forest on yellow sandy soils overlying Miocene
sandstone, at altitudes to 400 m. Recorded from Mulu NP; endangered.
2. COTYLELOBIUM Pierre
(Greek, kotyle = a small cup, lobos = a pod; the receptacle)
resak (preferred name)
Fl. For. Coch. 3 (1889) sub. t. 235; Merrill, EB (1921) 408; Ridley, FMP 1 (1922) 239; Foxworthy,
Malay. For. Rec. 10 (1932) 245; Slooten, Bull. Jard. Bot. Buitenz. 3, 10 (1929) 393, Bull. Jard. Bot.
Buitenz. 3, 12 (1932) 43; Masamune, EPB (1942) 484; Symington, Malay. For. Rec. 16 (1943) 232;
Ashton, MDB (1964) 56, MDBS (1968) 24, FM 1, 9 (1982) 340; Meijer & Wood, Sabah For. Rec. 5
(1964) 323; Burgess, TBS (1966) 226; Anderson, CLTS (1980) 110; PROSEA 5, 1 (1993) 139;
Kessler & Sidiyasa, TBSAEK (1994) 90; Coode et al. (eds.), CLBD (1996) 67. Synonym: Dyerella
F.Heim, Rech. Dipt. (1892) 123.
Main canopy and emergent trees with dense even domeshaped crowns; buttresses low,
stout, rounded. Bark at first greyish, smooth, hoopmarked, becoming irregularly flaky to
shaggy (in C. lanceolatum), flakes leaving scrollmarked surfaces; inner bark homogeneous
pale brown; dammar and sapwood as in Vatica. Twigs slender, greyish brown tomentose,
terete. Leaf buds small. Stipules fugaceous. Leaves epilose or tomentose below; blade
oblong to ovatelanceolate, margin revolute; lateral veins slender, unraised above and
hardly so below, arched, bifurcating towards the margin and anastomosing to form a
looped intramarginal vein, with many shorter intermediates similarly bifurcating;
intercostal venation reticulate, obscure; petiole short, not geniculate. Inflorescences
axillary or rarely terminal panicles. Flowers: buds ovoidlanceolate; corolla cream, petals
free, ellipticoblong; stamens to 15, in 3 whorls, filaments short and deltoid or filiform,
connate at base, anthers linear oblong, latrorse, setose along the margins, connectival
appendage slender, shorter than anther; ovary free, globose, pubescent around base, style
filiform, many times longer than ovary, stigma small, trifid. Fruits: calyx with 2 long and 3
short lobes, the lobes united at base into a shallow cup. Nuts free from calyx base, round in
crosssection.
79
Fig. 3. Cotylelobium burckii (A–L) and C. lanceolatum (M–P). A, leafy twig bearing post
anthesis flowers and very young fruits; B, detail of indumentum on lower leaf surface; C,
side view of tufted hairs; D, flower bud; E, open flower; F, open flower with sepals, petals
and several stamens removed; G, longitudinal section of gynoecium; H, adaxial view of
stamens; I, abaxial view of stamens; J, mature fruit; K, transverse section of mature fruit; L,
longitudinal section of mature fruit; M, lower leaf surface; N, detail of indumentum on
lower leaf surface; O, oblique top view of tufted hairs; P, mature fruit. (A–C from Egon
610, D–I from S 37031, J–L from Wong WKM 937; M–P from bb. 29668.)
80
Distribution. Five species distributed in Sri Lanka, Peninsular Thailand, Sumatra,

Peninsular Malaysia, and Borneo; 3 species in Sabah and Sarawak.
Ecology. Locally common in mixed dipterocarp, kerangas and lower montane forests at
altitudes to 1500 m, on humic yellow and white more or less sandy soils on well drained
sites.
Use. Cotylelobium melanoxylon reaches commercial size, and its hard dark brown
heartwood is used for heavy construction.
Notes. An interesting small genus with a curious disjunct distribution; its leaves recall those
of the archaic dipterocarp of the ancient sandstone Guyana Highlands, Pakaraimaea.
Species occurring in Sabah and Sarawak are sometimes difficult to distinguish, populations
sometimes exhibiting one or more characters intermediate between two species; the
commonest forms of such variation are noted under the relevant species.
Key to Cotylelobium species

1. Leaf blade sparsely puberulent or glabrescent, drying dull olivebrown…………………

……………………………………………………………………….3. C. melanoxylon
Leaf blade densely tomentose below……………………………………………………2
2. Leaf blade oblonglanceolate, larger, 8–12 × 3–4.5 cm, drying dull olivebrown above,
densely shortly evenly bright yellowish ochre tomentose below; intramarginal vein
close to margin………………………………………………………….…1. C. burckii
Leaf blade narrowly ovatelanceolate, smaller, 6–8 × 2.5–3 cm, drying dark greyish
brown above, densely dark grey scabridtomentose below; intramarginal vein distant
from the margin…………………………………………………….…2. C. lanceolatum
1. Cotylelobium burckii (F.Heim) F.Heim Fig. 3A–L

(W. Burck, 1848–1910, botanist of Dipterocarpaceae and Sapotaceae)
Rech. Dipt. (1892) 122; Slooten, Bull. Jard. Bot. Buitenz. 3, 9 (1927) 78; Ashton, Gard. Bull. Sing. 20
(1963) 243, op. cit. (1964) 57, op. cit. (1968) 24, op. cit. (1982) 342; Anderson op. cit. 110; PROSEA
op. cit. 142; Coode et al. (eds.) op. cit. 67. Basionym: Vatica burckii F.Heim, Bull. Mens. Soc. Linn.
Paris 2 (July 1891) 956. Type: Beccari PB 3260, Borneo, Sarawak, Marop (holotype PC).
Synonyms: Cotylelobium flavum Pierre, Fl. For. Coch. 4 (Oct. 1891) t. 258A, Merrill op. cit. 408,
Slooten op. cit. (1929) 396, Foxworthy op. cit. 246, Symington, Gard. Bull. S. S. 8 (1934) 36,
Masamune op. cit. 484, Brown, FTSB (1955) 95; C. asperum Slooten op. cit. (1929) 401.
Main canopy tree, to 40 m tall, to 70 cm diameter. Young parts, buds, leaf midrib below,
petiole, inflorescence, and ovary densely shortly evenly persistently buffpubescent. Leaves
thickly coriaceous, drying dull olivebrown above, densely bright ochreous tomentose
below; blade oblonglanceolate, 8–12 × 3–4.5 cm, base broadly cuneate to obtuse, margin
81
prominently revolute, apex with acumen to 0.7 cm long; lateral veins 10–12 pairs, hardly
elevated, comparatively straight; intramarginal vein close to the margin; petiole 1.5–2 cm
long. Inflorescences to 15 cm long, axillary or rarely terminal. Flowers: buds ovoid
lanceolate, to 10 × 4 mm, acute; calyx lobes linear, obtuse, 3 shorter lobes lanceolate, acute;
petals narrowly oblong, glabrous; stamens inner 5 slightly longer than the others,
connectival appendage c. ¼x as long as anther; ovary subglobose, densely tomentose, styles
slender, 3–4 times as long as ovary. Fruits: calyx densely tomentose, 2 longer lobes c. 1.5
cm broad, 3 shorter ones linear, to 1.2 cm long.
Vernacular name. Sarawak—resak durian (preferred name).
Distribution. Endemic in Borneo; absent from Sabah. In Sarawak known from Bau,
Bintulu, Kuching, Lawas, Marudi, Miri, Mukah, Sibu, and Sri Aman districts (e.g., S 17870,
S 37121, S 47027, S 66782, S 72947, and S 73349). Also occurring in Brunei (e.g., BRUN
70, FMS 32615, S 7805, and Wong WKM 935) and C and SE Kalimantan (e.g., bb. 19876,
bb. 23501 and bb. 29681).
Ecology. Locally common in kerangas forest on deep white sand podsols on raised beaches
behind present and Pleistocene coastlines, rare on sandstone, at altitudes to 300 m.
Vulnerable owing to the destruction of its habitat, but well represented in Bako NP and
occurring in Mulu NP.
2. Cotylelobium lanceolatum Craib Fig. 3M–P

(Latin, lanceolatus = in the form of a lance; the shape of leaf blade)
Bull. Misc. Inform. Kew (1913) 113; Ashton op. cit. (1982) 343; PROSEA op. cit. 142; Coode et al.
(eds.) op. cit. 67. Type: East Asiatic Co. H 20171, Thailand, Surat (holotype K). Synonyms:
Cotylelobium malayanum Slooten op. cit. (1932) 43, Foxworthy op. cit. 247, Symington Gard. Bull. S.
S. 9 (1938) 349, op. cit. (1943) 235, Browne op. cit. 95, Ashton op. cit. (1964) 58, Meijer & Wood op.
cit. 324, Anderson op. cit. 111; C. flavum auct. non Pierre: Ridley op. cit. 239, Slooten op. cit. (1929)
396, p.p., Foxworthy op. cit. 247.
Main canopy tree, to 45 m tall, to 1.2 m diameter. Bark eventually densely shaggy. Young
parts, buds, leaf venation below, petiole, and inflorescence densely greyish brown to rufous
scabridpubescent; leaf undersurface and ovary evenly so. Leaves thinly coriaceous, drying
dark greyish brown above, densely dark grey scabridtomentose below; blade narrowly
ovatelanceolate, 6–8 × 2.5–3 cm, base broadly cuneate, margin slightly revolute, apex with
acumen to 0.5 cm long; lateral veins 10–12 pairs, hardly raised below, joining into a
prominent looped intramarginal veins c. 2 mm from margin; petiole 0.6–1 cm long.
Inflorescences to 6 cm long, terminal or axillary. Flowers: buds ovoid, obtuse, to 8 × 3.5
mm; calyx densely bufftawny tomentose outside, sparsely so within, lobes deltoid, unequal;
petals narrowly oblong, obtuse, glabrous; inner stamens slightly longer than the rest,
filaments triangular, anthers narrowly oblong, with prominent setose margin, connectival
appendage short, slender; ovary subglobose, densely tomentose, style c. 3x as long as ovary.
Fruits: calyx scabridtomentose; 2 larger lobes oblong, to 4.5 × 1.2 cm, 3 shorter ones
broadly hastate, 1.2–2 × 0.4 cm.
82
Vernacular names. Sabah—resak kelabu (preferred name). Sarawak—resak batu

(preferred name).
Distribution. Peninsular Thailand to eastern coastal area of Peninsular Malaysia, Anambas

Is. and Borneo. Throughout Borneo where its habitat occurs. In Sabah known from
Sandakan, Sipitang and Tawau districts (e.g., SAN 18651, SAN 18652 and SAN 39272) and
in Sarawak from Bau, Bintulu, Kuching, Lawas, Limbang, Marudi, Miri, Mukah, and Serian
districts (e.g., S 12583, S 24065, S 28068, S 32850, S 38295, and S 47628). Also occurring
in Brunei (e.g., BRUN 414, BRUN 3045 and BRUN 5779) and SE Kalimantan (e.g., bb.
24015, bb. 32387, Kostermans 6371, Kostermans 9554, and Kostermans 12512A).
Ecology. Locally abundant in kerangas forest, on raised beach terraces, sandstone plateaux,
and in W Sarawak on organic soils over limestone; locally abundant in lower montane
kerangas forest on ridges, at altitudes to 1500 m. Common in Bako NP and occurring in
Mulu NP; probably not vulnerable.
Notes. Collections from some lowland localities, such as the Bako NP, in Sarawak, have an
even, hardly scabrous tomentum but are otherwise typical.
3. Cotylelobium melanoxylon (Hook.f.) Pierre Fig. 4.

(Greek, melanos = blackish, xylon = wood; the dark heartwood)
Fl. For. Coch. 3 (1889) t. 235; Merrill op. cit. (1921) 408; Slooten op. cit. (1927) 78, op. cit. (1929)
403, op. cit. (1932) 44; Masamune op. cit. 484; Symington op. cit. (1943) 236; Browne op. cit. 96;
Ashton op. cit. (1964) 59, op. cit. (1968) 24, op. cit. (1982) 341; Meijer & Wood op. cit. 324; Burgess
op. cit. 228; Anderson op. cit. 111; PROSEA op. cit. 143; Kessler & Sidiyasa op. cit. 90; Coode et al.
(eds.) op. cit. 67. Basionym: Anisoptera melanoxylon Hook.f., Trans. Linn. Soc. 23 (1860) 160. Type:
Lowe s.n., Borneo, Sabah, Labuan, Lubok Danau (holotype K). Synonyms: Vatica melanoxylon
(Hook.f.) Miq., Ann. Mus. Bot. Lugd.Bat. 3 (1867) 956; V. beccariana F.Heim op. cit. (1891) 955; V.
harmandii F.Heim op. cit. (1891) 955; Cotylelobium beccarii (F.Heim) Pierre op. cit. (1891) t. 258B,
Merrill op. cit. 408, Slooten op. cit. (1929) 405, Masamune op. cit. 484; C. harmandii (F.Heim)
F.Heim op. cit. (1892) 122, Merrill op. cit. 408, Masamune op. cit. 484; C. beccarianum (F.Heim)
F.Heim op. cit. (1892) 122; C. leucocarpum Slooten op. cit. (1929) 399, Masamune op. cit. 484; V.
leucocarpa (Slooten) Foxw. ex Den Berger & Endert, Med. Proefst. Boschw. 11 (1925) 130.
Emergent tree, to 60 m tall, to 1.5 m diameter. Young parts shortly more or less sparsely
greyish brown puberulent, fugaceous except on inflorescence, buds, calyx base and ovary,
and occasionally leaf undersurface. Leaves thinly coriaceous, satiny below, drying dull
olivebrown; blade lanceolate, oblong or ovate, 5–10 × 2–6 cm, base broadly cuneate or
obtuse, margin slightly revolute, apex with acumen to 0.8 cm long; lateral veins 10–13
pairs, hardly raised below; intramarginal vein strongly looped, at c. 2 mm from margin;
petiole 0.9–1.2 cm long. Inflorescences to 8 cm long, axillary. Flowers: buds ellipsoid,
obtuse, to 6 × 3 mm; stamens subequal, filaments c. 3x the length of anthers, slender,
tapering, connectival appendage c. ¼x the length of style; ovary ovoid, densely pubescent,
style filiform, c. 3x as long as ovary. Fruits: 2 longer calyx lobes oblong, obtuse, c. 4.5 ×
1.2 cm, 3 shorter ones hastateacute, 0.8–1.4 × 0.2–0.3 cm. Nuts ovoid, c. 0.6 cm across,
style remnant to 5 mm long.
Vernacular names. Sabah—resak tempurung (preferred name). Sarawak—resak hitam

(preferred name).
83
Fig. 4. Cotylelobium melanoxylon. A, fruiting leafy twig; B, detail venation on lower leaf
surface; C, side view of tufted hairs; D, mature fruit; E, part of inflorescence bearing flower
buds; F, flower bud; G, open flower with one sepal and two petals removed; H, longitudinal
section of gynoecium and adaxial view of stamens. (A–C from SAN 56120, D from S 15886,
E from SAN 57677, F–H from SAN 37642.)
84
Distribution. Sumatra, Peninsular Malaysia (NE Johor) and Borneo (widespread). In Sabah
recorded from Beaufort, Kota Kinabalu, Kudat, Labuk Sugut, Sandakan, Sipitang and
Tawau districts (e.g., KEP 80303, SAN 15259, SAN 17264, SAN 38807, and SAN 57688)
and in Sarawak from Bintulu, Kuching, Lundu, Marudi, Miri, and Sibu districts (e.g., S
11855, S 12475, S 15871, S 40315, and S 72955). Also occurring in Brunei (e.g., BRUN
292, BRUN 604, FMS 48302, NN 98, and S 1678) and E Kalimantan (e.g., Arifin &
Ambriansyah B 1528).
Ecology. Locally common in mixed dipterocarp forest on yellow sandy soils, and as a
smaller, canopy tree, in kerangas on podsols; at altitudes below 400 m and usually not far
from the coast. Well represented in Lambir and Mulu NPs, but vulnerable outside the parks
system.
Notes. The leaf becomes glabrous except in eastern parts of Borneo from Sandakan
southwards, where it may remain persistently sparsely puberulent below.
3. DIPTEROCARPUS Gaertn.f.
(Greek, di = two, pteron = wing, karpos = fruit; the twowinged fruit)
keruing (preferred name), kesugoi (Murut), resak (Iban), sugoi (Murut)
Fruct. 3 (1805) 50; Blume, Bijdr. Fl. Ned. Ind. (1825) 223; King, J. As. Soc. Beng. 62, 2 (1893) 89;
Ridley, FMP 1 (1922) 211; Slooten, Bull. Jard. Bot. Buitenz. 3, 8 (1927) 263; Foxworthy, Malay. For.
Rec. 10 (1932) 56; Symington, Malay. For. Rec. 16 (1943) 153; Browne, FTSB (1955) 102; Ashton,
Gard. Bull. Sing. 20 (1963) 233, MDB (1964) 16, MDBS (1968) 6, Gard. Bull. Sing. 31 (1978) 5, FM
1, 9 (1982) 291; Backer & Bakhuizen f., FJ 1 (1964) 328; Meijer & Wood, Sabah For. Rec. 5 (1964)
230; Burgess, TBS (1966) 99; Anderson, CLTS (1980) 111; PROSEA 5, 1 (1993) 166; Kessler &
Sidiyasa, TBSAEK (1994) 91; Coode et al. (eds.), CLBD (1996) 67; Newman et al., MDFBMHHW
(1998) 59.
Emergent or sometimes main canopy trees with irregular crowns arising from a few large
twisted ascending branches; buttresses stout, generally low, concave. Bark pale or dark
greyish to orangebrown, dotted with prominent pale warty lenticels, peeling off in irregular
usually large flakes; inner bark warm brown, hard, homogeneous; dammar slow to harden,
not usually as smears on bole but exuding from cut surfaces. Sapwood pale to dark orange
yellow, grading into rust to pinkish brown heartwood. Twigs with amplexicaul stipule scars.
Terminal (leaf) buds prominent, with speciesspecific shape and indumentum. Stipules
large, usually strapshaped. Leaves aestivation plicate; coriaceous, never silvery below,
more or less corrugated between the rather straight lateral veins; margin usually sinuate
towards apex; lateral veins stout and usually prominent below, curved only near the margin;
intercostal venation scalariform, usually wellspaced, rarely subreticulate; petiole usually
long, prominently geniculate. Inflorescences racemose, terminal or axillary; rachis short,
stout, zigzag, fewflowered, unbranched or sparingly so. Flowers large; buds spindle
shaped; calyx united at base into a smooth, angled, tuberculate or flanged tube enclosing but
free from ovary, with 5 valvate lobes; petals strapshaped, cream generally with a pink
median stripe, or pink; stamens 15–50, connate at base, falling in a group tardily, filaments
compressed, slender tapering, anthers large, elongate, tapering into a short or prominent
stout tapering connectival appendage; ovary ovoid, pubescent towards apex, tapering into a
filiform style, stigma obscure. Fruits usually large; calyx united at base into a tube
85
enclosing at least the basal half of the nut; tube smooth or adorned with 5 speciesspecific
ribs, tubercles, or flanges; lobes valvate and not incrassate at base, unequal, 2 longer ones
aliform, oblong to spatulate and more or less distinctly 3veined, and the other 3 short, or all
subequal and short. Nuts free within calyx tube, apex pubescent with short style remnant;
germination hypogeal; cotyledons without chlorophyll, intricately folded and trapped within
calyx tube, the seedling escaping by elongation of the cotyledonary petioles.
Distribution. About 70 species, distributed from Sri Lanka and S India to Myanmar,
Thailand, IndoChina, China (Yunnan), Sumatra, Peninsular Malaysia, Java, Borneo, the
Philippines, and Lesser Sunda Is. (Bali and Sumbawa). Forty one species occur in Borneo of
which 39 in Sabah and Sarawak.
Ecology. All inland lowland habitats in Borneo at altitudes to 900 m, but rare in mixed peat
swamp forest. Light demanders as juveniles, they are particularly common on ridges, river
banks, and in some kerangas, though some are specialists of other habitats. Individual
species often occur in extensive populations, more or less without the presence of others.
Uses. Medium hardwoods which readily take preservatives; used for heavy construction,
and formerly especially for railway sleepers. The resin was tapped, especially in seasonal
climates and not widely in our area, for tallow, varnish and paint manufacture. The wood of
ensurai (D. oblongifolius) was once esteemed for rice mortars.
Notes. Like Vatica but unlike most other dipterocarps, species of Dipterocarpus are
sometimes difficult to identify on leaves alone. The leaf buds though are prominent, and
their shape, and tomentum with that of the stipules, provide invaluable diagnostic
characters. Fallen twigs with buds attached can nearly always be found.
Key to Dipterocarpus species

(based on flowering and/or fruiting specimens)
1. Calyx tube of mature fruit smooth (without ribs, tubercles or flanges)............................2

Calyx tube of mature fruit ribbed, tuberculate or flanged…………………………...…10
2. Stamens at most 15…………………………………………………………………...…3

Stamens at least 20………………………………………………………………………4
3. Fruit calyx tube ellipsoidglobose, verrucoselenticellate………..….39. D. verrucosus

Fruit calyx tube ellipsoid, smooth……………………………………..…10. D. crinitus
4. Fruit calyx lobes all short, equal…………………………………….…37. D. tempehes

Fruit calyx lobes unequal, 2 large ones winglike………………………………………5
5. Stamens c. 25; short fruit calyx lobes less than 0.7 cm long, hardly recurved…………...
…………………………………………………………………………..5. D. caudiferus
Stamens c. 30; short fruit calyx lobes at least 0.8 cm long, becoming recurved and
revolute..........................................................................................................................…6
6. Twigs and leaf buds densely ferruginous tomentose……………………………………7

Twigs and leaf buds buff or golden tomentose, or glabrous………………………….…8
86
7. Leaf blade becoming glabrous………………………………………….…38. D. validus

Leaf blade persistently tomentose below………………………………....19. D. gracilis
8. Leaf bud persistently pubescent outside……….…4. D. caudatus (subsp. penangianus)

Leaf bud glabrous outside………………………………………………………………9
9. Inside of stipules densely pubescent; lateral veins at most 11 pairs…….….23. D. kerrii

Stipules entirely glabrous; lateral veins at least 11 pairs………………...21. D. hasseltii
10. Fruit calyx tube with prominent sharp or narrowly rounded ribs more or less confined to
the distal half, terminating in bosslike tubercles……………………….24. D. kunstleri
Fruit calyx tube either ribbed, angled or flanged, or tuberculate………………………11
11. Mature fruit calyx tube with 5 swelling ribs or tubercles below the neck, otherwise
spherical……………………………………………………………………………..…12
Mature fruit calyx tube angled, ridged, ribbed, or flanged………………………….…17
12. Mature fruit calyx tube ovoid………………………………..……..…2. D. applanatus

Mature fruit calyx tube subglobose or globose…………………………………….…13
13. Leaves with at most 16 pairs of lateral veins………………………………….…14..
14. Leaf blade pale ochreouspubescent below………………………….……33. D. rigidus

Leaf blade glabrous……………………………………………………………………15
15. Leaf bud and stipules outside glabrous, sometimes pruinose……….…9. D. costulatus
Leaf bud and stipules outside tomentose…………………………….…18. D. globosus
16. Fruit calyx lobes all short, subequal……………………………….…12. D. elongatus

Fruit calyx lobes unequal, 2 larger ones winglike……………….…22. D. humeratus
17. Fruit calyx tube with intricately folded flanges………………………………………18

Fruit calyx tube not flanged or if flanged, then the flanges straight or wavy but not
intricately folded……………………………………………………………………..20
18. Twigs terete; leaf blade concave, venation tomentose below…………25. D. lamellatus
Twigs compressed; leaf blade not concave, venation glabrous below………………19
19. Leaf blade ovatelanceolate, prominently corrugated, margin prominently revolute,

lateral veins at least 15 pairs…………………………………………….…26. D. lowii
Leaf blade broadly ovate, hardly corrugated, margin hardly or not revolute, lateral veins
at most 12 pairs………………………………………………….…31. D. pachyphyllus
20. Fruit calyx tube at least 1½x as long as broad, spindleshaped or narrowly ellipsoid, or
narrowly obovoid……………………………………………………………………21
87
Fruit calyx tube less than 1½x times as long as broad, more or less globose or broadly
ovoid though sometimes with flanges decurrent into pedicel and hence appearing
ellipsoid………………………………………………………………………………30
21. Fruit calyx tube angled or ribbed rather than flanged……………………………..…22

Fruit calyx tube distinctly flanged………………………………………………….…23
22. Leaf blade at least 15 cm long, broadly obovate to suborbicular………6. D. confertus

Leaf blade at most 12 cm long, narrowly ellipticlanceolate……………14. D. fagineus
23. Fruit calyx tube densely pale buffpubescent, with very narrow undulate flanges……….
…………………………………………………………………….29. D. oblongifolius
Fruit calyx tube glabrous; flanges, if narrow, straight………………………………24
24. Fruit calyx tube flanges at least 8 mm wide………………………………………..…25

Fruit calyx tube flanges at most 7 mm wide…………………………………………27
25. Stamens c. 30…………………………………………………….…20. D. grandiflorus

Stamens c. 15…………………………………………………………………………26
26. Young parts glabrous……………………………………………17. D. glabrigemmatus

Young twig, bud, stipule outside and petiole pubescent…………..32. D. palembanicus
27. Fruit calyx tube flanges of constant width…………………………….…28. D. nudus

Fruit calyx tube flanges widest in the distal half………………………………………28
28. Fruit calyx tube flanges more or less confined to distal half; leaf bud and leaf
undersurface pubescent; stamens c. 25……………………………..…3. D. borneensis
Fruit calyx tube flanges continuous from base; tomentum not as above; stamens, if
known, c. 15……………………………………………………………………………29
29. Leaf bud buff velutinous, leaf blade below sparsely so………………15. D. fusiformis
Leaf bud outside and leaf blade glabrous..………………………………27. D. mundus
30. Fruit calyx tube persistently pubescent……………………………………………..…31

Fruit calyx tube glabrescent……………………………………………………………32
31. Leaf including petiole glabrous…………………………………….16. D. geniculatus

Leaf below and petiole velutinous..…………….…7. D. conformis (subsp. borneensis)
32. Fruit calyx tube angled or ribbed rather than flanged………………...1. D. acutangulus
Fruit calyx tube truly flanged……………………………………………………….…33
33. Fruit calyx tube flanges exceeding 10 mm wide…………………………………….…34

Fruit calyx tube flanges less than 9 mm wide……………………………………….…35
34. Fruit calyx tube flanges not decurrent with pedicel, frequently undulate; leaf blade
glabrous…………………………………………………………....36. D. sublamellatus
Fruit calyx tube flanges decurrent into pedicel; leaf blade densely tomentose below……
………………………………………………………………………….35. D. stellatus
88
35. Leaves broadly obovate, apex obtuse or retuse……………………34. D. sarawakensis

Leaves narrowly ellipticovate, broadly ellipticovate to narrowly obovate, apex
subacute, acute, acuminate to cuspidate………………………………………………36
36. Leaf blade broader, 10–15 cm wide; fruit calyx tube flanges not continuous to base……
…………………………………………………………………………..8. D. coriaceus
Leaf blade narrower, 2–9.5 cm wide; fruit calyx tube flanges continuous to base…….37
37. Fruit calyx tube flanges broadest towards apex; leaf apex cuspidate.11. D. cuspidatus
Fruit calyx tube flanges of constant width; apex of mature leaf blade not cuspidate….38
38. Mature leaf blade 10–19 × 4.5–9.5 cm……………………………..…30. D. ochraceus

Mature leaf blade less than 10 × 4.5 cm………………………………13. D. eurynchus
Key to Dipterocarpus species

1. Leaf lateral veins more than 26 pairs…………………………………….………….…2

Leaf lateral veins less than 25 pairs………………………………………………….…3
2. Leaf blade at least 28 × 13 cm…………………………………………12. D. elongatus

Leaf blade at most 25 × 12 cm………………………………..…38. D. validus (in part)
3. Petiole at most 1.3 cm long………………………………………………………….…4

Petiole at least 1.3 cm long………………………………………………………..……6
4. Leaf apex retuse to obtuse; venation densely tawny longtomentose below……….….…

……………………………………………………………………..34. D. sarawakensis
Leaf apex acute or acuminate; venation sparsely pubescent or glabrescent below..……5
5. Leaf lateral veins at most 9 pairs. Trees of high shale ridges…………13. D. eurynchus
Leaf lateral veins at least 9 pairs. Trees of flood plains……….37. D. tempehes (in part)
6. Leaf undersurface persistently (at least sparsely) tomentose……………………………7

Leaf undersurface glabrous or becoming so……………………………………….…23
7. Leaf blade prominently concave…………………………………………………….…8

Leaf blade not concave (though more or less corrugated)………………………….…9
8. Petiole less than 4 cm long………………………………………..…25. D. lamellatus

Petiole more than 5 cm long……………………………………………6. D. confertus
9. Leaf bud glabrous, drying black, or occasionally sparsely tomentose……………..…10

Leaf bud persistently densely ochreous or brown tomentose……………………….…11
89
10. Leaf lateral veins at least 12 pairs; leaf blade drying greyish brown…………………….
…………………………………………………………………5. D. caudiferus (in part)
Leaf lateral veins at most 11 pairs; leaf blade drying warm rustbrown………………….
………………………………………………………………….17. D. glabrigemmatus
11. Leaf blade densely evenly pale ochreous or pinkish brown tomentose below...........…12
Leaf blade sparsely pubescent, or rustbrown, or scabridtomentose below..................14
12. Leaf blade at most 12 × 7 cm, thinly coriaceous; lateral veins 15–18 pairs……………...
…………………………………………………….7. D. conformis (subsp. borneensis)
Leaf blade at least 13 × 8 cm, thickly coriaceous; lateral veins 12–16 pairs…………..13
13. Tomentum pink. Trees of peat swamp forest……………………………8. D. coriaceus

Tomentum ochreyellow. Trees of mixed dipterocarp forest…………..…33. D. rigidus
14. Leaf blade with c. 20 pairs of lateral veins………………………………………..…15

Leaf blade with less than 18 pairs of lateral veins………………………………….…16
15. Leaf blade at least 20 × 12 cm; venation sparsely puberulent below....22. D. humeratus
Leaf blade at most 15 × 10 cm; venation persistently rustbrown tomentose below……..
…………………………………………………………………………….19. D. gracilis
16. Tomentum on twig and leaf bristlelike…………………………………………….….17

Tomentum, where present, short and even………………………………………….…19
17. Leaf apex with slender cuspidate acumen; lateral veins at most 9 pairs………………….
………………………………………………………………………..11. D. cuspidatus
Acumen of leaf apex, if present, broad and tapering; lateral veins at least 12 pairs…...18
18. Leaf blade elliptic, margin revolute, apex obtuse to shortly acuminate….10. D. crinitus
Leaf blade ovate, margin not revolute, acumen prominent………………35. D. stellatus
19. Petiole at least 3 cm long……………………………………………..16. D. geniculatus

Petiole at most 2.6 cm long………………………………………………………….…20
20. Twig stout; leaf bud ovoidlanceolate. Trees of high hills………….…30. D. ochraceus
Twig slender; leaf bud falcate to linearlanceolate. Trees of the lowlands………….…21
21. Leaf blade broadly ellipticobovate, prominently corrugated..37. D. tempehes (in part)
Leaf blade narrowly elliptic to lanceolate, hardly corrugated……………………..….22
22. Petiole at most 1.7 cm long………………………………….…14. D. fagineus (in part)

Petiole at least 2 cm long…………………………………………...…15. D. fusiformis
23. Leaf bud glabrous outside, drying black………………………………………………24

Leaf bud variously tomentose, drying yellowish, buff, brown or grey……………..…31
24. Leaf bud ovoid, broad and stout………………………..……1. D. acutangulus (in part)
Leaf bud long and slender (falcate)……………………………………………………25
90
25. Leaf lateral veins usually less than 11 pairs………………………………………..…26

Leaf lateral veins usually more than 11 pairs……………………………………….…27
26. Leaf blade narrowly elliptic to obovate, base narrowly obtuse………..…27. D. mundus
Leaf blade broadly elliptic, base cuneate………………………………..…23. D. kerrii
27. Petiole at most 2 cm long. Trees of banks of upland rivers………………………………

…………………………………………………………….29. D. oblongifolius (in part)
Petiole at least 2.5 cm long. Trees of mixed dipterocarp forest…………………….….28
28. Leaf blade broadly ellipticovate, thickly coriaceous and corrugated…..9. D. costulatus
Leaf blade narrowly elliptic, thinly coraiceous and hardly corrugated……………..…29
29. Intercostal venation widely scalariform to subreticulate……………….…28. D. nudus

Intercostal venation densely regularly scalariform……………………………………30
30. Leaf buds at first hairy……………………………………..…5. D. caudiferus (in part)

Leaf buds always glabrous………………………………………………21. D. hasseltii
31. Twigs at least 5 mm diameter apically; petioles at least 2 mm diameter; both stout…..32
Twigs at most 5 mm diameter apically; petioles at most 2 mm diameter; both
slender………………………………………………………………………………….36
32. Leaf lateral veins at least 15 pairs…………………………………………………..…33

Leaf lateral veins at most 15 pairs………………………………………………..……35
33. Twigs compressed……………………………………………………………26. D. lowii

Twigs terete……………………………………………………………………………34
34. Leaf lateral veins at least 22 pairs; bud and stipules shaggy rustbrown tomentose……..
……………………………………………………………………38. D. validus (in part)
Leaf lateral veins at most 17 pairs; bud and stipules shortly evenly buffpubescent……..
………………………………………………………………………20. D. grandiflorus
35. Petiole at least 3.5 cm long………………………………………….…2. D. applanatus

Petiole at most 2.5 cm long…………………………………………..…18. D. globosus
36. Tomentum on petiole, leaf blade and leaf bud sparse, greyish brown……………………
………………………………………………………………...5. D. caudiferus (in part)
Tomentum on petiole, leaf blade and leaf bud yellowish, buff, golden, or dark or rust
brown; dense at least on leaf bud…………………………………………………..…37
37. Leaf lateral veins at least 16 pairs…………………………………………………..…38

Leaf lateral veins at most 14 pairs………………………………………………….…39
38. Leaf bud densely pale yellowish longtomentose…………29. D. oblongifolius (in part)
Leaf bud densely minutely greyishbuff pubescent………………….…24. D. kunstleri
39. Twigs compressed……………………………………………….…31. D. pachyphyllus

Twigs terete…………………………………………………………………………….40
91
40. Leaf bud slender, more or less falcate, with apical tuft, tomentum otherwise very short
…………………………………………………………………………………….……41
Leaf bud ellipsoidovoid to broadly ovoid, stout, densely buff or rustbrown long
tomentose…...............................................................................................................…44
41. Leaf bud minutely goldenbrown pubescent, glistening due to the presence of minute
radiating shiny stellate hairs; leaf blade broadly ellipticovate, drying pinkish
brown…………………………………………………………………39. D. verrucosus
Leaf bud evenly greyishbuff pubescent, the hairs inclined towards the apex; leaf blade
elliptic, lanceolate, or obovate, drying pale rustbrown or orangebrown………….....42
42. Leaf blade broadly ellipticobovate, prominently corrugated…37. D. tempehes (in part)
Leaf blade narrowly elliptic, flat………………………………………………………43
43. Leaf lateral veins at least 10 pairs……………….4. D. caudatus (subsp. penangianus)

Leaf lateral veins at most 10 pairs…………………………..…14. D. fagineus (in part)
44. Leaf bud densely buff longtomentose………………………1. D. acutangulus (in part)

Leaf bud golden to rusttomentose or pale fulvoushirsute……………………….….45
45. Leaf lateral veins at least 12 pairs; leaf blade thinly coriaceous…..32. D. palembanicus
Leaf lateral veins at most 12 pairs; leaf blade coriaceous………………………….….46
46. Leaf bud rust to darkbrown tomentose. Trees of kerangas forest…….3. D. borneensis
Leaf bud pale fulvoushirsute. Trees of mixed dipterocarp forest, especially on flat land
……………………………………………………………………..36. D. sublamellatus
1. Dipterocarpus acutangulus Vesque

(Latin, angulus = angle, acutus = sharp; the ribs of fruit calyx tube)
C. R. Ac. Sci. Paris, 78 (1874) 626; Slooten op. cit. (1927) 321, Reinwardtia 5 (1961) 457; Masamune,
EPB (1942) 485; Symington op. cit. (1943) 166; Browne op. cit. 107; Ashton op. cit. (1963) 240, op.
cit. (1964) 22, op. cit. (1968) 11, op. cit. (1982) 322; Meijer & Wood op. cit. 235; Burgess op. cit. 99;
Anderson op. cit. (1980) 111; PROSEA op. cit. 172; Coode et al. (eds.) op. cit. 66; Newman et al. op.
cit. 76. Type: Beccari PB 2913, Borneo (holotype FI; isotype K). Synonyms: Dipterocarpus
tawaensis Slooten op. cit. (1927) 313, Merrill, PEB (1929) 201, Masamune op. cit. 488; D.
helicopteryx Slooten, Bull. Jard. Bot. Buitenz. 3, 16 (1940) 441.
Emergent tree, to 60 m tall, to 1.2 m diameter. Bark becoming pale buffbrown, flaky.
Young parts densely evenly buffpubescent, persistent only on leaf bud, stipules and ovary.
Twigs c. 3 mm diameter apically, terete, with somewhat swollen nodes. Leaf buds short,
stout, broadly ovoid, to 8 × 5 mm, buffpubescent but occasionally glabrous and drying
black. Stipules to 5 × 0.8 cm. Leaves coriaceous, drying yellowish brown, glabrous or
almost so below; blade elliptic to ovate, 7–10 × 3–6 cm, base obtuse or cuneate, margin
sinuate, apex with tapering acumen to 1 cm long; lateral veins 7–12 pairs, stout below;
petiole 1.5–2.5 cm long, to 2 mm diameter. Inflorescences: rachis axillary, slender, terete,
glabrous, to 6 cm long, c. 1.5 mm diameter. Flower: buds spindleshaped, to 1.5 × 0.5 cm;
calyx tube narrowly obconical, narrowly 5winged, shorter sepal lobes vestigial; stamens
15, anthers c. 6 mm long including appendages, narrowly deltoid; style exceeding anthers,
slender glabrous. Fruits: calyx tube globose, to 2.5 cm diameter, glabrescent, base
92
becoming impressed, 5ribbed from base to apex; 2 longer calyx lobes to 10 × 2.5 cm, 3
shorter ones deltoid, to 0.5 × 0.5 cm, revolute.
Vernacular names. Sabah—keruing merkah (preferred name). Sarawak—keruing beludu

(preferred name).
Distribution. Peninsular Malaysia (rare) and Borneo except southeastern parts. Widespread
in Sabah and known from Beaufort, Kinabatangan, Kota Kinabalu, Labuk Sugut, Lahad
Datu, Ranau, Sandakan, Semporna, Sipitang, and Tawau districts (e.g., SAN 15169, SAN
16183, SAN 18477, SAN 21408, and SAN 90469) and in Sarawak from Bintulu, Kapit,
Kuching, Lawas, Lundu, Miri, and Serian districts (e.g., S 13372, S 15145, S 22062, S
23800, and S 46477). Also occurring in Brunei (e.g., BRUN 927, BRUN 985, BRUN 2637,
BRUN 3348, and S 1669) and Kalimantan (e.g., bb. 10012 and bb. 19811).
Ecology. Locally common in mixed dipterocarp forest on yellow sandy and sandy clay
soils, at altitudes below 400 m. Scattered also on inland shale and sandstone ridges, at
altitudes to 1000 m. Well represented in Sarawak parks system and in Sepilok FR;
elsewhere vulnerable.
Notes. A variable species in leaf size and texture; the leaf being larger and more coriaceous
in W Sarawak than elsewhere where it may be difficult to distinguish from D. globosus
(q.v.). The Kinabalu endemic, D. ochraceus, appears to be descended from it.
2. Dipterocarpus applanatus Slooten Fig. 5.

(Latin, applanatus = flattened out; the leaf blade)
Bull. Jard. Bot. Buitenz. 3, 16 (1940) 443; Masamune op. cit. 485; Meijer & Wood op. cit. 237;
Burgess op. cit. 99; Ashton op. cit. (1968) 11, op. cit. (1982) 310; Anderson op. cit. (1980) 111;
PROSEA op. cit. 173; Newman et al. op. cit. 78. Type: Puasa SAN 1720, Borneo, Sabah, Sandakan
district (holotype K).
Emergent tree, to 50 m tall, to 1.5 m diameter; buttresses prominent, to 3 m tall. Bark

orange and greenishbrown mottled, thinly flaky. Leaf bud and stipule outside persistently
evenly bufftomentose; young twig, inside of stipules and base of inflorescence shortly buff
pubescent. Twigs to 11 mm diameter apically, stout, terete, pale but for slightly depressed
dark stipule scars. Leaf buds broadly ellipsoid to falcate, to 27 × 15 mm. Stipules to 5 × 2
cm. Leaves thickly coriaceous, somewhat undulate, glabrous below; blade broadly elliptic,
12–30 × 9–20 cm, base obtuse to subcordate, apex shortly abruptly acuminate; lateral veins
11–15 pairs, prominent below; intercostal venation remote, subreticulate; petiole to 6 cm
long, to 4 mm diameter, stout. Inflorescences: axillary or ramiflorous, glabrous, to 12 cm
long; rachis stout, unbranched, bearing to 5 flowers. Flower large, ellipsoid, to 4 × 1.5 cm;
calyx tube ellipsoid, smooth, sepal lobes subequal, ellipticoblong; stamens c. 50, anthers
linearlorate, c. 1.8 mm long with fleshy tapering connective appendage. Fruits: calyx tube
ovoid, to 5 × 4.5 cm, glabrous, with 5, to 8 mm wide sharp ribs most prominent distally,
sometimes absent in basal half; 2 major calyx lobes to 19 × 4.5 cm, 3 minor ones
suborbicular, to 1 × 1 cm, revolute.
Vernacular names. Sabah—keruing daun besar (preferred name). Sarawak—keruing

arong (preferred name).
93
Distribution. Endemic in Borneo with a disjunct range. In Sabah known from

Kinabatangan, Sandakan and Tawau districts (e.g., FMS 38844, SAN 17160, SAN 18752,
SAN 19677, and SAN 39151) and in Sarawak from Kuching and Serian districts (e.g., S
8587, S 9489, S 15201, S 41016, and S 41017). Also occurring in SE Kalimantan (e.g., bb.
11675, bb. 18135 and bb. 26595).
Ecology. Local but sometimes abundant in mixed dipterocarp forest, on flat land, low hills
and sometimes rocky slopes, on yellow sandy soils especially near the coast, at altitudes
below 200 m. Vulnerable.
3. Dipterocarpus borneensis Slooten

(of Borneo)
Bull. Jard. Bot. Buitenz. 3, 16 (1940) 445; Masamune op. cit. 485; Browne op. cit. 107; Ashton op. cit.
(1964) 24, op. cit. (1968) 11, op. cit. (1982) 319; Anderson op. cit. (1980) 111; PROSEA op. cit. 174;
Coode et al. (eds.) op. cit. 67; Newman et al. op. cit. 79. Type: bb. 26417, Borneo, Kalimantan, Pepas,
Muara Tewe (holotype BO; isotype KEP).
Main canopy or shortly emergent tree, to 30 m tall, to 70 cm diameter, often with crooked
trunk. Bark pale orangebrown to greyish brown, narrowly thinly flaking, becoming
shaggy. Young parts sparsely rustbrown pubescent; indumentum more or less persistent on
ovary, leaf venation below and petiole, and dense and persistent on leaf bud and stipule
outside. Twigs 3–5 mm diameter apically, terete. Leaf buds ovoid, to 7 × 3 mm. Stipules to
4 × 0.5 cm. Leaves coriaceous, drying greyish brown, sparsely pubescent to glabrous
below; blade broadly ovate to elliptic, 7–12 × 3–7 cm, base obtuse or broadly cuneate,
margin frequently wavy towards apex, apex with tapering acumen to 0.8 cm long; lateral
veins 9–12 pairs, slender but prominent below; petiole 1.5–2.5 cm long, to 2 mm diameter,
slender. Inflorescences to 6 cm long, singly or rarely doubly branched. Flower: buds to 4 ×
1 cm; stamens c. 25, anthers short, narrowly sagittate. Fruits: calyx tube narrowly obovoid
to ellipsoid, to 1.5 × 1 cm, glabrous, tapering at each end, with 5, c. 1 mm wide flanges in
the distal half; 2 major calyx lobes to 7.5 × 1.8 cm, 3 minor ones c. 0.5 × 0.4 cm, revolute
and recurved.
Vernacular names. Sarawak—keruing sindor (preferred name), resak kerangas (Iban).
Distribution. Sumatra and Borneo. In Borneo, common and widespread in Sarawak and
recorded from Bintulu, Kuching, Lawas, Lundu, Marudi, Samarahan, Serian, Sibu, and Sri
Aman districts (e.g., S 9855, S 12492, S 13732, S 16222, and S 16236) but absent in Sabah.
Also occurring in Brunei (e.g., BRUN 648, BRUN 830, FMS 34572, Niga 141, and S 2236)
and W and SE Kalimantan (e.g., bb. 11344, bb. 16472, bb. 32384, and Kostermans 6888).
Ecology. Locally abundant in kerangas forest, usually on deep podsol soils, on raised
beaches and plateaux, at altitudes to 400 m; locally also on shallow peat. Common in Bako
NP and occurring in Mulu NP, but vulnerable elsewhere owing to habitat destruction.
4. Dipterocarpus caudatus Foxw.

(Latin, caudatus = tailed; the narrow acumen of the leaf apex)
94
Fig. 5. Dipterocarpus applanatus. A, fruiting leafy twig; B, flower bud; C, longitudinal

section of flower bud; D, open flower; E, adaxial view of stamens; F, abaxial view of
stamens; G, longitudinal section of fruit; H, basal view of calyx tube. (A and H from SAN
39151, B, D–F from S 41017, C from SAN 35868, G from FMS 35395.)
95
Fig. 6. Dipterocarpus caudiferus. A, leafy twig; B, flower bud; C, longitudinal section of

open flower; D, adaxial view of stamens; E, abaxial view of stamens; F, fruit; G, basal view
of fruit calyx tube. (A, F–G from SAN 16847, B from FMS 38786, C–E from SAN 16335.)
96
Philip. J. Sci. 13 (1918) Bot. 177, Philip. J. Sci. 67 (1938) Bot. 256; Ashton op. cit. (1978) 8, op. cit.
(1982) 305. Lectotype (Ashton, 1978): Alvarez FB 21193, the Philippines, Luzon, Barrio Hibatac,
Camarines (hololectotype K).
Distribution. Sumatra, Peninsular Malaysia, Singapore, Borneo, and the Philippines.
Notes. Two subspecies, viz. subsp. caudatus and subsp. penangianus are recognised. Of
these, only subsp. penangianus occurs in Borneo.
subsp. penangianus (Foxw.) P.S.Ashton

(of Pulau Pinang, Peninsular Malaysia)
Gard. Bull. Sing. 31 (1978) 8, op. cit. (1982) 305; PROSEA op. cit. 174; Coode et al. (eds.) op. cit. 68;
Newman et al. op. cit. 80. Basionym: Dipterocarpus penangianus Foxw., op. cit. (1932) 72,
Symington op. cit. (1943) 185, Ashton op. cit. (1964) 43. Type: Haniff SFN 3484, Peninsular
Malaysia, Penang, Mt. Olivia (holotype SING; isotype K).
Emergent tree, to 50 m tall, to 1.2 m diameter. Bark orangered, becoming pale orangegrey
when exposed, irregularly flaky. Young parts shortly sparsely fugaceous buffpuberulent;
ovary, leaf bud and stipule outside densely persistently buffpubescent. Twigs terete, 1–3
mm diameter apically, slender. Leaf buds linearfalcate, to 17 × 3 mm, tufted at apex.
Stipules c. 3.5 × 0.5 cm. Leaves drying pale orangebrown, shiny above, glabrous or
almost so below, flat; blade ellipticlanceolate to ellipticoblanceolate, 7–11 × 3.5–4 cm,
base cuneate, margin frequently wavy distally, apex prominently narrowly acuminate,
acumen to 2 cm long; lateral veins 10–12 pairs, wellspaced, hardly raised below; petiole
1.5–2.5 cm long, to 1 mm diameter, slender. Inflorescences to 12 cm long, slightly
compressed. Flowers: buds to 3 × 0.8 cm; stamens c. 30, anthers short, linear. Fruits: calyx
tube to 2 cm diameter, obturbinate and tapering into the short pedicel, smooth; 2 major calyx
lobes to 14 × 3 cm, 3 minor ones to 0.8 × 0.4 cm, becoming recurved and revolute.
Vernacular name. Sarawak—keruing gasing (preferred name).
Distribution. Sumatra (Karimun, Musala), coastal Peninsular Malaysia, Singapore, and

Borneo. In Sabah recorded from Beaufort district (e.g., SAN 15231) and in Sarawak from
Limbang, Marudi and Serian districts (e.g., S 15204, S 23310, S 32208, and S 32329). Also
occurring in Brunei (e.g., BRUN 5724, BRUN 5734, FMS 48205, KEP 80125, and S 1674),
but not known in Kalimantan.
Ecology. Local but common where it occurs, in mixed dipterocarp forests on dry ridges, at
altitudes below 400 m, especially but not exclusively near the coast; formerly most
abundant along the coastal hills on sandy soils, but also on the acid volcanic rocks of Ulu
Arip, Balingian, and on high dacite ridges in the Hose mountains and Usun Apau in
Sarawak. Endangered by land conversion.
5. Dipterocarpus caudiferus Merr. Fig. 6.

(Latin, cauda = a tail, ferre = to bear; the narrow acumen of leaf apex)
Philip. J. Sci. 29 (1926) 398; Slooten op. cit. (1927) 302, op. cit. (1961) 459; Masamune op. cit. 485;
Browne op. cit. 107; Ashton op. cit. (1963) 236, op. cit. (1964) 25, op. cit. (1968) 11, op. cit. (1982)
299; Meijer & Wood op. cit. 239; Burgess op. cit. 99; Anderson op. cit. (1980) 111; PROSEA op. cit.
97
175; Coode et al. (eds.) op. cit. 69; Newman et al. op. cit. 81. Type: Castro & Melegrito 1709,
Borneo, Banguey Is. (holotype K). Synonyms: Dipterocarpus macrorrhinus Slooten op. cit. (1927)
300; D. kutaianus Slooten op. cit. (1940) 437, Masamune op. cit. 483.
Immense emergent tree, to 65 m tall, to 1.5 m diameter, with tall columnar bole and
prominent buttresses to 3.5 m tall. Bark pale pinkish grey, appearing smooth but thinly
regularly vertically flaked. Young parts more or less caducous sparsely silky greyish brown
hispid, persistent on young trees. Twigs c. 5 mm diameter apically, terete. Leaf buds
sparsely greyish brown hispid or glabrous, lanceolate, somewhat compressed, to 25 × 3
mm, occasionally drying black. Stipules c. 7 cm long. Leaves thinly coriaceous, hardly to
prominently corrugated, drying greyish brown, glabrous or sparsely hispid with greyish
brown hairs to 2 mm long; blade narrowly elliptic, 11–20 × 5–15 cm, base obtuse or
cuneate, margin frequently sinuate distally, apex with slender acumen to 0.8 cm long;
lateral veins 12–20 pairs, dense, slender but prominent below; intercostal venation slender,
densely, regularly scalariform; petiole 3–4 cm long, slender, at most to 2 mm diameter,
persistently hispid on the knee. Inflorescences to 15 cm long, simple or singly branched.
Flowers: buds to 5 × 0.8 cm; stamens c. 25, anthers narrowly oblong, tapering. Fruits:
calyx tube globose or obturbinate, to 3.5 cm diameter, tapering into pedicel, smooth; 2
major calyx lobes to 17 × 3 cm, 3 minor ones deltoid, to 0.6 × 0.7 cm, hardly recurved.
Vernacular name. Sabah and Sarawak—keruing putih (preferred name).
Distribution. Endemic in Borneo, absent from south and southwest parts. Common
throughout Sabah and recorded from most districts (e.g., SAN 16335, SAN 17003, SAN
18603, SAN 19416, SAN 20949, SAN 27363, and SAN 134504) and throughout Sarawak and
recorded from Bau, Kapit, Kuching, Lawas, Limbang, Lundu, Marudi, Miri, and Tatau
districts (e.g., S 1812, S 10159, S 13779, S 15504, and S 29681). Also occurring in Brunei
(e.g., BRUN 331, BRUN 2639, BRUN 3025, and S 1662) and E Kalimantan (e.g.,
Kostermans 6630, Kostermans 13277, Kostermans 13960, and Kostermans 13978).
Ecology. Locally common in mixed dipterocarp forest, at altitudes below 800 m, on well
structured clay soils; frequent on other clay soils, on shale, volcanic rocks and granodiorite.
Common in G. Gading NP and locally so in Mulu NP; probably not endangered.
6. Dipterocarpus confertus Slooten

(Latin, confertus = crammed together; perhaps referring to the indumentum)
Bull. Jard. Bot. Buitenz. 3, 8 (1927) 322, Bull. Jard. Bot. Buitenz. 3, 17 (1941) 104, op. cit. (1961)
460; Merrill op. cit. (1929) 201; Foxworthy op. cit. (1932) 62; Masamune op. cit. 485; Browne op. cit.
108; Ashton op. cit. (1964) 26, op. cit. (1968) 12, op. cit. (1982) 315; Meijer & Wood op. cit. 242;
Burgess op. cit. 99; Anderson op. cit. (1980) 111; PROSEA op. cit. 175; Kessler & Sidiyasa op. cit.
92; Coode et al. (eds.) op. cit. 68; Newman et al. op. cit. 82. Lectotype (Slooten, 1941): Elmer 21548,
Borneo, Sabah, Tawau district (hololectotype K; isolectotypes L, NY).
Mediumsized emergent tree, to 50 m tall, to 1.5 m diameter; bole rather short; crown fresh
green from below, largeleafed and diffuse. Bark regularly vertically cracked and flaky.
Twigs at first, buds, stipule outside, petiole, midrib above, venation below, ovary, and calyx
persistently long palefulvous tuftedhispid; leaf surface persistently or caducously so.
98
Twigs to 10 mm diameter apically, stout. Leaf buds to 20 × 17 mm, large, broadly ovoid,
obtuse or subacute. Stipules broadly ovate, c. 5 × 5 cm. Leaves concave, somewhat bullate,
chartaceous, drying pale honeybrown and quickly decomposing; blade broadly obovate to
suborbicular, (18–)22–35 × (14–)16–22 cm, base obtuse, generally subpeltate, apex obtuse
or shortly acuminate; lateral veins 9–12 pairs, wellspaced; petiole 5–6 cm long,
prominently articulated. Inflorescences to 7 cm long, simple or singly branched. Flowers:
buds to 4 × 1 cm; stamens c. 25, anthers narrowly oblong, tapering. Fruits: calyx tube
narrowly obovoid, c. 3 × 1.7 cm, tapering into pedicel, 5ribbed; 2 longer calyx lobes to 14
× 3 cm, glabrous within, 3 minor ones oblong, to 1.7 × 0.7 cm, recurved distally.
Vernacular name. Sabah and Sarawak—keruing kobis (preferred name).
Distribution. Endemic in Borneo; widespread except in the southwestern parts. Common in

Sabah and known from Kinabatangan, Kudat, Ranau, Sandakan, and Sipitang districts (e.g.,
SAN 15192, SAN 16372, SAN 16545, SAN A 3065, and SAN A 4676) but less common in
Sarawak and known from Kapit, Kuching, Lawas, Limbang, Miri, and Samarahan districts
(e.g., S 1830, S 13473, S 15762, S 19857, S 32353, and S 69176). Also occurring in Brunei
(e.g., BRUN 441, FMS 30518 and FMS 37111) and E Kalimantan (e.g., bb. 19283, bb.
24014, Kostermans 6212, Kostermans 6707, and Kostermans 7278).
Ecology. Locally frequent in mixed dipterocarp forest on leached yellow clay soils,
especially on undulating lands and low ridges, occasionally at altitudes to 800 m.
Vulnerable.
7. Dipterocarpus conformis Slooten

(Latin, conformis = of a similar form; alluding the great similarity in vegetative characters
with two other largeleafed species, viz. D. concavus and D. confertus)
Bull. Jard. Bot. Buitenz. 3, 17 (1941) 102. Type: bb. 29177, Sumatra, Aceh Province, Alur Buaya,
Langsa (holotype BO).
Distribution. Sumatra and Borneo.
Notes. Two subspecies are recognised, viz. subsp. conformis which is confined to Aceh, N
Sumatra and subsp. borneensis which is endemic in Borneo.
subsp. borneensis P.S.Ashton

(of Borneo)
Gard. Bull. Sing. 20 (1963) 28, op. cit. (1964) 28, op. cit. (1968) 12, op. cit. (1982) 321; Meijer &
Wood op. cit. 244; Burgess op. cit. 99; Anderson op. cit. (1980) 111; PROSEA op. cit. 176; Coode et
al. (eds.) op. cit. 68; Newman et al. op. cit. 84. Type: G.H.S Wood SAN 15102, Borneo, Sabah,
Sipitang district, Pangi (holotype L; isotypes KEP, SAN).
Mediumsized emergent tree, to 50 m tall, to 1.2 m diameter; crown densely symmetrical,

pale greyish green from below. Bark becoming unevenly irregularly flaky. Young parts
densely pale pinkish brown velutinous, persistent except for leaf upper surface. Twigs c. 3
mm diameter apically. Leaf buds ovoid, c. 8 × 4 mm, obtuse. Stipules ovate, c. 1.5 × 0.6
99
cm, concave. Leaves thinly coriaceous, more or less corrugated but not concave; blade
obovate, 9–12 × 5–7 cm, base obtuse to subcordate, narrowly subpeltate, apex with acumen
0.4–0.8 cm long; lateral veins 15–18 pairs, slender but prominent and dense below; petiole
1.7–2.5 cm long, slender, rugose on drying. Inflorescences to 6 cm long, terminal or
axillary, unbranched or singly branched. Flowers: buds to 3 × 0.8 cm; stamens c. 30, anther
linear, short. Fruits: calyx tube ellipsoid, c. 2.5 × 2 cm, neck narrowed to 1.2 cm diameter,
persistently velutinous, with 5, c. 3 mm wide rigid incrassate flanges from base to apex; 2
major calyx lobes to 10 × 2 cm, 3 minor ones ovate, to 0.8 × 0.8 cm, revolute.
Vernacular name. Sabah and Sarawak—keruing beludu kuning (preferred name).
Distribution. Endemic in Borneo. Known in Sabah from Beaufort and Labuk Sugut
districts (e.g., SAN 15102, SAN 27582 and SAN 97592) and in Sarawak from Bintulu, Kapit,
Lawas, and Limbang districts (e.g., S 14358 and S 32390). Also occurring in Brunei (e.g.,
BRUN 738, BRUN 2602 and BRUN 5673).
Ecology. Rare in mixed dipterocarp forest on clay soils over shale, generally in steep
country, at altitudes below 800 m. Vulnerable, possibly endangered.
8. Dipterocarpus coriaceus Slooten

(Latin, coriaceus = with leathery texture; the leaf blade)
Bull. Jard. Bot. Buitenz. 3, 8 (1927) 331; Masamune op. cit. 485; Symington op. cit. (1943) 171;
Browne op. cit. 108; Anderson, Gard. Bull. Sing. 20 (1963) 157, op. cit. (1980) 111; Ashton op. cit.
(1968) 12, op. cit. (1982) 324; Newman et al. op. cit. 85. Type: bb. 8154, Borneo, SE Kalimantan,
Tawai Baru, near Kuala Kapuas (holotype BO; isotype L).
Mediumsized emergent tree, to 40 m tall, to 1 m diameter; crown irregular, diffuse. Bark

irregularly flaky. Leaf under surface, petiole and twig shortly densely evenly persistently
pinkish brown pubescent; leaf bud, stipule outside, midrib below, twig and petiole of young
tree at first densely hairy with hairs to 2 mm long. Twigs c. 10 mm diameter apically, stout.
Leaf buds ovoiddeltoid c. 25 × 10 mm. Stipules narrowly ovate, c. 4 × 2 cm. Leaves
thickly coriaceous, prominently corrugated; blade broadly ellipticovate, 16–21 × 10–15
cm, base obtuse to broadly cuneate, apex subacute; lateral veins 14–16 pairs, prominent
below, sunken above; intercostal venation obscure; petiole 4.5–6 cm long, stout,
prominently geniculate. Inflorescences and flowers unknown. Fruits: pedicels to 7 mm
long, prominent; calyx tube broadly ovoid, to 3 × 2.2 cm, glabrescent, with 5 stout flanges
or ribs to 5 mm wide, tapering abruptly apically and gradually towards base; 2 major calyx
lobes to 14 × 2.5 cm, 3 minor ones elliptic, to 1.2 × 0.6 cm, hardly revolute.
Vernacular name. Sarawak—keruing paya (preferred name).
Distribution. E Sumatra, C Peninsular Malaysia, S and W Borneo. In Borneo, known in

Sarawak from Kuching, Lundu, Mukah, Samarahan, Sibu, and Sri Aman districts (e.g., S
555, S 556, S 9820, and S 14584) and W and SE Kalimantan (e.g., bb. 7450, bb. 8154 and
bb. 9442).
100
Ecology. Very local; as scattered individuals or semigregarious populations in mixed peat

swamp forest on the inner margins. Critically endangered.
9. Dipterocarpus costulatus Slooten

(Latin, costulatus = slightlyribbed; the leaf blade)
Bull. Jard. Bot. Buitenz. 3, 8 (1927) 315, op. cit. (1941) 105; Masamune op. cit. 486; Symington op.
cit. (1943) 174; Browne op. cit. 108; Meijer & Wood op. cit. 244; Ashton op. cit. (1968) 112, op. cit.
(1982) 310; PROSEA op. cit. 177; Anderson op. cit. (1980) 112; Newman et al. op. cit. 87. Type:
Sahak CF 1908, Peninsular Malaysia, Negeri Sembilan, Kuala Pilah, Senaling Inas FR (holotype
KEP).
Emergent tree, to 50 m tall, to 1.2 m diameter. Bark pale greenish cream, patchily peeling
but sustaining an overall smooth appearance. Parts glabrous but for densely shortly buff
pubescent inner surface of stipules, bud scales, and ovary. Twigs c. 7 mm diameter apically,
at first somewhat compressed. Leaf buds falcate, to 30 × 6 mm, drying black. Stipules
lorate, c. 16 × 2 cm. Leaves thickly coriaceous, corrugated; blade broadly elliptic to ovate,
12–20 × 7–14 cm, base cuneate, apex obtuse or shortly acuminate; lateral veins 11–14
pairs, prominent below; petiole 3–6 cm long. Inflorescences to 20 cm long, once branched.
Flowers: buds to 3 × 5 cm; stamens c. 24, anther linear. Fruits: pedicels to 6 mm long,
prominent; calyx tube subglobose, to 1.5 × 2 cm, with 5 prominent flangelike median
tubercles; 2 major calyx lobes to 20 × 4 cm, 3 minor ones suborbicular, to 0.7 × 0.6 cm,
revolute.
Vernacular name. Sabah and Sarawak—keruing kipas (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah known from Tawau
district, including Sebatik Is. (e.g., FMS 38859, SAN 18470, SAN 18559, SAN 19643, and
SAN 25026) and in Sarawak from Kapit, Kuching and Lundu districts (e.g., S 4798, S
15422, S 21451, S 23805, and S 43192). Also occurring in W Kalimantan (e.g., bb. 6271).
Ecology. Common in Peninsular Malaysia but very local and uncommon in Sabah and
Sarawak, on occasionally flooded alluvium and in the ecotone between mixed dipterocarp
forest and kerangas forest, on sandy soils and shallow peat, at altitudes below 200 m.
Highly vulnerable owing to forest conversion.
10. Dipterocarpus crinitus Dyer

(Latin, crinitus = having tufts of long weak hairs; the living plant parts)
Fl. Brit. Ind. 1 (Jan. 1874) 296, J. Bot. 12 (1874) 103 & 154; King op. cit. 90; Merrill, EB (1921) 398;
Ridley op. cit. (1922) 214; Slooten op. cit. (1927) 288; Masamune op. cit. 486; Foxworthy op. cit.
(1932) 66; Symington op. cit. (1943) 175; Browne op. cit. 108; Ashton op. cit. (1964) 29, op. cit.
(1968) 13, op. cit. (1982) 299; Meijer & Wood op. cit. 246; Burgess op. cit. 99; Anderson op. cit.
(1980) 112; PROSEA op. cit. 177; Coode et al. (eds.) op. cit. 68; Newman et al. op. cit. 88. Type:
Maingay 196, Peninsular Malaysia, Malacca (holotype K). Synonym: Dipterocarpus hirtus Vesque
op. cit. (March 1874) 627.
Large emergent tree, to 60 m tall, to 2 m diameter; bole frequently misshapen; crown

goldengreen from below. Bark weathering pale greyish pink, irregularly patchily flaky;
inner bark relatively thick, creamcoloured. Plant parts tuftedsetose with goldenbrown
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bristlelike hairs c. 3 mm long, shorter on leaf blade; leaf upper surface fugaceous
pubescent. Twigs c. 3 mm diameter apically. Leaf buds oblong, c. 5 × 2 mm, obtuse.
Stipules lanceolate, c. 3 × 0.5 cm. Leaves chartaceous, not concave, persistently folded
between the veins; blade elliptic, 6–9 × 3–5 cm, base obtuse, margin revolute, apex obtuse
or shortly acuminate; lateral veins 13–15 pairs, slender but prominent below; petiole 1.5–
2.5 cm long, slender. Inflorescences to 12 cm long, singly branched, terminal or axillary.
Flowers: stamens 15, anthers linear. Fruits subsessile; calyx tube ellipsoid, to 1.8 × 0.8 cm,
smooth, 2 major calyx lobes to 8 × 1.5 cm, 3 minor ones deltoid, c. 0.3 cm long, acute.
Vernacular name. Sabah and Sarawak—keruing mempelas (preferred name).
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, and Borneo. In Sabah

recorded from Sipitang and Tawau districts (e.g., SAN 15171, SAN 15287, SAN 26894, and
SAN A 3296) and in Sarawak from Belaga, Bintulu, Kapit, Kuching, Lawas, Lundu, Marudi,
Miri, and Serian districts (e.g., S 9478, S 10121, S 15125, S 16629, and S 23897). Also
occurring in Brunei (e.g., BRUN 3137, BRUN 3268 and BRUN 5043) and W, C and SE
Kalimantan (e.g., Amiril P6. 247, bb. 27768, bb. 31415, and Kostermans 12765).
Ecology. Common in mixed dipterocarp forest on sandy clay soils, at altitudes to 800 m.
This species flowers sporadically outside mass flowering events, but seeds have high
infertility though juveniles are common. Well represented in the parks system; vulnerable
elsewhere.
11. Dipterocarpus cuspidatus P.S.Ashton

(Latin, cuspidatus = pointed at the end; the leaf blade)
Gard. Bull. Sing. 23 (1967) 261, op. cit. (1968) 13, op. cit. (1982) 324; Anderson op. cit. (1980) 112;
Newman et al. op. cit. 90. Type: Ilias S 15821, Borneo, Sarawak, Bintulu district, Labang FR
(holotype K; isotypes KEP, L).
Mediumsized emergent tree, to 50 m tall, to 1.2 m diameter. Bark rustbrown. Leaf buds
densely persistently long bufftomentose; twig, petiole, midrib above and venation below
sparsely pubescent with persistent pale greyish brown bristlelike hairs; parts otherwise
glabrous. Twigs c. 1 mm diameter apically, very slender. Leaf buds oblong, c. 7 × 3 mm.
Leaves chartaceous, hardly corrugated; blade narrowly ellipticovate, 6–11 × 2–4 cm, base
obtuse or occasionally broadly cuneate, apex cuspidate, acumen to 2 cm long, slender,
prominent; lateral veins 8–9 pairs, slender but prominent below as also the midrib; petiole
1.3–1.8 cm long, very slender. Inflorescences to 5 cm long, terminal or axillary, singly
branched. Flowers unknown. Fruits: calyx tube subglobose, to 1.7 × 2 cm, with 5, c. 2.5
mm wide rigid flanges from base to apex, broadest towards apex, glabrescent; 2 major
calyx lobes to 8 × 2 cm, 3 minor ones ovate, to 0.7 × 0.5 cm, obtuse, becoming revolute.
Vernacular name. Sarawak—keruing runcing (preferred name).
Distribution. Endemic in Borneo and confined to Segan FR, Bintulu district, Sarawak (e.g.,
S 15138, S 22061, S 22064, and S 27116).
Ecology. Formerly scattered in mixed dipterocarp forest on sandy clay soils on low hills, at
altitudes below 100 m. Endangered owing to land conversion and logging.
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12. Dipterocarpus elongatus Korth.

(Latin, elongatus = attenuated; the leaf shape)
Kruidk. (1841) 62; Merrill op. cit. (1921) 398; Slooten op. cit. (1927) 272, op. cit. (1961) 473;
Masamune op. cit. 486; Ashton op. cit. (1963) 237, op. cit. (1982) 312; PROSEA op. cit. 178; Coode
et al. (eds.) op. cit. 68; Newman et al. op. cit. 91. Type: Korthals s.n. (= RHL Sheet No. 91062130),
Borneo, Kalimantan, Barito, Sg. Punin (holotype L). Synonyms: Dipterocarpus apterus Foxw., op.
cit. (1932) 77, Masamune op. cit. 485; D. megacarpus Madani, Sandakania 2 (1993) 1.
Large emergent tree, to 60 m tall, to 1.7 m diameter; crown open, spreading; buttresses
stout, to 3 m tall. Bark greyish pink, overall smooth but thinly irregularly flaky. Twig, leaf
bud, stipule outside, petiole, inflorescence and calyx at first densely very long rufousbrown
tufted tomentose, the tufts separating and caducous as parts expand; ovary apex persistently
tomentose. Twigs to 1.5 cm diameter apically, terete, reddish brown, becoming cracked and
thinly flake. Leaf buds falcate, to 60 × 15 mm. Stipules c. 15 × 2.5 cm. Leaves coriaceous,
prominently corrugated, drying dark rustbrown below; blade elliptic, 28–50 × 13–20 cm,
base obtuse, apex shortly abruptly acuminate; lateral veins 25–38 pairs, dense, prominent
below; intercostal venation distantly scalariform; petiole 5–7 cm long, c. 5 mm diameter,
stout. Inflorescences to 12 cm long, terminal or axillary, rarely branched. Flowers
unknown. Fruits: calyx tube ovoid becoming globose, to 5 × 5.5 cm, tomatoshaped with 5
distal swellings, constricted to c. 1.5 cm diameter at the neck, crowned by 5 subequal,
obtuse, recurved lobes to 0.8 cm long.
Vernacular names. Sarawak—keruing latek (preferred name), kudan (Murut), ran (Iban).
Distribution. E Sumatra, Peninsular Malaysia, Anambas Archipelago, and Borneo (except

south and northwest parts). In Sabah known from Kinabatangan district (e.g., SAN 133997
and Wong et al. WKM 2342) and in Sarawak from Bintulu district (e.g., S 286, S 11736, S
15131, and S 27103). Also occurring in Brunei (e.g., BRUN 767, BRUN 2603, BRUN 5638,
Coode 7081, and FMS 28683) and W Kalimantan (e.g., bb. 7855 and Kostermans 9566).
Ecology. Now rare, but formerly probably quite common and locally gregarious. In the
floodplains where the rivers come out from the hills, especially on sandy alluvium;
occasional on narrow riverine terraces further inland, at altitudes below 300 m. The fruit
float in water. Critically endangered.
13. Dipterocarpus eurynchus Miq.

(Greek, eu = well, rhynchos = snout(ed); the sapling leaf acumen)
Fl. Ned. Ind., Suppl. (1862) 485; Slooten op. cit. (1927) 273, 302; Ashton op. cit. (1963) 238, op. cit.
Coode et al. (eds.) op. cit. 68; Newman et al. op. cit. 92. Type: Teijsmann HB 3255, Sumatra, Bangka
(holotype U). Synonyms: Dipterocarpus eurynchoides Scheff., Nat. Tijd. Ned. Ind. 31 (1870) 346; D.
appendiculatus Scheff., op. cit. 347, Merrill op. cit. (1921) 397, Masamune op. cit. 485; D. basilanicus
Foxw., op. cit. (1918) 179.
Mediumsized emergent tree, to 65 m tall, to 1 m diameter but usually smaller; bole

frequently misshapen; crown distinctive when the old leaves turn coppery red before falling.
Bark pale greyish brown, irregularly flaky and eventually shaggy. Leaf buds densely
persistently long bufftomentose; twig, petiole and venation below sparsely but more or less
103
persistently greyish pubescent, stipules and ovary apex persistently so. Twigs c. 2 mm
diameter apically, muchbranched. Leaf buds conical, to 10 × 3 mm. Stipules c. 3 × 0. 8 cm.
Leaves thinly coriaceous, drying dull yellow to pale orangebrown; blade ellipticobovate,
4–6(–10) × 2–3.5(–4. 5) cm, base cuneate, apex subacute to shortly acuminate; lateral veins
8–9 pairs, ascending; petiole 0.6–0.9 cm long, slender. Inflorescences to 6 cm, simple or
singly branched. Flowers unknown. Fruits: calyx tube subglobose, to 1.7 × 2 cm,
glabrescent, with 5, rigid, c. 2.5 mm wide flanges from base to apex; 2 major calyx lobes to
8 × 2 cm, 3 minor ones ovate, to 0.7 × 0.5 cm, becoming revolute.
Vernacular name. Sarawak—keruing baran (preferred name).
Distribution. Sumatra, Peninsular Malaysia, Borneo, and the Philippines. In Sarawak

known from Kapit, Limbang and Miri districts (e.g., BRUN 3073, S 23818, S 23888, and S
60135). Also occurring in Brunei (e.g., BRUN 2600, BRUN 5235, BRUN 5254, and BRUN
5257).
Ecology. In Sarawak and Brunei rare, scattered on high ridges of upper dipterocarp forest at
600–1000 m altitude. Occurring in Mulu NP but vulnerable elsewhere.
14. Dipterocarpus fagineus Vesque

(Latin, fagineus = like a beech (Fagus) tree; the leaves)
C. R. Ac. Sci. Paris 78 (March 1874) 625; King op. cit. 94; Merrill op. cit. (1921) 398; Ridley op. cit.
(1922) 216; Slooten op. cit. (1927) 318; Symington, Bull. Misc. Inform Kew (1937) 318, op. cit.
(1943) 177; Masamune op. cit. 486; Browne op. cit. 108; Ashton op. cit. (1968) 14, op. cit. (1978) 10;
Anderson op. cit. (1980) 112; Newman et al. op. cit. 93. Type: Beccari PB 3008, Borneo, Sarawak, G.
Matang (holotype P). Synonyms: Dipterocarpus prismaticus Dyer, J. Bot. 12 (April 1874) 104; D.
pseudofagineus Foxw., op. cit. (1932) 82.
Mediumsized emergent tree, to 45 m tall, to 1 m diameter. Bark greyish brown, flaky.

Young twig, leaf bud, parts of flower exposed in bud, stipule outside, petiole, and
inflorescence shortly evenly greyish brown pubescent, becoming glabrous on the
inflorescence. Twigs c. 2 mm diameter apically, terete. Leaf buds falcate, to 10 × 2.5 mm.
Stipules to 2 × 0.3 cm. Leaves thinly coriaceous, flat, drying pale rustbrown, sparsely
pubescent especially along midrib and lateral veins or glabrescent below; blade narrowly
elliptic to lanceolate, 4–9(–12) × 1.5–4(–5.5) cm, base cuneate, apex acuminate, acumen
slender to 0.8 cm long; midrib flat above, prominent below; lateral veins 8–10 pairs, slender
but elevated below; petiole 1.3–1.7 cm long, slender, to 2 mm diameter, prominently
geniculate. Inflorescences to 5 cm long, unbranched or singly branched. Flowers: buds c. 2
× 0.6 cm; stamens c. 15. Fruits: calyx tube subglobose to ellipsoid, to 1 × 0.8 cm, tapering
into a c. 6 mm long pedicel, with 5 slender acute ribs from base or distally only; 2 major
calyx lobes to 8 × 1.5 cm; 3 shorter ones deltoid, to 0.6 × 0.5 cm.
Vernacular name. Sarawak—keruing pipit (preferred name).
Distribution. Sumatra (Riau and Lingga Archipelagoes), Peninsular Malaysia, and Borneo.
Rare in Sarawak, known only from Kapit, Kuching (G. Matang) and Simunjan (G. Gaharu)
districts (e.g., S 15539, S 43809 and S 57613).
104
Ecology. Rare in mixed dipterocarp forest on hills. Conservation status unknown.
Notes. A clearly defined species outside Borneo but the Sarawak specimens, including the
type, are hardly distinguishable from D. acutangulus, and bear close resemblance also to D.
borneensis.
15. Dipterocarpus fusiformis P.S.Ashton

(Latin, fusiformis = spindleshaped; the fruit calyx tube)
Gard. Bull. Sing. 31 (1978) 12, op. cit. (1982) 319; Kessler & Sidiyasa op. cit. 93; Newman et al. op.
cit. 93. Type: Singh SAN 39170, Borneo, Sabah, Sandakan district, Mile 81, Labuk Road (holotype L;
isotype SAN).
Large emergent tree. Bark greyish. Young parts densely buffvelutinous; persistent on leaf
buds, stipule outside and ovary apex; sparsely so on twig, leaf below and petiole; caducous
elsewhere. Twigs slender, c. 2 mm diameter apically. Leaf buds linearfalcate, to 15 × 3
mm. Stipules lanceolate, c. 3 × 0.6 cm. Leaves coriaceous, hardly corrugated, drying
yellowish brown; blade elliptic or narrowly ovate, 7–17 × 3–7 cm, base cuneate or obtuse,
apex prominently acuminate, acumen slender to 12 cm long; lateral veins 13–17 pairs,
slender but prominent below; petiole 2–2.6 cm long, slender. Inflorescences and flowers
unknown. Fruits: calyx tube ellipsoid, to 2.8 × 1.8 cm, glabrous, with 5, to 6 mm wide
incrassate flanges or narrow ridges continuous from base to apex but generally broadest
distally; 2 major calyx lobes to 10 × 2.6 cm, 3 minor ones suborbicular, to 0.5 × 0.5 cm,
subrevolute.
Distribution. Endemic in Borneo. In Sabah recorded from Kinabatangan, Lahad Datu,

Ranau, Sandakan, Semporna, and Tawau districts (e.g., SAN 15379, SAN 16952, SAN
17178, SAN 63787, SAN 97110, and SAN 100176). Also occurring in Kalimantan (e.g.,
Sidiyasa S 462).
Ecology. In mixed dipterocarp forest on wellstructured and welldrained clay soils, on

basic volcanic and shale substrates, at altitudes to 600 m. Endangered by land conversion.
16. Dipterocarpus geniculatus Vesque

(Latin, geniculatus = with bent knee; the petiole)
C. R. Ac. Sci. Paris 78 (March 1874) 626; Merrill op. cit. (1921) 398; Slooten op. cit. (1927) 317, op.
cit. (1941) 99; Masamune op. cit. 486; Meijer & Wood op. cit. 248; Burgess op. cit. 900; Browne op.
cit. 109; Ashton op. cit. (1968) 14, op. cit. (1982) 320; Anderson op. cit. (1980) 112; PROSEA op. cit.
179; Newman et al. op. cit. 94. Type: Beccari PB 3034, Borneo, Sarawak (holotype P). Synonym:
Dipterocarpus angulatus Dyer, J. Bot. 12 (April 1874) 104.
Emergent tree, to 60 m tall, to 2 m diameter; crown wide, largeleaved. Bark chocolate

brown, becoming irregularly, eventually shaggily flaked. Young twig, leaf buds, stipule
outside, ovary, fruit calyx tube, and inflorescence persistently densely evenly pale cream
pubescent with short hairs; fruit calyx lobes, petiole and leaf venation below sparsely more
or less caducously so. Twigs to 13 mm diameter apically, with rows of elongate lenticels
and sinuate stipule scars. Leaf buds broadly ovoid, c. 25 × 20 mm. Stipules lanceolate, c. 6
× 2 cm. Leaves coriaceous, weakly corrugated, shiny above, sparsely pubescent or
glabrous below, drying pale chocolatebrown; blade elliptic to obovate, base obtuse, apex
105
obtuse to shortly acuminate; lateral veins 10–12 pairs, prominent below, wellspaced;
petiole 3–10 cm long, stout, glabrous. Inflorescences to 24 cm long, axillary, singly or
occasionally doubly branched. Flowers: buds to 3.5 × 1cm; stamens c. 30, anthers short,
narrowly oblong. Fruits: calyx tube glabrescent, obovoidglobose, c. 1.5 × 1.5 cm, tapering
to pedicel, with 5 stout continuous ridges to 3 mm wide towards the undulate distal ends; 2
major calyx lobes 12–15 × 2.5–4 cm, 3 minor calyx lobes cordate, c. 1 × 1.5 cm, the sides
recurved.
Vernacular names. Sabah—keruing tangkai panjang (preferred name). Sarawak—keruing

kerubung (preferred name).
Distribution. Endemic in Borneo.
Notes. In Sabah and Sarawak, two subspecies, viz. subsp. geniculatus and subsp. grandis,
are recognised.
Key to subspecies
Twigs to 7 mm diameter apically. Leaf blade 7–12 × 5–7 cm; petiole 3–5 cm long. Two
major fruit calyx lobes to 12 × 2.5 cm…………………………………...……………………
subsp. geniculatus
Known in Sarawak from Belaga, Bintulu, Kuching, Lundu, and Miri districts (e.g., S
11054, S 15144, S 16475, S 32411, S 68679, and SFN 36109) and in W Kalimantan
from the Lower Kapuas drainage (e.g., bb. 8322). In mixed dipterocarp forest on low
hills and valleys on leached sandy and sandy and silty clay soils, at altitudes to 400 m;
locally common. Vulnerable though locally frequent in Bako NP.
Twigs to 13 mm diameter apically. Leaf blade 20–35 × 12–16 cm; petiole to 10 cm long.
Two major fruit calyx lobes to 15 × 4 cm…………………………………………………….
subsp. grandis P.S.Ashton
(Latin, grandis = large; the twig and associated organs)
Gard. Bull. Sing. 20 (1963) 240, op. cit. (1964) 11; Meijer & Wood op. cit. 248; Coode et al.
(eds.) op. cit. 68; Newman et al. op. cit. 95. Type: Dzulkiple S 1870, Borneo, Brunei, Ladan Hills
FR (holotype KEP).
Known from Sipitang district in Sabah (e.g., SAN 15132 and SAN A 4051) and from
Limbang and Miri districts in Sarawak (e.g., S 1829 and S 46505). Also occurring in
Brunei (e.g., BRUN 3064, FMS 30586 and FMS 35666). As subsp. geniculatus, and on
ultrabasic rocks in E Sabah. Vulnerable though common in Lambir NP.
17. Dipterocarpus glabrigemmatus P.S.Ashton

(Latin, glabrus = hairless, gemmatus = budded; the glabrous leaf buds)
Gard. Bull. Sing. 31 (1978) 11, op. cit. (1968) 19 (‘32. Dipterocarpus sp.’), op. cit. (1982) 318. Type:
Jugah S 23849, Borneo, Sarawak, Kapit district, Bt. Raya (holotype K; isotypes KEP, SAR).
Mediumsized to large tree. Midrib and veins sparsely pubescent, ovary densely so; parts
otherwise glabrous. Twigs c. 3 × 2 mm apically, somewhat compressed and ribbed, shiny.
Leaf buds ovoid, c. 6 × 4 mm, acute, glabrous, drying black, occasionally sparsely
tomentose. Leaves glabrous except on the midrib and veins, drying warm rustbrown, more
106
or less corrugated; blade broadly ovate, 6–9 × 4.5–6 cm, base obtuse, margin prominently
sinuate distally, apex with an acumen to 0.5 cm long; midrib prominent below; lateral veins
10–11 pairs, prominent below; petiole 1.5–2.5 cm long, slender, drying rugose.
Inflorescences to 10 cm long, unbranched. Flowers: buds to 2.5 × 1.2 cm; calyx tube
broadly flanged; stamens c. 15, anthers linear. Fruits unknown.
Distribution. Endemic in Borneo. Known in C Sarawak from Kapit district (e.g., S 22339, S
23163, S 24224, S 24250, and Smythies s.n.) and in C and E Kalimantan.
Ecology. In mixed dipterocarp forest on shale ridges, at altitudes to 600 m. Rare and
endangered.
18. Dipterocarpus globosus Vesque

(Latin, globosus = spherical; the fruit calyx tube)
C. R. Ac. Sci. Paris 78 (March 1874) 627; Merrill op. cit. (1921) 398; Slooten op. cit. (1927) 304, op.
cit. (1941) 98; Masamune op. cit. 486; Browne op. cit. 109; Ashton op. cit. (1964) 33, op. cit. (1968)
15, op. cit. (1982) 311; Burgess op. cit. 100; Anderson op. cit. (1980) 112; PROSEA op. cit. 180;
Coode et al. (eds.) op. cit. 68; Newman et al. op. cit. 97. Type: Beccari PB 2914, Borneo, Sarawak,
Matang (holotype P). Synonyms: Dipterocarpus beccarianus Vesque op. cit. 627; D. beccarii Dyer, J.
Bot. 12 (April 1874) 103.
Large emergent tree, to 65 m tall, to 2 m diameter; buttresses to 3.5 m tall and wide, stout,
concave, many. Bark pale greyish brown, cleanly vertically cracked and thinly flaking.
Young parts at first persistently shortly chestnut pubescent; leaf buds and stipule outside
persistently densely long chestnut tomentose. Twigs 5–8 mm diameter apically, stout. Leaf
buds ovoid, c. 1 × 0.8 cm, acute. Stipules narrowly lanceolate, c. 7 × 0.7 cm, acute. Leaves
thickly coriaceous, corrugated, glabrous or glabrescent below; blade broadly ovate, 10–14 ×
7–9 cm, base broadly cuneate, margin sinuate, apex broadly acuminate, acumen to 0.4 cm
long; midrib and veins below fugaceous puberulent; lateral veins 12–14 pairs, wellspaced,
prominent below; petiole 2–2.5 cm long, stout, at least 2 mm diameter. Inflorescences to 8
cm long, axillary, singly branched. Flowers unknown. Fruits: calyx tube subglobose or
globose, to 3.5 cm long and broad, more or less 5ribbed from the impressed base to the
constricted apex, most prominent and slightly tuberculate distally; 2 major calyx lobes to 15
× 3.5 cm, 3 minor ones deltoid, 0.4–0.5 cm long and broad, becoming recurved and
revolute.
Vernacular name. Sabah and Sarawak—keruing buah bulat (preferred name).
Distribution. Endemic in Borneo. Known in Sarawak from Bintulu, Kuching, Lundu, Miri,
Serian and Simunjan districts (e.g., S 10058, S 21421, S 22052, S 27176, and S 27968). Also
occurring in Brunei (e.g., BRUN 5540, FMS 30488, FMS 35531, and FMS 48175).
Ecology. In mixed dipterocarp forest on deep yellow sandy soils, on coastal hills at altitudes
below 400 m. One of the most abundant large dipterocarps on sandy coastal hills between
Mukah (Sarawak) and Andulau forest (Brunei), including the Lambir Hills NP. Endangered
outside parks system.
107
19. Dipterocarpus gracilis Blume

(Latin, gracilis = slender; the twig and petiole)
Bijdr. Fl. Ned. Ind. (1825) 224; Fl. Java 2 (1829) 20; Slooten op. cit. (1927) 276, op. cit. (1940) 434;
Masamune op. cit. 486; Symington op. cit. (1943) 177; Ashton op. cit. (1963) 235, op. cit. (1964) 35,
op. cit. (1968) 15, op. cit. (1982) 301; Backer & Bakhuizen f. op. cit. 329; Meijer & Wood op. cit.
250; Burgess op. cit. 100; Anderson op. cit. (1980) 112; PROSEA op. cit. 180; Kessler & Sidiyasa op.
cit. 94; Coode et al. (eds.) op. cit. 68; Newman et al. op. cit. 98. Type: Blume 1256 (= RHL Sheet Nos.
902146–113, 114 & 118), Java, G. Parang (holotype L; isotype NY). Synonyms: Dipterocarpus
marginatus Korth., op. cit. 64; D. fulvus Blume, Mus. Bot. Lugd.Bat. 2 (1852) 37; Anisoptera
palembanica Miq., op. cit. (1862) 191, 485; D. bancanus Burck, Ann. Jard. Bot. Buitenz. 6 (1887)
196; D. skinneri King op. cit. 91; D. vanderhoevenii Koord. & Valeton, Bull. Inst. Bot. Buitenz. 2
(1899) 3; Shorea mollis Boerl., Cat. Hort. Bog. 2 (1901) 110. (For other synonyms, cf. Ashton op. cit.
(1982) 301).
Large emergent tree, to 50 m tall, to 1.5 m diameter; crown rufous from below. Bark
reddish brown, becoming thinly patchily irregularly flaky. Living exposed parts and ovary
more or less densely rufous scabridtomentose, persistent except on leaf upper surface and
calyx. Twigs c. 3 mm diameter apically. Leaf buds narrowly conical, c. 10 × 3 mm, obtuse.
Stipules narrowly lanceolate, c. 5 cm long. Leaves thinly coriaceous, not concave, drying
rufous brown; blade elliptic to ovate, 8–15 × 4–10 cm, base obtuse, apex shortly acuminate;
lateral veins 12–20 pairs, dense; petiole 2–2.5 cm long, slender. Inflorescences to 9 cm
long, terminal or axillary, singly branched. Flower: buds c. 2.5 × 0.8 cm; stamens c. 30,
anthers linear. Fruits: calyx glabrescent; tube globose, c. 2 cm diameter, smooth; 2 major
calyx lobes to 14 × 2.5 cm, 3 minor ones ovate, to 2.2 × 1 cm, becoming recurved and
revolute.
Vernacular name. Sabah and Sarawak—keruing kesat (preferred name).
Distribution. Andamans and Chittagong to the Philippines and W Java; throughout Borneo
but not in the western parts. In Sabah recorded from Beaufort, Keningau, Kinabatangan,
Kudat, Lahad Datu, Sandakan, Semporna, Sipitang, and Tawau districts (e.g., SAN 15094,
SAN 15510, SAN 16347, SAN 16406, and SAN 19141) and in Sarawak from Belaga, Kapit,
Lawas, Limbang, and Marudi districts (e.g., S 1694, S 13986, S 16534, S 22260, and S
22735). Also occurring in Brunei (e.g., BRUN 3226, FMS 35591 and SAN 17116) and SE
Kalimantan (e.g., bb. 19800).
Ecology. In mixed and upper dipterocarp forest on humusrich clay soils, at altitudes to 800
m. Usually rare, as scattered trees, but occasionally locally common, especially on volcanic
soils in E Sabah lowlands. Probably endangered.
20. Dipterocarpus grandiflorus (Blanco) Blanco Plate 2C.

(Latin, grandis = large, florus = flower; with large flowers)
Fl. Filip. ed. 2 (1845) 314, Fl. Filip. ed. 3, 2 (1878) 218; King op. cit. 95; Merrill, Sp. Blancoan.
(1918) 268, op. cit. (1921) 398, op. cit. (1929) 201; Slooten op. cit. (1927) 333; Masamune op. cit.
486; Symington op. cit. (1943) 178; Meijer & Wood op. cit. 252; Burgess op. cit. 100; Ashton op. cit.
(1982) 317; PROSEA op. cit. (1993) 180; Newman et al. op. cit. (1998) 99. Basionym: Mocanera
grandiflora Blanco, Fl. Filip. ed. 1 (1837) 451. Neotype (designated here): Merrill Sp. Blancoan. 119
(= US 903794), the Philippines, Bataan Province, Luzon, Mt. Lamao (K; US). Synonyms:
Dipterocarpus blancoi Blume op. cit. (1852) 35; Vatica trigyna Griff., Notul. 4 (1854) 514; D.
108
motleyanus Hook.f., Trans. Linn. Soc. 23 (1860) 159; D. griffithii Miq., Ann. Mus. Bot. Lugd.Bat. 1
(1864) 213; D. pterygocalyx Scheff., op. cit. 347.
Mediumsized, occasionally large, emergent tree, to 45 m tall, to 1.5 m diameter; crown

dense, dark, irregular. Bark pale greyish to yellowish brown, appearing smooth but
becoming irregularly flaky. Leaf bud, outside of stipules, parts exposed in bud, ovary apex,
and sometimes twig densely evenly pale buffpubescent; parts otherwise glabrous. Twigs
terete, to 12 mm diameter apically, stout; internodes short, the leaves clustered round the
ends. Leaf buds ovoid, acute, to 20 × 10 mm. Stipules oblonglanceolate, to 18 × 5 cm.
Leaves thickly coriaceous, hardly corrugated, glabrous, drying pinkish brown; blade
broadly ovate, 10–18 × 5–12 cm, base obtuse to subcordate, apex with acumen to 1 cm
long; lateral veins 15–17 pairs, prominent below, spreading; petiole to 9 cm long, at least 2
mm diameter, stout. Inflorescences to 18 cm long, unbranched, slender, borne in dense
groups behind the leaves. Flowers: buds to 3.5 × 1.3 cm; stamens c. 30, anthers linear
lanceolate, tapering. Fruits: pedicel stout; calyx tube glabrous, ellipsoid, to 7 × 3.5 cm, with
5, to 15 mm wide prominent coriaceous pruinose flanges from base to apex; 2 major calyx
lobes to 22 × 3 cm, 3 minor ones elliptic, to 2 × 1.5 cm, subrevolute.
Vernacular name. Sabah—keruing belimbing (preferred name).
Distribution. Andamans, Myanmar, Thailand, Vietnam, Sumatra, Peninsular Malaysia,

Borneo, and the Philippines. In Borneo known only in Sabah and recorded from Kota
Kinabalu, Kudat, Lahad Datu, Papar, Sandakan, and Semporna districts (e.g., SAN 18854,
SAN 19749, SAN 27192, SAN A 287, and SAN A 1247).
Ecology. Locally common on dry ridges and low hills, at altitudes below 400 m, within 150
km of the coast; spreading inland, especially on ridges, in drier climates. Vulnerable.
21. Dipterocarpus hasseltii Blume

(J.C. van Hasselt, 1797–1823, a Dutch botanist)
Fl. Java 2 (1829) 22; Slooten op. cit. (1927) 280, op. cit. (1940) 436; Foxworthy op. cit. (1932) 67;
Masamune op. cit. 486; Symington op. cit. (1943) 180; Backer & Bakhuizen f. op. cit. 329; Meijer &
Wood op. cit. 255; Ashton op. cit. (1978) 8, op. cit. (1982) 306; PROSEA op. cit. 181; Kessler &
Sidiyasa op. cit. 94; Newman et al. op. cit. 101. Type: van Hasselt s.n. (= RHL Sheet Nos.
902146210–211), Java, Banten, Menes (holotype L). Synonyms: Dipterocarpus quinquegonus Blume
op. cit. (1852) 36; D. balsamiferus Blume op. cit. (1852) 37, Merrill op. cit. (1921) 397, Masamune
op. cit. 485; D. lampongus Scheff., op. cit. 146; D. subalpinus Foxw. in Elmer, Leafl. Philip. Bot. 6
(1913) 1950. (For other synonyms, cf. Ashton op. cit. (1982) 306).
Stately emergent tree, to 55 m tall, to 2 m diameter, with dense hemispherical crown and tall
buttresses. Bark overall smooth, thinly flaky, greyish brown. Parts exposed in flower bud
and ovary apex buffpubescent; veins below sparsely so, usually caducous; parts otherwise
glabrous. Twigs c. 4 × 2 mm apically, somewhat compressed, drying black. Leaf buds
falcate, to 20 × 5 mm, glabrous, drying black. Stipules loratelanceolate, c. 12 × 1 cm,
glabrous. Leaves thinly coriaceous, hardly corrugated, glabrous, drying dull greyish
brown; blade narrowly elliptic, 9–16 × 5–10 cm, base cuneate, margin often prominently
wavy, apex with acumen to 1 cm long; lateral veins 11–14 pairs, slender but prominent
below, ascending; intercostal venation densely regularly scalariform; petiole 2.5–4 cm long,
slender. Inflorescences to 10 cm long, axillary, hardly branched. Flowers: bud c. 3 × 1 cm;
stamens c. 30, anthers linear, somewhat tapering. Fruits: pedicels to 3 mm long; calyx tube
109
globose, to 3 cm diameter, smooth; 2 major calyx lobes to 22 × 3 cm, 3 minor ones

suborbicular, to 1.5 × 0.3 cm, somewhat revolute.
Vernacular name. Sabah—keruing kerukup kecil (preferred name).
Distribution. Peninsular Thailand, Sumatra, Peninsular Malaysia, W Java, Nusa Tenggara

(Bali Is.), Borneo, and the Philippines. In Borneo recorded from Kinabatangan, Pensiangan,
Sandakan, and Tawau districts in Sabah (e.g., FMS 55342, KEP 70522, SAN 15517, SAN
18464, and SAN 23596) and E Kalimantan (e.g., Kostermans 4886 and Kostermans 4956).
Ecology. In mixed dipterocarp forest on lower slopes and moist (but not swampy) valleys,
especially on fertile soils over basic volcanic rock, at altitude to 500 m. Critically
endangered owing to forest conversion.
22. Dipterocarpus humeratus Slooten Fig. 7.

(Latin, humerus = shoulder; the articulated petiole)
Bull. Jard. Bot. Buitenz. 3, 8 (1927) 308; Ashton op. cit. (1963) 237, op. cit. (1964) 36, op. cit. (1968)
15, op. cit. (1982) 311; Meijer & Wood op. cit. 257; Burgess op. cit. 100; Anderson op. cit. (1980)
112; PROSEA op. cit. 181; Coode et al. (eds.) op. cit. 68; Newman et al. op. cit. 102. Lectotype
(designated here): bb. E 1150, Sumatra, Palembang, Lematang Hilir, near G. Megang (hololectotype
BO; isolectotype K).
Emergent tree, to 50 m tall, to 1 m diameter; buttresses to 2 m tall. Bark pinkish brown,

irregularly flaking. Young twig, petiole, midrib and vein below cream puberulent; leaf bud
and stipule outside shaggily fulvous hispid; ovary and raceme goldenbrown pubescent.
Twigs c. 1 cm diameter apically, stout, becoming thinly flaky. Leaf buds narrowly conical,
to 50 × 15 mm, obtuse. Stipules narrowly lanceolate, c. 10 × 2 cm. Leaves thickly
coriaceous, corrugated, drying chocolatebrown, sparsely pubescent below; blade broadly
ovate, 20–38 × 12–23 cm, base obtuse, apex obtuse to shortly acuminate; lateral veins c. 20
pairs, prominent, wellspaced; petiole 4–6 cm long, stout, prominently geniculate.
Inflorescences to 21 cm long, terminal or axillary, hardly or not branched. Flowers: buds c.
5 × 2 cm; stamens c. 40, anthers linear, tapering. Fruits subsessile; calyx tube globose, to 4
× 3.5 cm, with 5 obtuse tubercles distally; 2 major calyx lobes to 18 × 5 cm, 3 minor ones
broadly ovate, to 1.5 × 1.5 cm, subcordate, with obtuse apex, revolute back to back.
Vernacular names. Sabah—keruing kerukup (preferred name). Sarawak—keruing latek

bukit (preferred name).
Distribution. Sumatra and Borneo. In Sabah known from Kinabatangan, Ranau, Sandakan,
and Tawau districts (e.g., FMS 48854, SAN 15393, SAN 18568, and SAN A 4142) and in
Sarawak from Lawas, Limbang and Tatau districts (e.g., BRUN 805, S 24955 and S 30229).
Also occurring in Brunei (e.g., BRUN 329, BRUN 330 and BRUN 2618) and SE Kalimantan
(e.g., bb. 26595).
Ecology. In mixed dipterocarp forest on welldrained clay soils; on low hills in eastern
Sabah, and high shale ridges at 200–700 m altitude in Sarawak and Brunei. Occurring in
Mulu NP; probably vulnerable elsewhere.
110
Fig. 7. Dipterocarpus humeratus. A, leafy twig; B, detail of indumentum on lower leaf

surface; C, flower bud; D, longitudinal section of open flower; E, adaxial view of stamens;
F, abaxial view of stamens; G, side view of almost mature fruit; H, side view of mature fruit
with the larger calyx lobes removed; I, basal view of mature fruit. (A–B from FMS 48864, C
from SAN 15393, D–F from FMS 38752, G from SAN 17327, H–I from SAN 31397.)
111
23. Dipterocarpus kerrii King

(A.F.G. Kerr, 1877–1942, introduced western medical practise to Siam, prodigious amateur
botanist)
J. As. Soc. Beng. 62, 2 (1893) 93; Ridley op. cit. (1922) 215; Slooten op. cit. (1927) 295; Foxworthy
op. cit. (1932) 69; Symington op. cit. (1943) 181; Meijer & Wood op. cit. 259; Burgess op. cit. 101;
Ashton op. cit. (1982) 305; PROSEA op. cit. 182; Newman et al. op. cit. 103. Lectotype (designated
here): Kerr 7438/7349, S Thailand, Pattani (hololectotype K). Synonyms: Dipterocarpus obconicus
Foxw. in Elmer op. cit. (1913) 1951; D. perturbinatus Foxw., op. cit. (1918) 177; D. cuneatus Foxw.,
op. cit. (1918) 178.
Large emergent tree, to 50 m tall, to 1.2 m diameter; crown dense, compact, dark. Bark pale
greyish to yellowish brown, thinly irregularly flaky. Parts of petals exposed in bud, inside of
bud scales and stipules, and ovary densely silky cream pubescent; parts otherwise glabrous.
Twigs c. 3 mm diameter apically, drying black. Leaf buds lanceolatefalcate, to 12 × 3 mm,
glabrous, drying black. Stipules linearlanceolate, c. 8 × 0.5 cm. Leaves coriaceous, drying
dark chocolatebrown, glabrous below; blade broadly elliptic, 8–13 × 3.3–7 cm, base
cuneate, apex with acumen less than 0.5 cm long; lateral veins (7–)9–11 pairs, slender but
prominent below, ascending; intercostal venation densely scalariform, very slender, hardly
raised; petiole 2–2.8 cm long, slender. Inflorescences to 8 cm long, unbranched or singly
branched. Flowers: buds c. 2.5 × 1 cm; stamens c. 30, anthers subauriculate, narrowly
deltoid. Fruits: pedicels to 3 mm long, stout; calyx tube globose to turbinate, to 3.5 cm
diameter, smooth; 2 major calyx lobes to 14 × 3 cm, 3 minor ones suborbicular, to 1 × 1
cm, subrevolute.
Vernacular name. Sabah—keruing gondol (preferred name).
Distribution. Andamans, Myanmar, Peninsular Thailand, Sumatra, Peninsular Malaysia,

Borneo, and the Philippines. In Borneo only known in the eastern parts of Sabah and
recorded from Kinabatangan, Labuk Sugut and Sandakan districts (e.g., SAN 16247, SAN
16329, SAN 16350, SAN 96959, and SAN 97202).
Ecology. In mixed dipterocarp forest on welldrained clay soils on low hills, at altitudes to
400 m, near the coast in aseasonal climates. Locally frequent in scattered localities down the
east coast of Sabah. Highly vulnerable, possibly endangered.
24. Dipterocarpus kunstleri King

(H.H. Kunstler, 1837–1887, King’s plant collector in Perak)
J. As. Soc. Beng. 62, 2 (1893) 96; Ridley op. cit. (1922) 217; Slooten op. cit. (1927) 327; Symington
op. cit. (1943) 182; Ashton op. cit. (1978) 10, op. cit. (1982) 309; PROSEA op. cit. 182; Coode et al.
(eds.) op. cit. 69; Newman et al. op. cit. 104. Type: King’s collector 3798, Peninsular Malaysia,
Perak, Larut (holotype K). Synonyms: Dipterocarpus speciosus Brandis, J. Linn. Soc. Bot. 31 (1895)
38; D. exalatus Slooten ex Wood, Gard. Bull. Sing. 17 (1960) 486, Slooten op. cit. (1961) 462, Ashton
op. cit. (1964) 31, op. cit. (1968) 14, Meijer & Wood op. cit. 246, Burgess op. cit. 100, Anderson op.
cit. (1980) 112.
Main canopy tree, to 40 m tall, to 1 m diameter; bole relatively short. Bark pale orange
brown, patchily thinly flaky. Leaf venation puberulent or glabrous below; leaf buds and
stipule outside persistently densely pale grey adpressed puberulent; parts otherwise
glabrous. Twigs slender, to 5 mm diameter apically, often slightly ribbed or compressed,
112
with prominent stipule scars. Leaf buds narrowly falcate, to 15 × 3.5 mm. Stipules linear, to
7 × 0.8 cm. Leaves chartaceous, glabrous, drying greyish brown below; blade elliptic to
broadly lanceolate, 13–22 × 7–10 cm, base cuneate, apex shortly acuminate (mature trees);
lateral veins 16–18 pairs, slender but prominent below, ascending; petiole 2–3 cm long,
slender, to 2 mm diameter. Inflorescences to 22 cm long, singly branched, terminal or
axillary. Flowers: buds c. 3.5 × 1.5 cm; stamens c. 30, anthers narrow, tapering. Fruits:
calyx tube ellipsoid, to 5 × 2.5 cm, tapering gradually to the base and to the constricted
neck, 5ribbed or almost winged, the ribs c. 7 mm wide and 4 mm thick apically, either
confined to the distal half or tapering to the base, terminating apically as obtuse tubercles;
2 major calyx lobes to 11 × 1.5 cm, of variable length and sometimes subequal with minor
lobes, 3 minor ones to 0.6 × 0.5 cm, recurved, thickened.
Vernacular names. Sabah—keruing rapak (preferred name). Sarawak—keruing kuntum

putih (preferred name).
Distribution. Sumatra, Peninsular Malaysia, Borneo, and the Philippines. In Borneo known
in Sabah from Beaufort, Kinabatangan, Lahad Datu, Papar, Ranau, Sandakan, Sipitang, and
Tawau districts (e.g., FMS 38783, SAN 16037, SAN 17177, SAN 21273, SAN 38149, and
SAN 97486) and in Sarawak from Kapit, Lundu, Limbang, Marudi and Miri districts (e.g., S
1813, S 7959, S 22330, S 23330, and S 23351). Also occurring in Brunei (e.g., BRUN 138,
BRUN 298, BRUN 903, and BRUN 3075) and E and SE Kalimantan (e.g., bb. 18424, bb.
20644, bb. 25609, Kostermans 7254, and Kostermans 9745).
Ecology. Locally frequent in mixed dipterocarp forest on leached silty clay soils, mainly in
moist valleys and lower hillsides, at altitudes to 400 m. Most abundant in E Sabah, in
scattered localities elsewhere. Highly vulnerable though occurring in Bako and Mulu NPs.
25. Dipterocarpus lamellatus Hook.f.

(Latin, lamellatus = thinly plated; the fruit calyx tube flanges)
Trans. Linn. Soc. 23 (1860) 159; Merrill op. cit. (1921) 399; Slooten op. cit. (1927) 347; Masamune
op. cit. 487; Meijer & Wood op. cit. 261; Burgess op. cit. 101; Ashton op. cit. (1982) 312; Coode et al.
(eds.) op. cit. 69; Newman et al. op. cit. 105. Type: Motley 159, Borneo, Labuan (holotype K).
Tall emergent tree, to 55 m, to 1.2 m diameter; crown golden from below. Exposed living
parts and ovary persistently yellowish brown scabridhirsute, leaf blade and fruit calyx
sparsely so; parts otherwise densely scabridhirsute. Twigs terete, c. 4 mm diameter
apically. Leaf buds ovoid, to 9 × 7 mm, acute. Stipules lanceolate, c. 3 × 0.8 cm. Leaves
chartaceous, boatshaped and concave; blade elliptic, 13–16 × 6–9 cm, base narrowly
obtuse, apex with abrupt, slender acumen to 0.8 cm long; lateral veins 15–17 pairs, slender
but prominent below, shallowly sunken above; intercostal venation distinctly elevated
below, evident above; petiole 3–4 cm long, slender. Inflorescences to 8 cm long, axillary,
hardly branched. Flowers unknown. Fruits: pedicels to 4 mm long, slender; calyx tube
subglobose, to 1.8 cm long including the 5 densely convoluted flanges; 2 major calyx lobes
to 14 × 2.5 cm, 3 minor ones ovate, to 1.4 × 0.7 cm, revolute.
Vernacular name. Sabah—keruing jarang (preferred name).
113
Distribution. Endemic in Borneo; known only from Beaufort Hill FR and Labuan, in SW
Sabah (e.g., SAN 16251 and the type) and Ladan Hills, Tutong district in Brunei (e.g., Wong
1652).
Ecology. With Shorea dispar one of the rarest and most endangered of all dipterocarps.
Found in mixed dipterocarp forest on yellow sandy soil on hills, at altitudes below 200 m,
within 50 km of the coast. Probably extinct in Sabah due to the loss of its habitat.
26. Dipterocarpus lowii Hook.f. Fig. 8.

(Sir Hugh Low, 1824–1905; Officer of the British East India Company, plant collector)
Trans. Linn. Soc. 23 (1860) 160; Merrill op. cit. (1921) 399, op. cit. (1929) 201; Slooten op. cit.
(1927) 433; Masamune op. cit. 487; Symington op. cit. (1943) 183; Browne op. cit.110; Ashton op.
cit. (1964) 37, op. cit. (1968) 15, op. cit. (1982) 313; Meijer & Wood op. cit. 261; Burgess op. cit. 101;
cit. 106. Type: Lowe s.n., Borneo, Labuan (holotype K). Synonym: Dipterocarpus undulatus Vesque
op. cit. 153.
Large emergent tree, to 55 m tall, to 2 m diameter; crown spreading, rather flat, diffuse,
largeleaved; buttresses to 2.5 m tall, prominent. Bark chocolatebrown, thinly evenly flaky
and vertically cracked. Twigs, midrib and vein below more or less densely caducously pale
goldenyellow pubescent; exposed parts of buds, ovary, stipule outside, and inflorescence
persistently so. Twigs to 10 × 3 mm across apically, compressed, stout, with broad swollen
stipule scars. Leaf buds conical to falcate, c. 15 × 9 mm, obtuse, pale goldenyellow
pubescent. Stipules deltoid, to 4 × 3 cm. Leaves thickly coriaceous, prominently
corrugated, drying pale brown and more or less glabrous below; blade ovatelanceolate,
15–20 × 6–10 cm, base obtuse to cordate, margin prominently revolute, apex obtuse to
narrowacuminate with an acumen to 0.6 cm long; lateral veins 15–20 pairs, prominent and
glabrous below; petiole 1.5–3 cm long, stout. Inflorescences to 3 cm long, stout, terminal or
axillary, simple or singly branched. Flowers: buds to 4 × 1.2 cm; stamens c. 30, anthers
narrowly oblong. Fruits: calyx tube globose, c. 4 × 4 cm, including the 5 intricately
convoluted flanges; 2 major calyx lobes to 14 × 3.5 cm, 3 minor ones broadly ovate, to 2 × 2
cm, recurved and concurrent with flanges.
Vernacular name. Sabah and Sarawak—keruing sol/shol (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo, known in Sabah from
Beaufort, Kinabatangan, Lahad Datu, Sipitang, Tawau, and Tenom districts (e.g., SAN
15013, SAN 15119, SAN 36004, SAN 37834, and SAN 40611) and in Sarawak from Baram,
Bintulu, Kuching, Lawas, Limbang, and Lundu districts (e.g., S 1826, S 9097, S 10068, S
16944, and S 42981). Also occurring in Brunei (e.g., BRUN 3285, FMS 35657 and Wong
WKM 1610) and C, SE and E Kalimantan (e.g., Argent 9485, bb. 18329 and Meijer 2552).
Ecology. Local but common where it occurs, in mixed dipterocarp forest on low
sedimentary hills on yellow sandy soils, at altitudes to 400 m. In many localities in Sabah
and Sarawak, it also occurs on ultrabasic rocks. Locally frequent in Lambir and Mulu NPs,
114
Fig. 8. Dipterocarpus lowii. A, fruiting leafy twig; B, flower bud; C, longitudinal section of
flower bud; D, adaxial view of stamens; E, abaxial view of stamens; F, longitudinal section
of fruit. (A from SAN 40611, B–E from S 10068, F from S 1826.)
115
27. Dipterocarpus mundus Slooten

(Latin, mundus = elegant; the neat appearance of the glabrous parts)
Bull. Jard. Bot. Buitenz. 3, 16 (1940) 446; Masamune op. cit. 487; Ashton op. cit. (1968) 15, op. cit.
(1982) 319; Anderson op. cit. (1980) 112; PROSEA op. cit. 183; Newman et al. op. cit. 107.
Lectotype (here designated): bb. 26033, Kalimantan, Melawi, Bt. Ransah, Catit (hololectotype BO;
isolectotype KEP).
Emergent tree, to 50 m tall, to 1 m diameter. Parts glabrous but for the inner surface of
stipules. Twigs 2–3 mm diameter apically, slender. Leaf buds linearfalcate, to 14 × 3 mm,
glabrous outside, drying black. Stipules linear, to 4 × 0.6 cm. Leaves thinly coriaceous,
drying rich brown below; blade narrowly elliptic to obovate, 5.5–16 × 2.5–7.5 cm, base
narrowly obtuse, apex with tapering acumen to 0.8 cm long; lateral veins 8–10 pairs,
slender but prominent below; petiole 1.6–3 cm long, slender. Inflorescences to 6 cm long,
axillary, unbranched. Flowers: buds to 2 × 0.8 cm; stamens c. 15, anther narrowly oblong.
Fruits: pedicels c. 3 mm long; calyx tube glabrous, narrowly ellipsoid or narrowly obovoid,
to 3 × 1.2 cm, with 5, rigid flanges to 7 mm wide, widest in the distal half; 2 major calyx
lobes to 11 × 2.7 cm, 3 minor ones broadly ovoid, to 0.8 × 0.8 cm, obtuse, somewhat
recurved.
Vernacular name. Sarawak—keruing matang (preferred name).
Distribution. Endemic in Borneo and known only from the central mountains of the Kapuas
and Rajang hinterlands. In Sarawak recorded from Kapit and Miri districts (e.g., Ashton
s.n., S 19046, S 29648, S 29649, and S 57980). Also occurring in W Kalimantan (e.g., bb.
27022, bb. 28135 and bb. 29650).
Ecology. In Sarawak, scattered in mixed dipterocarp forest in small groups along the narrow
sandstone and shale ridges of the Rajang Series, on shallow clayrich soils, at 400–600 m
altitude. Vulnerable.
28. Dipterocarpus nudus Vesque

(Latin, nudus = naked; the glabrous parts)
C. R. Ac. Sci. Paris, 78 (March 1874) 150; Merrill op. cit. (1921) 399; Slooten op. cit. (1927) 325;
Masamune op. cit. 487; Browne op. cit. 110; Ashton op. cit. (1964) 38, op. cit. (1968) 16, op. cit.
(1982) 320; Anderson op. cit. (1980) 113; Coode et al. (eds.) op. cit. 69; Newman et al. op. cit. 108.
Type: Beccari PB 2905, Borneo, Sarawak, G. Matang (holotype P). Synonym: Dipterocarpus
pentapterus Dyer, J. Bot. 12 (April 1874) 106 & 152.
Emergent tree, to 50 m tall, to 1.3 m diameter; bole frequently malformed; crown dark and
dense, somewhat flat. Bark typical. Parts glabrous but for the buffpubescent ovary apex
and inner surface of stipules. Twigs 2–3 mm diameter apically, slender. Leaf buds linear
falcate, 15–30 × 2–3 mm, drying black. Stipules linear, to 6 × 0.5 cm. Leaves thinly
coriaceous, hardly corrugated, drying chocolatebrown below; blade more or less narrowly
elliptic, 11–14 × 4–6 cm, base cuneate or narrowly obtuse, margin undulate and somewhat
sinuate distally, apex with slender acumen to 0.6 cm long; lateral veins (8–)11–14 pairs,
dense, ascending, slender but prominent below; intercostal venation widely scalariform to
subreticulate; petiole 2.5–4.5 cm long, slender. Inflorescences to 18 cm long, terminal or
axillary, singly branched, zigzag. Flowers: buds to 4 × 0.9 cm, slender; stamens c. 15,
anther narrowly oblong, tapering. Fruits: calyx tube glabrous, narrowly ellipsoid to
116
spindleshaped, c. 2.5 × 1 cm, with 5, rigid flanges to 3 mm wide; 2 major calyx lobes to 9.5
× 3 cm, 3 minor ones c. 0.4 × 0.4 cm, obtuse, subrevolute.
Vernacular name. Sarawak—keruing licin (preferred name).
Distribution. Endemic in Borneo. Known in Sarawak from Bintulu, Kapit, Kuching,

Mukah, Limbang, Lundu, Simunjan, Sri Aman, and Tatau districts (e.g., S 6501, S 9481, S
773, BRUN 3095, and BRUN 3193).
Ecology. Widespread but uncommon in mixed dipterocarp forest on hills, at altitudes to 600
m, on yellow sandy soils and on shallow shale derived silty clays. Vulnerable.
Notes. In the western part of its range this species has broader more coriaceous leaves, with
narrowly obtuse base and 8–12 pairs of lateral veins (as compared with 12–14 pairs of
lateral veins in its eastern range). They differ from one another almost as much as both do
from D. mundus.
29. Dipterocarpus oblongifolius Blume Plates 2D–F.

(Latin, oblongus = rather long, folius = leaf; with oblong leaves)
Mus. Bot. Lugd.Bat. 2 (1852) 36; King op. cit. 95; Merrill op. cit. (1921) 399; Ridley op. cit. (1922)
216; Slooten op. cit. (1927) 338; Masamune op. cit. 487; Symington op. cit. (1943) 184; Browne op.
cit. 110; Ashton op. cit. (1964) 39, op. cit. (1968) 16, op. cit. (1982) 317; Meijer & Wood op. cit. 364;
Burgess op. cit. 101; Anderson op. cit. (1980) 113; PROSEA op. cit. 183; Coode et al. (eds.) op. cit.
69; Newman et al. op. cit. 109. Type: Korthals s.n. (= RHL Sheet Nos. 902146156–159), Borneo,
Kalimantan (holotype L). Synonyms: Dipterocarpus stenopterus Vesque op. cit. 625; D. pulcherrimus
Ridl., Trans. Linn. Soc. Bot. 3 (1893) 283.
Stout trunked leaning tree, to 30 m tall, to 1.5 m diameter; bole elliptic in crosssection
towards base, branching low; crown spreading, with hanging branch endings, dark green
shiny leaves and conspicuous applered young leafy shoots. Bark pale greyish brown, thinly
irregularly flaked. Exposed living parts and ovary, leaf surface and veins above excepted,
densely evenly pale creamyellow pubescent, persistent only on stipule outside, calyx and
ovary. Twigs 2–3 mm diameter apically, slender. Leaf buds linear, c. 20 × 3 mm,
compressed, glabrous or densely tomentose, drying yellowish brown or black. Stipules
lorate, to 15 × 1.5 cm, obtuse. Leaves thinly coriaceous, glabrous, drying greyish brown;
blade narrowly elliptic to lanceolate, 14–18(–25 in juveniles) × 4–7(–9) cm, base cuneate,
apex tapering to 1 cm slender acumen; lateral veins 16–20 pairs, slender; petiole 1.7–2 cm
long, slender. Inflorescences to 18 cm long, terminal or axillary, simple or singly branched.
Flowers: buds c. 4 × 1 cm; stamens c. 15, anthers narrowly oblong, tapering. Fruits:
pedicels to 2 mm long; calyx tube densely pale buffpubescent, narrowly ovoid to spindle
shaped, to 3 × 0.9 cm, only slightly constricted at neck, with 5, c. 1 mm wide wavy flanges
from base to apex; 2 major calyx lobes to 12 × 1.5 cm, 3 minor ones narrowly deltoid, c. 1 ×
0.3 cm, recurved.
Vernacular names. Sabah—keruing neram (preferred name). Sarawak—ensurai (Iban),

gansurai (Iban), laran (Kayan).
117
Distribution. Peninsular Thailand, Peninsular Malaysia and Borneo. In Sabah widespread

and known from Beaufort, Beluran, Labuk Sugut, Ranau, Sandakan, Sipitang, Tawau, and
Tenom districts (e.g., SAN 5498, SAN 6726, SAN 24959, SAN 69924, and SAN 95397) and
in Sarawak from Belaga, Betong, Bintulu, Kapit, Lawas, Limbang, Marudi, Miri, Song, and
Sri Aman districts (e.g., S 15451, S 29568, S 56490, S 57497, and S 64888). Also occurring
in Brunei (e.g., BRUN 353, BRUN 401, FMS 39607, KEP 80118, and Wong WKM 1156)
and Kalimantan (e.g., bb. 20633, Kostermans 12664, Mogea 3548).
Ecology. Gregarious along the banks of white water rivers, on stable soils over bedrock,
below flood and above normal river level, at altitudes to 400 m. Frequently flowering and
fruiting, the masses of pink calyces a glory of the inland rivers. Occurring in Mulu NP;
decimated in many areas outside the park but not yet vulnerable.
30. Dipterocarpus ochraceus Meijer

Acta Bot. Neerl. 12 (1963) 351; Meijer & Wood op. cit. 26; Burgess op. cit. 100; Ashton op. cit.
(1982) 325; Newman et al. op. cit. 110. Type: Meijer SAN 24200, Borneo, Sabah, Ranau district, Bt.
Kulang (holotype K; isotypes KEP, L, SAN).
Mediumsized emergent (large for its habitat) tree, to 45 m tall, to 1.2 m diameter. Bark
rustbrown, becoming coarsely irregularflaky. Young parts densely ochreous velutinous,
persistent on buds, stipule outside and twigs; becoming sparse on petiole and venation
below; caducous elsewhere. Twigs c. 4 mm diameter apically, stout. Leaf buds ovoid
lanceolate, to 20 × 8 mm. Leaves coriaceous, corrugated, sparsely pubescent and drying
pale yellowish brown below; blade broadly ellipticovate, 10–19 × 4.5–9.5 cm, base cuneate
or rarely obtuse, apex acute with acumen to 1.5 cm long; lateral veins 11–15 pairs, slender
but prominent below; intercostal venation dense, elevated below; petiole 1.5–2.5 cm long.
Inflorescences and flowers unknown. Fruits: pedicels to 3 × 2 mm, prominent; calyx tube
glabrescent, broadly ellipsoid, to 1.8 × 1.5 cm, with 5, to 2 mm wide continuous rigid
flanges; 2 major lobes to 8 × 1.5 cm, 3 minor ones ovate, to 0.6–3 × 0.6 cm, subrevolute.
Vernacular name. Sabah—keruing ranau (preferred name).
Distribution. Endemic in Borneo; recorded only in Sabah from Bt. Kulong and Bt.
Tampurango, Ranau district (e.g., SAN 20670, SAN 22353, SAN 22371, and the type).
Ecology. In scattered groups along ridges in upper dipterocarp forest, at 600–700 m altitude,
on the ultrabasic and basic volcanic mountains of the Kinabalu flanks but mainly outside the
National Park boundary. Endangered.
31. Dipterocarpus pachyphyllus Meijer

(Greek, pachys = thick, phullon = leaf; with thick leaf texture)
(1964) 41, op. cit. (1968) 16, op. cit. (1982) 313; Anderson op. cit. (1980) 113; Coode et al. (eds) op.
cit. 69; Newman et al. op. cit. 111. Lectotype (designated here): G.H.S. Wood SAN 15100, Borneo,
Sabah, Padas Gorge (hololectotype K; isolectotypes KEP, L, SAN).
118
Emergent tree, to 45 m tall, to 1.2 m diameter; crown becoming flat; buttresses to 1 m tall.
Bark pale chocolatebrown, vertically cracked and evenly flaky. Young parts at first evenly
shortly pale yellowish brown pubescent, fugaceous on all but exposed parts in bud, outside
of stipules, ovary apex, and inflorescence. Twigs to 4 × 2.5 mm apically, compressed,
slender, with prominently raised stipule scars. Leaf buds linearfalcate, c. 18 × 4 mm.
Stipules narrowly deltoid, c. 4 × 1.2 cm. Leaves thickly coriaceous, hardly corrugated,
drying yellowish brown and glabrous below, somewhat shiny; blade broadly ovate, 9–17 ×
5–9 cm, base obtuse to subcordate, margin hardly revolute, apex with broad acumen to 1 cm
long; lateral veins 10–12 pairs, distant, prominent below, coalescing to an indistinct
intramarginal vein; petiole 2–3.4 cm long, to 2 mm diameter, slender, prominently
geniculate. Inflorescences to 5 cm long, terminal or axillary, simple. Flowers: buds to 1.5 ×
0.5 cm; stamens 23–25, anther narrowly oblong. Fruits: calyx pruinose, tube subglobose, to
2.5 × 2 cm, including the 5 tightly convoluted flanges; 2 major calyx lobes to 13 × 3 cm, 3
minor ones broadly deltoid, to 1 × 1.5 cm, recurved.
Vernacular names. Sabah—keruing daun tebal (preferred name). Sarawak—keruing sol

padi (preferred name).
Distribution. Endemic in Borneo. In Sabah widespread and known from Beaufort,

Keningau, Kinabatangan, Lahad Datu, Sandakan, Sipitang, Tawau, and Tenom districts
(e.g., KEP 80471, SAN 15053, SAN 16433, SAN 16473, and SAN 93675) and in Sarawak
from Bintulu, Kapit, Marudi, Miri and Tatau districts (e.g., Hotta 15812, S 22032, S 22043,
and S 22715). Also occurring in Brunei (e.g., BRUN 365 and S 1671).
Ecology. Scattered in mixed dipterocarp forest on leached clay soils on low hills and slopes,
at altitudes to 400 m. Occurring in Mulu NP, elsewhere vulnerable.
32. Dipterocarpus palembanicus Slooten

(of Palembang, Sumatra)
Bull. Jard. Bot. Buitenz. 3, 8 (1927) 336; Symington op. cit. (1943) 185; Ashton op. cit. (1968) 16, op.
cit. (1982) 318; PROSEA op. cit. 184; Newman et al. op. cit. 113. Lectotype (Ashton, 1978): bb.
T.134, Feb. 1924, Sumatra, Palembang, Lematang Hilir, near G. Megang (hololectotype BO;
isolectotypes KEP, L).
Emergent tree, to 50 m tall, to 1.2 m diameter; crown compact, dense, hemispherical. Bark
at first rich orangebrown, becoming irregularly flaky. Twigs terete, 2–3 mm diameter
apically, finely cracked and flaky. Leaf buds ellipsoidovoid or broadly ovoid, c. 7 × 4 mm,
obtuse, densely golden to rusttomentose. Stipules narrowly hastate, to 3 cm long. Leaves
thinly coriaceous, somewhat corrugated, drying pinkish brown and glabrous below; blade
elliptic to ovate and 10–14 × 5–9 cm, or oblongovate and 7–11 × 3–6 cm, base obtuse or
cuneate, margin sinuate, apex with slender acumen to 2 cm long; lateral veins 12–14 pairs,
slender but prominent below, rather dense; intercostal venation dense; petiole 2–3 cm or to
1.5 cm long, to 2 mm diameter, pubescent. Inflorescences to 6 cm long, terminal or axillary,
unbranched or singly branched. Flowers: buds c. 3.5 × 1.5 cm; stamens c. 15, anther linear
tapering. Fruits: calyx glabrous, pruinose, tube narrowly ellipsoid, to 3.5 × 1.5 or 5.5 × 2
cm, tapering towards each end, with 5 broad thin flanges from base to apex, obtuse to
subcordate at base or the flanges to 15 mm wide, undulate, auriculate at base and apex; 2
major calyx lobes to 10 × 3.5 cm, revolute and subcordate at base, 3 minor ones broadly
orbicular, to 0.5 × 1 cm, recurved and with revolute undulate margins.
119
Vernacular name. Sabah—keruing palembang (preferred name).
Distribution. Sumatra, Peninsular Malaysia, Singapore and Borneo.
Notes. Two subspecies, viz. subsp. borneensis and subsp. palembanicus are recognised in
Borneo.
Key to subspecies
Twigs, leaf buds and stipules outside densely fulvous hispid; petiole, midrib above and leaf
blade below densely shortpubescent, caducous on twigs and petiole, otherwise persistent.
Leaf blade elliptic to ovate, 10–14 × 5–9 cm; petiole 2–3 cm long. Fruit calyx tube to 3.5 ×
1.5 cm, flanges to 8 mm wide, obtuse to subcordate at base………………………………….
subsp. palembanicus
As the species. In Borneo recorded from Kuching, Kapit and Miri districts in Sarawak
(e.g., S 22678, S 39095, S 46438, S 46529, and S 68426). Locally frequent along ridges
in mixed dipterocarp forest, at altitudes to 600 m. Occurring in Lambir NP, elsewhere
vulnerable.
Twigs, petiole, midrib on both sides and lateral veins below densely reddish brown
pubescent, caducous on twigs, persistent and long on leaf buds, stipules and petiole. Leaf
blade oblong to ovate, 7–11 × 3–6 cm; petiole to 1.5 cm long. Fruit calyx tube to 5.5 × 2
cm, flanges to 15 mm wide, undulate, auriculate at base and apex…………………………...
subsp. borneensis P.S.Ashton
(of Borneo)
Gard. Bull. Sing. 31 (1978) 12, op. cit. (1982) 319; Anderson op. cit. (1980) 113; PROSEA op.
cit. 184; Coode et al. (eds.) op. cit. 69; Newman et al. op. cit. 114. Type: G.H.S. Wood SAN A
1747, Borneo, Sabah, Beaufort district (holotype L; isotypes KEP, SAN).
Endemic in Borneo, and recorded from Beaufort, Kinabatangan, Labuk Sugut,
Sandakan, and Tawau districts in Sabah (e.g., SAN 16452, SAN 17772, SAN 26890, and
SAN 28102) and from Belaga, Bintulu and Kapit districts in Sarawak (e.g., S 14719, S
15101, S 22204, S 23828 and S 32309). Also occurring in Brunei (e.g., BRUN 2617 and
BRUN 5742), and C and E Kalimantan (e.g., Wilkie 94310). In mixed dipterocarp forest
on clayrich soils over sedimentary rocks, on ridges, at altitudes to 600 m. Occurring in
Lambir NP, elsewhere vulnerable.
33. Dipterocarpus rigidus Ridl.

(Latin, rigidus = stiff; the leaf blade)
J. Str. Br. Roy. As. Soc. 82 (1920) 171, op. cit. (1922) 217; Slooten op. cit. (1927) 347, op. cit. (1961)
463; Masamune op. cit. 488; Symington op. cit. (1943) 186; Browne op. cit. 110; Ashton op. cit.
(1968) 16, op. cit. (1982) 310; Anderson op. cit. (1980) 113; PROSEA op. cit. 185; Newman et al. op.
cit. 114. Type: Foxworthy 1190, Peninsular Malaysia, Johor, Endau, Sg. Maoung (holotype K).
Large emergent tree, to 50 m tall, to 1 m diameter; crown golden brown from below, dense.
Bark rustbrown, vertically cracked and shallowly patchily flaked. Twig, petiole, midrib
above, blade undersurface, petal outside, and ovary apex shortly densely evenly ochreous
yellow pubescent, blade above fugaceously so; leaf bud and stipule outside longer
tomentose; parts otherwise glabrous. Twigs c. 8 mm diameter apically, stout. Leaf buds
conical, to 15 × 8 mm, acute. Stipules lorate, c. 7 × 1 cm. Leaves thickly coriaceous,
corrugated; blade ovate, 13–25 × 8–16 cm, base broadly cuneate, apex with broad acumen
120
to 1 cm long; lateral veins 12–16 pairs, with midrib stoutly prominent below, somewhat
sunken above; petiole 3–6 cm long. Inflorescences to 15 cm long, glabrescent, usually
unbranched. Flowers: buds c. 3.5 × 1 cm; stamens c. 24, anthers lineartapering. Fruits:
pedicels c. 10 mm long, stout; calyx tube subglobose, to 5 × 4.5 cm, with 5 distal tubercles
becoming obscure at maturity; 2 major calyx lobes to 18 × 5 cm, 3 minor ones suborbicular,
to 0.8 × 0.8 cm, revolute.
Vernacular name. Sarawak—keruing utap (preferred name).
Distribution. Sumatra (including Riau and Lingga Archipelagoes), Peninsular Malaysia,

Anambas Archipelago, and Borneo. In Sarawak known from Bintulu, Kuching, Lawas, and
Lundu districts (e.g., S 10063, S 15826, S 22056, S 27192, and S 42985). Also occurring in
C and W Kalimantan (e.g., bb. 27741 and Yamada K 9536).
Ecology. In Sarawak, locally common and confined to mixed dipterocarp forest on yellow
sandy soils, on low hills not far from the coast. Endangered by forest conversion though
present in Bako NP.
Notes. Specimens of juvenile trees of D. costulatus and D. globosus are sometimes

tomentose and difficult to distinguish from those of this species.
34. Dipterocarpus sarawakensis F.G. Browne ex Slooten Fig. 9.

(of Sarawak)
Reinwardtia 5 (1961) 465; Ashton op. cit. (1964) 43, op. cit. (1968) 17, op. cit. (1982) 323; Anderson
op. cit. (1980) 69; Coode et al. (eds.) op. cit. 69; Newman et al. op. cit. 116. Type: Mead S 5, Borneo,
Sarawak, Kuching district, Semengoh FR (holotype KEP).
Large emergent tree, to 55 m tall, to 1.2 m diameter; crown hemispherical, with the old
leaves turning coppery red before falling. Bark orangebrown, irregularly flaked. Twigs,
parts exposed in bud, stipule outside, inflorescence, ovary apex, midrib on both surfaces,
and venation below densely persistently goldenbrown scabridtomentose; lateral veins
above and intercostal venation below sparsely so; blade margin setose. Twigs c. 3 mm
diameter apically. Leaf buds ovoid, c. 8 × 3 mm, acute. Stipules narrowly ovate, c. 1.5 cm
long. Leaves thinly coriaceous; blade broadly obovate, 5.5–8 × 3.5–5 cm, flat, margin
straight and revolute towards the cuneate base, sinuate towards the obtuse or retuse apex;
lateral veins 7–8 pairs, distant, ascending; petiole 0.7–1 cm, stout. Inflorescences to 7 cm
long, unbranched or singly branched. Flowers: buds to 2.5 × 1.5 cm; stamens c. 15, anthers
narrowly oblong, relatively short, stout. Fruits subsessile; calyx glabrescent; tube spindle
shaped, to 2.5 × 1.3 cm, broadest towards base, tapering towards the ends, with 5 papery
flanges to 8 mm wide distally, tapering to the base or somewhat above it, tapering abruptly
apically and running into lobes; 2 major calyx lobes to 9 × 2 cm, 3 minor ones narrowly
ovate, to 1.5 × 0.5 cm, acute, revolute.
121
Vernacular name. Sarawak—keruing layang (preferred name).
Distribution. Peninsular Malaysia (East coast, one record) and Borneo. In Sarawak
recorded from Bintulu, Kuching, Lundu, and Mukah districts (e.g., S 9770, S 13713, S
18097, S 23749, and S 27194). Also occurring in Brunei (e.g., BRUN 928, BRUN 5154 and
FMS 37090) and C Kalimantan.
Ecology. Scattered in mixed dipterocarp forest, rarely frequent in isolated localities on

leached sandy and sandy clay soils, on low hills, at altitudes to 400 m. Recorded from Mulu
NP but vulnerable, perhaps endangered elsewhere.
35. Dipterocarpus stellatus Vesque

(Latin, stellatus = starlike; the stellate hairs)
C. R. Ac. Sci. Paris, 78 (March 1874) 626; Merrill op. cit. (1921) 400; Slooten op. cit. (1927) 335, op.
cit. (1961) 465; Masamune op. cit. 488; Browne op. cit. 111; Ashton op. cit. (1963) 239, op. cit.
(1968) 17, op. cit. (1982) 323; Anderson op. cit. (1980) 113; Newman et al. op. cit. 118. Lectotype
(designated here): Beccari PB 2555/2907, Borneo, Sarawak, Matang (hololectotype P). Synonym:
Dipterocarpus nobilis Dyer, J. Bot. 12 (April 1874) 106.
Emergent tree, to 65 m tall, to 1 m diameter; crown somewhat flat, diffuse; buttresses to 1.5
m tall. Bark rich pinkish brown, appearing smooth, with irregular somewhat curled thin
flakes. Young twig, leaf bud, stipule outside, midrib on both surfaces, and petiole densely
persistently long tufted rustbrown scabridtomentose, venation below sparsely so. Twigs
2–7 mm diameter apically, ribbed and cracked, with prominent petiole scars. Leaf buds
spherical, c. 7 × 5 mm. Stipules deltoid, c. 2 × 0.8 cm. Leaves thinly coriaceous, not
concave; blade ovate, 10–25 × 5–16 cm, base cordate or obtuse, margin not revolute, apex
with tapering acumen 0.8–1.4 cm long; lateral veins 12–16 pairs, wellspaced; petiole 2–5
cm long. Inflorescences 10–20 cm long, axillary, unbranched or singly branched. Flowers:
buds to 6 × 2 cm; stamens c. 30; anther lineartapering, Fruits sessile or on short stout
pedicels; calyx pruinose, tube glabrous, obturbinate, to 5 × 2 cm, with the nut enclosed in
the distal half and the base a slender cylinder, with 5, to 12 mm wide thin undulate flanges
from base to apex and concurrent with lobes; 2 major calyx lobes to 12 × 3 cm, 3 minor
ones deltoid, c. 1 × 1 cm, with undulate revolute margin.
Vernacular names. Sabah—keruing bulu (preferred name). Sarawak—keruing gunung

(preferred name).
Distribution. Endemic in the northern parts of Borneo.
Notes. Two subspecies, viz. subsp. parvus and subsp. stellatus, are recognised.
Key to subspecies
Twigs to 7 mm diameter apically. Leaf blade 20–25 × 12–16 cm, base cordate, apex with
acumen to 1.4 cm long; petiole 2–5 cm long. Inflorescences to 20 cm long………………….
subsp. stellatus
In Sarawak recorded from Kuching, Lundu and Serian districts (e.g., S 7992, S 8946, S
10182, and S 15383). Also occurring in W Kalimantan (e.g., Church et al. 1808 and
122
Fig. 9. Dipterocarpus sarawakensis. A, fruiting (young) leafy twig; B, detail of indumentum

on lower leaf surface; C, indumentum on leaf bud, inflorescence and young fruit; D, flower
bud; E, longitudinal section of flower bud; F, open flower; G, adaxial view of stamens; H,
abaxial view of stamens; I, mature fruit; J, basal view of mature fruit calyx tube. (A–C from
SA 1288, D–H from S 23749, I–J from BRUN 5154.)
123
Church et al. 1814). Locally common in hill mixed dipterocarp and upper dipterocarp
forests on sandy clay soils derived from sandstone and siliceous acid igneous rocks, at
altitudes to 800 m. Recorded from G. Gading and Kubah NPs; probably not vulnerable.
Twigs to 2 mm diameter apically. Leaf blade 10–15 × 5–7 cm, base obtuse, apex with
acumen to 0.8 cm long; petiole at most 2 cm long. Inflorescences to 10 cm long……………
subsp. parvus P.S.Ashton
(Latin parvus = small; the vegetative and reproductive parts)
Gard. Bull. Sing. 20 (1963) 239, op. cit. (1964) 45, op. cit. (1968) 17, op. cit. (1982) 323; Meijer
& Wood op. cit. 268; Burgess op. cit. 101; Anderson op. cit. (1980) 113; Coode et al. (eds.) op.
cit. 70; Newman et al. op. cit. 119. Type: Ashton BRUN 3176, Borneo, Brunei, Bangar (holotype
K; isotype KEP).
Recorded from Beaufort, Kinabatangan, Sandakan, Sipitang, and Tawau districts in
Sabah (e.g., SAN 15131, SAN 15403, SAN 24837, and SAN A 3126) and from Belaga,
Bintulu, Marudi and Miri districts in Sarawak (e.g., S 3016 and S 22057). Also
occurring in Brunei (e.g., BRUN 3008 and BRUN 3138) and E Kalimantan (e.g.,
Ambriansyah & Arifin Berau 1061, Ambriansyah & Arifin Berau 1089, Kessler et al.
Berau 690, and Kessler et al. Berau 749). Scattered in mixed dipterocarp forest, on
ridges and undulating land, on clay soils derived from shale and acid volcanic rocks, at
altitudes to 700 m. Occurring in Lambir and Mulu NPs but vulnerable and possibly
endangered elsewhere.
36. Dipterocarpus sublamellatus Foxw.

(Latin, sub = somewhat, lamellatus = layered; the flanges of the fruit calyx tube)
Malay. For. Rec. 10 (1932) 92; Slooten op. cit. (1941) 108; Symington op. cit. (1943) 189; Meijer &
Wood op. cit. 269; Ashton op. cit. (1968) 17, op. cit. (1982) 322; Anderson op. cit. (1980) 113;
PROSEA op. cit. 185; Newman et al. op. cit. 119. Type: Watson KEP 6061, Peninsular Malaysia,
Johor, KluangMersing Road (holotype KEP).
Large emergent tree, to 70 m tall, to 3 m diameter; buttresses to 3 m tall and out. Bark dark
orangebrown, thickly patchily flaky. Leaf bud and stipule outside densely long pale fulvous
hirsute (rarely glabrous), ovary shortly so, venation below sparsely caducously so; parts
otherwise glabrous. Twigs terete, c. 3 mm diameter apically, dark with prominent pale
stipule scars. Leaf buds ellipsoidovoid, to 12 × 8 mm, subacute. Stipules lorate, to 4 × 0.8
cm, acute. Leaves coriaceous, corrugated, drying chocolatebrown and glabrous below;
blade ovate to elliptic (young trees), 5–15 × 3.5–8 cm, base broadly cuneate, apex with
short, slender acumen; lateral veins 8–12 pairs, ascending, prominent below; petiole 1.5(–3)
cm long, to 2 mm diameter, slender. Inflorescence: axillary; rachis to 5 cm long, hardly or
not branched, somewhat compressed, fewflowered. Flowers: buds to 3.5 × 1 cm; stamens
c. 24, anther lineartapering. Fruits subsessile; calyx tube glabrescent, globose, to 3 × 3 cm,
with 5, to 15 mm wide thickened but undulate flanges tapering into the lobes and auriculate
at base; 2 major calyx lobes to 12 × 3 cm, 2 minor ones suborbicular, to 0.5 × 0.7 cm,
revolute.
Vernacular name. Sabah and Sarawak—keruing kerut (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo recorded in Sabah from
Sebatik Is., Tawau district (e.g, SAN 19627) and in Sarawak from Bintulu, Kapit and
Kuching districts (e.g., S 6539, S 6542, S 18428, and Sinclair 10296). Also occurring in
Kalimantan (e.g., Argent 94109, Suzuki K 9745 and Wilkie 94314).
124
Ecology. Locally common, on low hills but mostly on periodically flooded sandy clay
alluvium, sometimes with shallow peat, at altitudes below 400 m. Vulnerable owing to land
conversion.
37. Dipterocarpus tempehes Slooten

(tempehes = a local name in E Kalimantan)
Reinwardtia 5 (1961) 468; Meijer & Wood op. cit. 270; Burgess op. cit. 101; Ashton op. cit. (1968)
18, op. cit. (1982) 297; Anderson op. cit. (1980) 113; PROSEA op. cit. 185; Kessler & Sidiyasa op.
cit. 95; Newman et al. op. cit. 120. Type: Rutten 126, Borneo, Kalimantan, Bulungan, Sg. Sajau
(holotype U).
Main canopy, occasionally low emergent tree, to 45 m tall, to 1.2 m diameter; buttresses to
2 m tall, 3 m out. Bark yellowish brown and dark greenish brown, thinly irregularly flaking.
Twig, parts exposed in leaf bud, outside of stipules and petals, raceme, ovary apex, and
petiole shortly densely persistently pale buffpuberulent; leaf venation below and stipule
inside sparsely so; venation above caducously so. Twigs terete, 2–4 mm diameter apically,
slender, stipule scars darker than twig. Leaf buds falcate, to 12 × 3 mm. Stipules lorate, to 8
× 1.2 cm. Leaves thickly coriaceous, corrugated, drying warm rustbrown, sparsely
pubescent or glabrous below; blade broadly elliptic to obovate, 6–12 × 3.5–8 cm, base
broadly cuneate, apex acute to shortly abruptly acuminate; lateral veins 9–12 pairs,
ascending, prominent below; petiole 1–2 cm long, to 2 mm diameter. Inflorescences to 2.5
cm long, axillary, unbranched. Flowers: buds to 3 × 0.8 cm; stamens c. 30, anthers
narrowly oblong, tapering. Fruits subsessile; calyx glabrous; tube turbinate, to 4 × 4 cm,
lenticellate, smooth, with 5 vestigial lobes round the protruding nut apex.
Vernacular names. Sabah—keruing asam (preferred name). Sarawak—keruing tempayan

(preferred name).
Distribution. Endemic in Borneo. Known in Sabah from Labuk Sugut, Lahad Datu,
Sandakan, and Tawau districts (e.g., SAN 17792, SAN 17794 and SAN 37891) and in
Sarawak from Bintulu, Kuching, Lawas, and Lundu districts (e.g., S 14583, S 15823, S
16580, SA 554, and SA 659). Also occurring in S and E Kalimantan (e.g., Argent & Amiril
93104, bb. 10543, bb. 17840, Kostermans 5274, and Kostermans 6211).
Ecology. Local, but where occurring often common, on clayrich periodically flooded
alluvium, at altitudes below 300 m. Vulnerable, possibly endangered by forest conversion.
38. Dipterocarpus validus Blume

(Latin, validus = mighty; a big tree)
Mus. Bot. Lugd.Bat. 2 (1852) 36; Merrill op. cit. (1921) 400; Slooten op. cit. (1927) 272; Masamune
op. cit. 488; Ashton op. cit. (1963) 237, op. cit. (1982) 301; PROSEA op. cit. 185; Newman et al. op.
cit. 122. Type: Korthals s.n., Borneo, Kalimantan, G. Sakumbang (holotype L). Synonyms:
Dipterocarpus warburgii Brandis op. cit. 32, Slooten op. cit. (1927) 305, op. cit. (1961) 473, Meijer &
Wood op. cit. 273, Burgess op. cit. 101; D. affinis Brandis op. cit. 31; D. lasiopodus Perkins, Fragm.
Fl. Philip. (1904) 22; D. woodii Merr., op. cit. (1926) 399, Slooten op. cit. (1927) 303.
125
Emergent tree, to 50 m tall, to 1.8 m diameter; crown rather flat, diffuse, largeleaved;
buttresses to 2.5 m tall. Twig, leaf bud, stipule outside, petiole, base of inflorescence, and
ovary apex densely persistently evenly long rufoustomentose, becoming distinctly tufted as
the twig and petiole expand; lateral veins and midrib below, and rest of inflorescence
sparsely puberulent. Twigs terete, c. 8 mm diameter apically, rather stout. Leaf buds
lanceolatefalcate, to 90 × 13 mm. Stipules lorate, to 20 × 3 cm. Leaves thinly coriaceous,
corrugated, drying warm brown and glabrous below; blade ellipticoblong to ovate, 15–25
× 7.5–12 cm (to 40 × 20 cm in juveniles), base cuneate to obtuse (narrowly peltate in young
trees), margin undulate, prominently sinuate distally, apex acute to narrowly acuminate with
acumen to 1 cm long; midrib elevated above; lateral veins 22–28 pairs, slender but
prominent below, dense, shallowly sunken above; petiole 3.5–5 cm long, at least 2 mm
diameter, stout. Inflorescences to 14 cm long, hardly or not branched, axillary. Flowers:
buds to 3 × 0.9 cm; stamens c. 30, anthers slender, tapering, auriculate. Fruits: calyx tube
turbinate and tapering into pedicel, to 4 × 3.5 cm, smooth; 2 major calyx lobes to 25 × 3.5
cm, 3 minor ones suborbicular, to 0.8 × 0.6 cm, becoming recurved and revolute.
Vernacular name. Sabah—keruing kosugoi (preferred name).
Distribution. Borneo and the Philippines. In Borneo recorded from Kuala Penyu, Kudat,
Kinabatangan, Lahad Datu, Ranau, Tawau, and Tenom districts in Sabah (e.g., SAN 16402,
SAN 19551, SAN A 2140, SAN A 2492, and SAN A 4224) and SE and E Kalimantan (e.g., bb.
19783 and Kostermans 6775).
Ecology. Locally abundant, sometimes gregarious, in lower floodplains behind the

mangrove, and near river banks further inland; occasional on low hills on clayrich soils, at
altitudes to 200 m. Endangered by forest conversion.
39. Dipterocarpus verrucosus Foxw. ex Slooten

(Latin, verrucosus = warty; the lenticellate fruit calyx tube)
Bull. Jard. Bot. Buitenz. 3, 8 (1927) 293; Merrill op. cit. (1929) 201; Foxworthy op. cit. (1932) 71;
Masamune op. cit. 488; Symington op. cit. (1943) 31; Browne op. cit. 111; Ashton op. cit. (1964) 46,
op. cit. (1968) 19; Meijer & Wood op. cit. 272; Burgess op. cit. 101; Anderson op. cit. (1980) 113;
PROSEA op. cit. 186; Coode et al. (eds.) op. cit. 70; Newman et al. op. cit. 123. Lectotype
(designated here): Mitchell CF 0313, Peninsular Malaysia, Selangor, Kajang, Bangi FR (hololectotype
KEP; isolectotypes K, L).
Large emergent tree, to 60 m tall, to 1.5 m diameter; crown compact, hemispherical. Bark
orangebrown, thinly irregularly flaking. Twig, petiole, midrib and vein below, and flower
calyx densely caducously adpressed goldenbrown lustrous puberulent; leaf bud and
inflorescence densely, stipule sparsely, persistently so. Twigs c. 2 mm diameter apically,
terete or slightly compressed, stipule scars somewhat swollen. Leaf buds falcate or
occasionally conical, to 12 × 3 mm, with terminal tuft. Stipules narrowly oblong, to 8 × 0.8
cm, obtuse. Leaves coriaceous, glabrous, almost flat, drying distinctive pinkish red below
and slightly glazed; blade broadly ellipticovate, 6–12 × 3.5–6 cm, base cuneate, apex
subacute to broadly acuminate with acumen to 0.5 cm long; lateral veins 9–14 pairs, well
spaced, not prominent; intercostal venation wellspaced, somewhat sinuate; petiole 1.3–2 cm
long, to 2 mm diameter. Inflorescences to 9 cm long, axillary, unbranched or singly
branched. Flowers: buds to 2.5 × 1.2 cm; stamens c. 15, anthers filiformtapering. Fruits
subsessile; calyx tube ellipsoidglobose, tapering at base, 1.5–1. 8 × 0.6–0.8 cm, verrucose
lenticellate; 2 major calyx lobes to 8 × 1.5 cm, 3 minor ones deltoid, to 0.3 cm long, acute.
126
Vernacular name. Sabah and Sarawak—keruing merah (preferred name).
Distribution. Peninsular Thailand, Peninsular Malaysia and Borneo. In Sabah known from
Lahad Datu, Semporna, Sipitang and Tawau districts (e.g., SAN 15197, SAN 17175, SAN
18466, SAN 43697, and SAN 129564) and in Sarawak from Bau, Belaga, Bintulu, Kapit,
Lawas, Limbang, Lundu, Miri, Semarahan, Serian, and Sri Aman districts (e.g., S 14454, S
20291, S 22325, S 67568, and S 76735). Also occurring in Brunei (e.g., BRUN 3388 and
BRUN 5743) and W, C and E Kalimantan (e.g., Kostermans 13277 and Wilkie 94305).
Ecology. A species of shale and volcanic ridges and low hills, at altitudes below 800 m.
Curiously rare, even missing in most parts of Borneo but locally common along the high
ridges of the Limbang (Sarawak), southernmost Crocker Range (Sabah), and Amo district
(Brunei). Occurring in Mulu NP but vulnerable.
4. DRYOBALANOPS Gaertn.f.
(Greek, dryas = a nymph associated with oaks, balanops = acorn; the acornlike nut)
kapur (preferred name)
Fruct. 3 (1805) 49; Foxworthy, Malay. For. Rec. 10 (1932) 103; Slooten, Bull. Jard. Bot. Buitenz. 3,
12 (1932) 1; Symington, Malay. For. Rec. 16 (1943) 191; WyattSmith, Malay. For. 18 (1955) 145;
Browne, FTSB (1955) 111; Ashton, MDB (1964) 48, MDBS (1968) 20, FM 1, 9 (1982) 371; Meijer &
Wood, Sabah For. Rec. 5 (1964) 276; Burgess, TBS (1966) 117; Anderson, CLTS (1980) 114;
PROSEA, 1 (1993) 186; Coode et al. (eds.), CLBD (1996) 70; Newman et al., MDFDMHHW (1998)
125. Synonyms: Pterygium Correa, Ann. Mus. Nat. Hist. Paris 8 (1806) 397, p.p.; Baillonodendron
F.Heim, Bull. Mens. Soc. Linn. Paris 2 (1890) 867.
Very large emergent trees (except D. keithii) with immense cauliflowershaped crowns, the
minor branches numerous, the twigs bunched; buttresses narrowly rounded, often large
hardly branching plank. Bark yellowish brown, becoming irregularly flaky, sometimes
shaggy; inner bark pale brown, uniform, finely fibrous. Sapwood pale yellowish brown,
darkening gradually toward heartwood. Parts smelling more or less intensely aromatic of
camphor. Leaf buds small, ovoid, more or less compressed. Twigs slender, more or less
ribbed owing to the decurrent leaf traces. Stipules linear, small, early caducous. Leaves
coriaceous; blade lanceolate, ovate or oblong, apex prominently slenderacuminate; midrib
obscurely sunken above, slender but prominent below; lateral veins very many, very slender
and parallel, almost straight, fusing into a straight intramarginal vein at or close to the
margin, equal and without intermediates (D. keithii, D. oblongifolia excepted), unraised (D.
keithii excepted); intercostal venation obscure (D. keithii excepted); petiole slender,
channelled above, not geniculate. Inflorescences paniculate, terminal or axillary, lax, few
flowered; bracts and bracteoles minute, fugaceous. Flowers: buds spindleshaped, ellipsoid,
lanceolate or ovoid, acute or obtuse; sepals equal or subequal, imbricate, united at base into
an obconical tube tapering into pedicel; petals white or cream, broadly elliptic, subacute,
glabrous, hardly contorted in bud, connate at base and falling more or less in a rosette;
stamens c. 30 (c. 40 in D. oblongifolia), glabrous, subequal or unequal, epipetalous,
filaments broad and compressed, connate at base, broad at base, tapering and filiform
distally, anthers bright yellow, large, linear, latrorse, connectival appendage small, stout,
slightly or hardly exceeding the length of anthers; ovary small, ovoid, glabrous, without
stylopodium, style filiform, to 3x the length of ovary, stigma minute. Fruits: calyx with
127
incrassate basal cup partially enclosing but free from the nut, 5 lobes equal to subequal,
becoming valvate, obtuse, short or winglike, subrotate. Nuts large, with short apiculate
style remnant; pericarp woody, thin, splitting into 3 equal valves at germination; cotyledons
reniform, epigeal, on long slender hypocotyl; first two pairs of leaves opposite with short
intervening internode. Saplings of Massart’s model, that is with more or less horizontal, that
is plagiotropic, branches with distichous leaf arrangement clustered in false whorls owing to
intermittent growth of the vertical leader (compare, e.g., with Diospyros).
Distribution. Confined to W Malesia; seven species in Sumatra, Peninsular Malaysia and

Borneo; all occurring in Sabah and Sarawak.
Ecology. Frequently characteristically semigregarious, among the most abundant species of

the emergent canopy trees, dominating the skyline of the forests they inhabit (D. keithii
excepted). In mixed dipterocarp, kerangas and coastal mixed peat swamp forests on a
variety of soils, at altitudes below 800 m. The species flower more frequently than other
dipterocarps, and their saplings are generally abundant below parent trees. They are both
relatively shade tolerant and relatively fast growing in response to light; characteristics
which may help explain the frequent abundance of mature trees.
Uses. The timber, which differs in density and strength between species, is an important
moderately heavy and durable construction timber. The camphor obtained from the wood of
D. aromatica attracted early Arab traders, then being worth more than its weight in gold in
the Middle East and used for incense and perfume.
Key to Dryobalanops species

(based on all diagnostic characters)
1. Fruit calyx lobes shorter than the nut..…………6. D. oblongifolia (subsp. oblongifolia)
Fruit calyx lobes spatulate or oblongspatulate, longer than the nut……………………2
2. Leaf blade usually longer than 14 cm; lateral veins distinctly unequal, raised below,
obscure and sunken above; margin revolute throughout………………..…4. D. keithii
Leaf blade shorter than 12 cm; lateral veins subequal, unraised or hardly so below, not
sunken above; margin only partially or not revolute……………………………………3
3. Fruit calyx lobes 0.8–2 cm wide; calyx base fused into a cup to 2 cm diameter and at
least 0.5 cm deep……………………………………………………………………….4
Fruit calyx lobes to 1.3 cm wide; calyx base fused into a cup not more than 0.8 cm
diameter nor deeper than 0.5 cm……………………………………………………..…5
4. Leaf blade narrowly lanceolate, margin revolute at base…………….…5. D. lanceolata

Leaf blade broadly ovate, not revolute at base……………………….…1. D. aromatica
5. Leaf blade 5–8 × 1–3 cm, glabrous…………………………………….….2. D. beccarii

Leaf blade 5–11 × 2–4 cm, more or less tomentose below………………………….….6
128
6. Tomentum persistent, even, dark goldenbrown. Style 3–4x the length of ovary. Nuts
ellipsoidovoid, to 2 × 1.3 cm……………………………………………..…3. D. fusca
Tomentum more or less caducous, flocculent, rufous. Style 2x the length of ovary. Nuts
ovoid, to 1 × 0.6 cm……………………………………………………….…7. D. rappa
Key to Dryobalanops species

1. Leaf lateral veins unequal, distinctly raised below, sunken above; leaf margin narrowly
revolute throughout. In forest on clay soils in moist valleys……………….4. D. keithii
Leaf lateral veins subequal, more or less unraised below; leaf margin at most partially
revolute. In other types of habitats………………………………………………………2
2. Leaf blade more or less tomentose below (saplings sometimes excepted)…………...…3

Leaf blade glabrous………………………………………………………………..……4
3. Tomentum persistent, even, dark goldenbrown. In kerangas forest……..…3. D. fusca

Tomentum at least partiallly flocculent, rufous, caducous. In peat swamp forest...……...
……………………………………………………………………………….7. D. rappa
4. Leaf blade oblong, or narrowly lanceolate………………………………………………5

Leaf blade ovatelanceolate to oblonglanceolate………………………………………6
5. Leaf blade narrowly lanceolate, margin revolute at base. In mixed dipterocarp forest on
clay soils……………………………………………………………...…5. D. lanceolata
Leaf blade oblong, margin not revolute at base. In hill forest on moist clay soils and
seasonal swamp forest………………………….6. D. oblongifolia (subsp. oblongifolia)
6. Leaf blade oblonglanceolate to ovatelanceolate, thinly coriaceous; lateral veins

distinctly paler than blade with distinct fine intramarginal vein visible within margin.
Bark dark yellowish tawny. In mixed dipterocarp forest on shallow welldrained
soils……………………………………………………………………….2. D. beccarii
Leaf blade broadly ovate, thickly coriaceous; lateral veins not paler than blade,
indistinct; intramarginal vein obscure. Bark warm rustbrown. In mixed dipterocarp
forest on deep yellow sands……………………………………………1. D. aromatica
1. Dryobalanops aromatica Gaertn.f., nom. cons. Plates 3A–B.

(Latin, aromaticus = spicelike; the smell of the dammar)
Fruct. 3 (1805) 49; Merrill, EB (1921) 401; Foxworthy op. cit. 105; Slooten op. cit. (1932) 7;
Masamune, EPB (1942) 489; Symington, op. cit. (1943) 194; WyattSmith op. cit. 148; Browne op.
cit. 114; Ashton, Gard. Bull. Sing. 20 (1963) 241, op. cit. (1964) 49, op. cit. (1968) 375, op. cit. (1982)
375; Meijer & Wood op. cit. 278; Burgess op. cit. 118; Anderson op. cit. (1980) 114; Coode et al.
(eds.) op. cit. 70; Newman et al. op. cit. 130. Type: Miller s.n., Sumatra, Tapanuli (holotype BM).
Synonyms: Arbor camphorifera Rumph., Herb. Amb. Cap. 82 (1755) 67; Pterigium teres Correa,
Ann. Mus. Hist. Nat. Paris 10 (1807) 159; Dryobalanops camphora Colebr., As. Res. 12 (1816) 535;
Shorea camphorifera (Rumph.) Roxb., Fl. Ind. ed. Carey 2 (1832) 616; Dipterocarpus dryobalanops
Steudel, Nom. Bot. ed. 2, 1 (1840) 518; Dipterocarpus teres (Correa) Steudel op. cit. 518;
Dryobalanops junghuhnii Becc., Nelle For. Born. (1902) 554; Dryobalanops vriesii Becc. op. cit. 554;
129
Fig. 10. Dryobalanops beccarii. A, flowering leafy twig; B, detail of venation on lower leaf
surface; C, flower bud; D, open flower; E, adaxial view of stamens; F, mature fruit. (A–B
from S 23922, C from SAN 36331, D–E from S 7612, F from S 8591.)
130
Dryobalanops sumatrensis (J.F. Gmelin) Kosterm., Blumea 33 (1988) 346 (based on Laurus
sumatrensis J.F. Gmelin, Syst. Veg. (1791 & 1796) 650).
Immense tree, to 65 m tall, to 2 m diameter. Bark warm pale rustbrown, becoming evenly
flaky, later shaggy. All parts strongly aromatic. Young exposed fleshy parts yellowish
lepidote, caducous. Twigs c. 1 mm diameter apically, smooth, terete, slender. Leaves
thickly coriaceous, glabrous; blade broadly ovate (sometimes broader than long), 4–6 × 2–
4 cm, base cuneate to obtuse, margin at most only partially revolute, apex with slender
acumen to 1.5 cm long; lateral veins subequal, indistinct, not paler than the blade;
intramarginal veins obscure; petiole 0.5–1 cm long. Inflorescences to 7 cm long; axis
angular on drying. Flowers: buds spindleshaped, to 9 × 4 mm; sepals glabrous, lanceolate,
obtuse; petals more or less oblong, acute; stamens shorter than the style, filaments slender,
tapering, anthers linear, connectival appendage linear, somewhat exceeding anther apex;
style c. 2x the length of ovary, glabrous. Fruits glabrous; calyx base cupshaped, 0.6–0.8
cm deep, 0.8–1.5 cm diameter, becoming constricted at the rim; lobes spatulate, obtuse, 4–6
× 0.8–2 cm, tapering to 3–5 mm wide at base. Nuts ovoid with a short style remnant, to 3 ×
1.5 cm, with a constriction at the calyx cup rim.
Vernacular names. Sabah—kapur barus (preferred name). Sarawak—kapur peringgi

(preferred name), kapur ranggi (Iban), keladan (Iban).
Distribution. Sumatra (including Musala, Lingga and Singkep Is.), Peninsular Malaysia and
Borneo. In Borneo recorded in Sabah from Labuk Sugut, Papar and Sipitang districts (e.g.,
FMS 35242, FMS 35441, SAN 15148, SAN 50639, and SAN A 3298) and in Sarawak from
Bintulu, Kapit, Lawas, Limbang, and Miri districts (S 1790, S 4842, S 15104, S 22054, and
S 46493). Also occurring in Brunei (e.g., BRUN 508, BRUN 3001, FMS 34573, and Niga
NN 228).
Ecology. Locally the most abundant emergent trees in mixed dipterocarp forest on deep
humic yellow sandy soils with a propensity for ridges, at altitudes below 400 m. Vulnerable
though abundant in Lambir NP and recorded from Mulu NP.
Notes. The leaves of juveniles are indistinguishable from those of D. beccarii, except by the
stronger aroma when crushed, being less coriaceous, with distinct venation and visible
intramarginal vein.
2. Dryobalanops beccarii Dyer Fig. 10, Plate 3C.

(Odoardo Beccari, 1843–1920, Italian explorer and botanist)
J. Bot. 12 (April 1874) 100; Beccari op. cit. (1902) 572; Merrill op. cit. 401; Slooten op. cit. (1932)
36; Masamune op. cit. 489; WyattSmith op. cit. 149; Browne op. cit. 115; Ashton op. cit. (1963) 242,
op. cit. (1964) 51, op. cit. (1968) 22, op. cit. (1982) 375; Meijer & Wood op. cit. 280; Burgess, op. cit.
118; Anderson, op. cit. (1980) 114; PROSEA, op. cit. 191; Kessler & Sidiyasa, TBSAEK (1994) 95;
Coode et al. (eds.) op. cit. 70; Newman et al. op. cit. 131; Chua & Saw, Gard. Bull. Sing. (2003) 1.
Lectotype (Slooten, 1932): Beccari PB 2553, Borneo, Sarawak, Matang (hololectotype K;
isolectotype PC). Synonym: Dryobalanops oocarpa Slooten op. cit. (1932) 33, WyattSmith op. cit.
155.
131
Large emergent tree, to 65 m tall, to 2 m diameter. Bark becoming dark yellowish tawny,
prominently irregularly flaky but rarely shaggy. Parts only faintly aromatic. Exposed young
fleshy parts fugaceous puberulent, otherwise entirely glabrous. Twigs to 1 mm diameter
apically, slender, terete, smooth. Leaves thinly coriaceous, glabrous; blade ovatelanceolate
to oblonglanceolate, 5–8 × 1–3 cm, base cuneate, margin at most only partially revolute,
often wavy, apex with slender acumen to 1.7 cm long; lateral veins subequal, usually paler
than blade and distinctly visible albeit hardly or not raised; intramarginal vein distinct, c. 1
mm within margin; petiole 0.7–1 cm long, very slender. Inflorescences to 10 cm long,
irregularly doubly branched; axis wrinkled on drying. Flowers: buds spindleshaped, to 10
× 4 mm, acute; sepals equal, narrowly deltoid, subacute; petals broadly elliptic, obtuse;
stamens subequal, almost 2/3x the length of style, anthers narrowly oblong, acute,
connectival appendage short, erect, slightly exceeding anther; style 2–3x the length of
ovary. Fruits: calyx at base with obconical cup to 0.8 cm diameter, to 0.5 cm deep, tapering
into pedicel and without constricted rim; calyx lobes narrowly oblongspatulate, subacute,
to 6.5 × 0.8 cm, tapering to 2 mm wide at base. Nuts ovoid to globose, acute, to 1.4 cm
diameter and long, seated on the small calyx cup thereby pushing the lobes out to a rotate
angle.
Vernacular names. Sabah—kapur merah (preferred name). Sarawak—kapur bukit

(preferred name), keladan (Iban).
Distribution. Peninsular Malaysia (SE Johor) and Borneo (excepting the southern and
southwestern parts). In Borneo, localised in Sabah and recorded from Beaufort,
Kinabatangan, Kudat, Labuk Sugut, Pitas, Sandakan, and Sipitang districts (e.g., KEP
38720, KEP 80252, SAN 36331, SAN 121326, and SAN A 4341) but widespread in Sarawak
and recorded from Bau, Bintulu, Kapit, Kuching, Lundu, and Miri districts (e.g., S 367, S
BRUN 5282) and W, E and S Kalimantan (e.g., bb. 26578, bb. 29733, Kostermans 5995,
and Kostermans 6782).
Ecology. In mixed dipterocarp forest, locally among the most abundant emergent trees,
elsewhere scattered; on shallow leached soils over both sandstone and shale on high narrow
ridges, at altitudes to 700 m. Particularly abundant on coastal sandstone formations,
including near the mouths of Sabah east coast rivers but also on seasonally swamped
alluvium there; locally common on the high narrow inland ridges from G. Menuku, the
Klingkang Range, Bako and Mulu NPs in Sarawak to the Crocker Range in W Sabah.
Though its habitat is greatly reduced by logging, the species is probably not yet vulnerable.
3. Dryobalanops fusca Slooten

(Latin, fuscus = darkcoloured; the indumentum)
Bull. Jard. Bot. Buitenz. 3, 12 (1932) 39; Masamune op. cit. 489; WyattSmith op. cit. 150; Ashton op.
cit. (1968) 22, op. cit. (1982) 377; Anderson op. cit. (1980) 114; PROSEA op. cit. 191; Newman et al.
op. cit. 133. Lectotype (designated here): bb. 11468, Borneo, Kalimantan, Sambas, Paloh
(hololectotype BO).
Emergent tree, to 60 m tall, to 1.5 m diameter; bole frequently misshapen; crown distinctly
coppery from below. Bark becoming dark goldenbrown, irregularly coarsely flaking.
Exposed fleshy parts, leaf blade, flower and fruit excepted, densely shortly evenly
persistently pale rufousbrown tomentose; blade undersurface similarly goldenbrown
132
tomentose. Twigs c. 1–2 mm diameter apically, slender, muchbranched. Leaves

coriaceous; blade broadly lanceolate, 5–10 × 2–4 cm, base obtuse to broadly cuneate,
margin not or only partially revolute, apex subcaudate with acumen to 1.5 cm long; lateral
veins subequal, more or less unraised below; intercostal venation obscured by tomentum;
petiole 0.5–1 cm long. Inflorescences singly branched; axis terete, to 5 cm long. Flowers:
buds lanceolate, to 12 × 3 mm; sepals equal, narrowly loratedeltoid, obtuse; petals
lanceolate; stamens reaching below style apex, filaments c. 2/3x the length of anthers,
anthers narrowly oblong, tapering, connectival appendage erect, extending somewhat above
anther apex; style 3–4x the length of ovary. Fruits glabrous; pedicels to 6 mm long,
prominent; calyx cup shallow, to 0.3 cm deep, to 0.7 cm diameter; calyx lobes spatulate,
obtuse, to 6 × 1.3 cm, tapering to 4 mm wide at base. Nuts ellipsoidovoid, apiculate, to 2 ×
1.3 cm.
Vernacular name. Sarawak—kapur empedu (preferred name).
Distribution. Endemic in Borneo; recorded from Bintulu and Lundu districts in Sarawak
(e.g., S 2451, S 7999, S 9625, and S 18629) and from W Kalimantan (e.g., bb. 8895, bb.
11352 and bb. 14514).
Ecology. Locally once the commonest emergent tree in kerangas forest, usually on raised
beaches, at altitudes to 200 m. Rare and endangered.
4. Dryobalanops keithii Symington Fig. 11.

(H.G. Keith, 1899–?, one time Conservator of Forests in Sandakan)
Gard. Bull. S. S. 10 (1939) 379; Slooten, Bull. Jard. Bot. Buitenz. 3, 16 (1940) 449, Reinwardtia 5
(1961) 475; Masamune op. cit. 489; WyattSmith op. cit. 151; Meijer & Wood op. cit. 283; Burgess
op. cit. 118; Ashton op. cit. (1982) 373; PROSEA op. cit. 191; Newman et al. op. cit. 134. Type:
Keith FMS 44382, Borneo, Sabah, Tawau district, Tiaggau, Kalabakan (holotype KEP; isotype KEP).
Main canopy to low emergent tree, to 40 m tall, to 2 m diameter; bole frequently misshapen;
buttresses small and few. Bark smooth or eventually thinly flaky, pale ochreousbrown.
Outside of calyx and inflorescence fugaceous puberulent; parts otherwise glabrescent.
Twigs at first c. 2 mm diameter apically, verrucoselenticellate, becoming smooth. Leaves
coriaceous; blade narrowly oblong, lanceolate or oblanceolate, 14–33 × 5–10 cm, base
obtuse to cordate, margin narrowly revolute throughout, apex with slender acumen to 1 cm
long; lateral veins unequal, many and parallel but wellspaced, with prominent intermediate
veins, arched within margin forming a distinct intramarginal vein, slender but prominently
raised below, obscurely sunken above; intercostal venation visible but indistinct, reticulate;
petiole 0.7–1.2 cm long, stout. Inflorescences terminal or subterminalaxillary, singly or
doubly branched, to 14 cm long. Flowers: buds ellipsoid, to 10 × 4 mm, acute; sepals
narrowly deltoid, subacute; petals ovate, acute; stamens subequal, slightly shorter than style,
filaments c. half the length of anthers, anthers linear, tapering, connectival appendage
acicular, short but distinctly exceeding anther apex; style c. 2.5x the length of ovary. Fruits:
pedicels to 4 mm long; calyx tube obconical, shallow, to 0.6 cm deep, 1 cm diameter; calyx
lobes broadly spatulate, obtuse, to 4 × 2 cm, tapering to c. 8 mm wide above the cup. Nuts
depressed ovoid, apiculate, to 1.6 × 1.5 cm.
Vernacular name. Sabah—kapur gumpait (preferred name).
133
Distribution. Endemic in Borneo; recorded from Kinabatangan, Sandakan, Tawau, and

Tenom districts in Sabah (e.g., FMS 44430, SAN 18732, SAN 97485, and SAN A 2587) and
from the extreme NE of Kalimantan (e.g., bb. 17944 and bb. 18159).
Ecology. Once locally abundant trees on welldrained but moist clay soils on low hills, at
altitudes to 250 m, never far from streams and often on their banks. Vulnerable owing to
forest conversion.
5. Dryobalanops lanceolata Burck

(Latin, lanceolatus = shaped like the head of a spear; the leaf blade)
Ann. Jard. Bot. Buitenz. 6 (1887) 244; Merrill op. cit. 401; Slooten op. cit. (1932) 28; Keith op. cit.
37; Masamune op. cit. 489; WyattSmith op. cit. 152; Browne op. cit. 116; Ashton op. cit. (1964) 52,
op. cit. (1968) 22, op. cit. (1982) 374; Meijer & Wood op. cit. 285; Burgess op. cit. 118; Anderson op.
cit. (1980) 70; PROSEA op. cit. 191; Coode et al. (eds.) op. cit. 70; Newman et al. op. cit. 135. Type:
Cult. in Hort. Bog. Sub VIIID 58 (from Beccari via Martin), Borneo, Sarawak, Bintulu district, Ulu
Kemena (holotype BO). Synonym: Dryobalanops kayanensis Becc., op. cit. 551.
Tall emergent tree, to 80 m tall, to 2.5 m diameter; bole tall, columnar; crown immense, like
a rising thunder cloud; buttresses concave. Bark dark leaden green, thinly irregularly
flaking. Leaf buds and stipules fugaceous puberulent; parts otherwise glabrous. Twigs c.
1.5 mm diameter apically, slender, smooth, pale lenticellate. Leaves thinly coriaceous,
glabrous, drying reddish brown; blade narrowly lanceolate, 7–10 × 2–3.5 cm, base cuneate,
margin revolute at base only, apex with slender acumen to 1 cm long; lateral veins
subequal, unraised below; petiole c. 1 cm long, slender. Inflorescences terminal or axillary,
simple or singly branched; axis ribbed, to 6 cm long. Flowers: buds spindleshaped, to 12 ×
5 mm, obtuse; sepals subequal, lanceolate, obtuse; petals lanceolate, acute, glabrescent;
stamens subequal, shorter than the style, anthers linear, connectival appendage hardly
exceeding anther apex; style c. 3x the length of ovary, glabrous. Fruits: calyx impressed at
pedicel, the base united into a shallow cup with waisted rim, to 2 cm diameter, to 0.5 cm
deep, with a hollow tubercle at the base of each lobe; calyx lobes spatulate, obtuse, to 9 × 2
cm, tapering to 3–5 mm wide at base. Nuts ovoid to globose, subacute, to 2 cm long and
broad.
Vernacular names. Sabah and Sarawak—kapur paji (preferred name). Sarawak—paji

(Iban), sesuan (Murut).
Distribution. Endemic in Borneo. Common and widespread in Sabah, particularly in the

eastern parts and recorded from Kudat, Lahad Datu, Papar, Sandakan, Sipitang, Tawau, and
Tenom districts (e.g., SAN 15189, SAN 17844, SAN 18004, SAN 27281, and SAN 83425)
and in Sarawak from Belaga, Bintulu, Kapit, Lawas, Limbang, Marudi, Miri, Sibu, Sri
Aman, and Tatau districts (e.g., S 1286, S 1428, S 15592, S 22410, and S 32386). Also
occurring in Brunei (e.g., BRUN 311, FMS 34562 and FMS 39604), and E and SE
Kalimantan (e.g., bb. 22866, bb. 29394, Kostermans 5406, and Kostermans 6209).
Ecology. In mixed dipterocarp forest on clayrich soils, mostly on lower slopes but
occasionally to 700 m altitude; most abundant, becoming the dominant emergent on deep
moist but wellaerated soils on calcareous shale and basic igneous rocks, therefore
threatened outside strictly managed parks. The saplings are shade tolerant, surviving many
years and expanding extensive horizontal to slightly pendent plagiotropic branches with
only slight extension of the leader shoots; but once a gap is formed young paji grow in
134
Fig. 11. Dryobalanops keithii. A, flowering leafy twig; B, detail of indumentum on lower
leaf surface; C, flower bud; D, postanthesis flower; E, longitudinal section of open flower;
F, adaxial view of stamens; G, abaxial view of stamens; H, ripe fruit with one calyx lobe
removed. (A–D from SAN 18732, E–G from FMS 55279, H from SAN 123771.)
135
height at comparable rates to the fastest pioneers. Common in Sepilok FR and Danum
Valley Conservation Area, Sabah and locally so in Lambir and Mulu NPs, Sarawak;
vulnerable elsewhere owing to land conversion.
6. Dryobalanops oblongifolia Dyer Fig. 12, Plate 3D.

(Latin, oblongus = rather long, folium = leaf; the leaf shape)
J. Bot. 12 (1874) 100; Foxworthy op. cit. 118; Slooten op. cit. (1932) 22; Symington op. cit. (1943)
196; WyattSmith op. cit. 153, p.p.; Browne op. cit. 116; Ashton op. cit. (1968) 23, Gard. Bull. Sing.
31 (1978) 25, op. cit. (1982) 373. Type: Beccari PB 2533, Borneo, Sarawak, Matang (holotype K).
Distribution. Sumatra, Peninsular Malaysia and Borneo.
Notes. Two subspecies are recognised, viz. subsp. oblongifolia and subsp. occidentalis
P.S.Ashton, with the former occurring in Borneo and the latter in Sumatra and Peninsular
Malaysia.
subsp. oblongifolia
Beccari, Nelle For. Born. (1902) 550; Merrill op. cit. 401; Masamune op. cit. 489; Browne op. cit.
116; Ashton op. cit. (1968) 23, op. cit. (1978) 26, op. cit. (1982) 373; Anderson op. cit. (1980) 114;
PROSEA op. cit. 192; Newman et al. op. cit. 136. Synonyms: Baillonodendron malayanum F.Heim,
op. cit. (1890) 867; Dryobalanops abnormis Slooten op. cit. (1940) 449.
Emergent tree, to 60 m tall, to 2 m diameter. Bark dark yellowish brown, flaky,

occasionally becoming shaggy. Inflorescence densely evenly rufouspubescent; young
shoots fugaceous pale fulvouspubescent; parts otherwise glabrescent. Twigs c. 2 mm
diameter apically, smooth, shiny. Leaves thinly coriaceous, frequently undulate, glabrous,
drying rustbrown, paler above than below; blade oblong, 6–20 × 4.5–5(–6.5) cm, base
broadly cuneate to obtuse, margin not revolute at base, apex cuspidate with acumen to 1.5
cm long; lateral veins subequal, distinct but only slightly elevated below, with visible
branching intermediate veins; petiole 0.5–1.2 cm long. Inflorescences terminal or axillary,
singly branched, to 14 cm long. Flowers: buds ovoid, to 8 × 5 mm, glabrous; sepals ovate,
acute; petals oblong; stamens c. 40, unequal, reaching to below the style apex, filaments c.
half the length of anthers, broadly lorate, anthers linear, connectival appendage more or less
erect, exceeding anther apex; style 2–3x as long as ovary, glabrous. Fruits subsessile; calyx
lobes shorter than the nut, incrassate, deltoid, acute, often reflexed, to 0.5 × 0.7 cm,
bordering a massive incrassate basal cup to 1.5 cm deep and diameter. Nuts ellipsoid to
obovoid, to 3.5 × 2.7 cm, more or less mucronate, prominently lenticellate.
Vernacular names. Sarawak—kapur kelansau (preferred name), kelansau (Iban).
Distribution. Endemic in Borneo. Recorded in Sarawak from Kapit, Kuching, Lundu,

Mukah, and Serian districts (e.g., S 4432, S 18628, S 19652, and S 27008). Also occurring in
W, C and E Kalimantan (e.g., bb. 16952, bb. 30212, bb. 31221, and Wilkie 94309).
Ecology. Locally common in mixed dipterocarp forest, on sandy clay soils, on undulating
land, in valleys and on slopes, at altitudes to 600 m; frequently as small groups of mature
individuals but not seen to dominate the canopy over large areas. Vulnerable, though
occurring in Kubah NP.
136
Fig. 12. Dryobalanops oblongifolia subsp. oblongifolia. A, flowering leafy twig; B, detail of
venation on lower leaf surface; C, detail of indumentum on lower leaf surface; D, detail of
indumentum on inflorescence and flowers; E, postanthesis flower; F, longitudinal section
of postanthesis flower; G, mature fruit; H, longitudinal section of mature fruit. (A–F from S
23993, G–H from S 41496.)
137
7. Dryobalanops rappa Becc.

(from an Iban word—kerapa = a shallow swamp; the habitat)
Nelle For. Born. (1902) 572; Merrill op. cit. 401; Slooten op. cit. (1932) 41; Keith op. cit. 37;
Masamune op. cit. 489; Browne op. cit. 116; WyattSmith op. cit. 155; Anderson, Gard. Bull. Sing. 20
(1963) 157, op. cit. (1980) 114; Ashton op. cit. (1963) 242, op. cit. (1964) 53, op. cit. (1968) 23, op.
cit. (1978) 25, op. cit. (1982) 377; Meijer & Wood op. cit. 289; Burgess op. cit. 118; PROSEA op. cit.
192; Coode et al. (eds.) op. cit. 70; Newman et al. op. cit. 137. Type: Beccari s.n., Borneo, Sarawak,
Kuching (holotype FI).
Emergent tree, to 55 m tall, to 1.5 m diameter. Bark becoming rufousbrown, shaggily

flaky. Exposed fleshy parts, leaf blade above, flower and fruit, and sapling excepted,
densely powdery more or less persistently rufoustomentose. Twigs c. 1.5 mm diameter
apically, smooth but for minute warty lenticels. Leaves coriaceous, drying mauvebrown
above, rufous to greyish tomentose below; blade ovatelanceolate, 6–11 × 2.5–4 cm, base
obtuse, margin only partially revolute, apex with slender acumen to 1 cm long; lateral veins
subequal, unraised below; petiole 0.6–1 cm long, stout. Inflorescences terminal or axillary,
doubly branched, to 8 cm long. Flowers: buds ellipsoid, to 10 × 5 mm, acute; sepals
narrowly deltoid, obtuse; petals cream, lanceolate; stamens reaching to below style apex,
filaments c. 2/3x the length of anthers; anthers narrowly oblong, connectival appendage
erect, extending somewhat above anther apex; style c. 2x as long as ovary, glabrous. Fruits:
calyx with shallow basal cup to 0.8 cm diameter, to 0.3 cm deep; calyx lobes thinly
coriaceous, linearspatulate, to 5 × 0.6 cm, subacute, tapering to 3.5 mm wide at base. Nuts
ovoid, to 1 × 0.6 cm, mucronate.
Vernacular name. Sabah and Sarawak—kapur paya (preferred name).
Distribution. Endemic in Borneo. Recorded in Sabah from Beaufort, Papar and Sipitang
districts (e.g., FMS 41106, FMS 49021, SAN 27991, SAN 78005, and SAN A 1704), and in
Sarawak from Bintulu, Kuching, Lawas, Limbang, Lundu, Marudi, Miri, Sematan and Sibu
districts (e.g., Haviland 2226, S 1270, S 1504, S 4850, and S 11242). Also occurring in
Brunei (e.g., BRUN 5105, BRUN 5106, FMS 28696, FMS 30640, and Wong WKM 917).
Ecology. In coastal mixed peat swamp forest, particularly over sands behind the sea beach
but also on clay, and in lower montane kerangas, at altitudes to 900 m. Vulnerable, though
occurring in Mulu NP
5. HOPEA Roxb., nom. cons.

(John Hope, 1725–1786, the first Regius Keeper of the Royal Botanic Garden, Edinburgh)
luis (Iban), selangan (Malay)
Pl. Corom. 3 (1811) 7; King, J. As. Soc. Beng. 62, 2 (1893) 125; Ridley, FMP 1 (1922) 234;
Foxworthy, Malay. For. Rec. 10 (1932) 113; Symington, Malay. For. Rec. 16 (1943) 108; Browne,
FTSB (1955) 116; Ashton, Gard. Bull. Sing. 20 (1963) 254, MDB (1964) 89, MDBS (1968) 37, Gard.
Bull. Sing. 31 (1978) 28, FM 1, 9 (1982) 391; Meijer & Wood, Sabah For. Rec. 5 (1964) 203; Backer
& Bakhuizen f., FJ 1 (1964) 330; Burgess, TBS (1965) 128; Anderson, CLTS (1980) 114; PROSEA,
5, 1 (1993) 238 & 255; Kessler & Sidiyasa, TBSAEK (1994) 96; Coode et al. (eds.), CLBD (1996)
71; Newman et al., MDFBMHHW (1998) 139. Synonyms: Neisandra Rafin., Sylv. Tell. (1838) 163;
Petalandra Hassk., Cat. Hort. Bog. (1858) 104; Balanocarpus Bedd., For. Man. Bot. (1873) 236 bis;
138
Hancea Pierre, For. Fl. Coch. 4 (1891) sub t. 244; Pierrea auct. non Hance: Heim, Bull. Mens. Soc.
Linn. Paris 2 (1891) 958; Dioticarpus Dunn, Bull. Misc. Inform. Kew (1920) 337.
Main canopy and subcanopy, occasionally low emergent trees, with narrow buttresses, often
flying buttresses, sometimes stilt roots. Bark variously smooth, chocolatebrown mottled, or
flaky or fissured; inner bark uniform pale yellowish brown, finely fibrous. Sapwood
variously moderately soft to very hard, creamwhite to straw yellow; heartwood pale to dark
chocolatebrown. Twigs generally slender, terete, smooth, brown, with short inconspicuous
stipule scars. Leaf buds generally minute. Stipules linear, early caducous except in
juveniles. Leaves small or mediumsized, occasionally large; venation pinnate, without
intramarginal veins, either ‘hopeatype’, i.e. with all lateral veins more or less parallel and
reaching the leaf margins, without intermediates, and with scalariform intercostal venation
(in sect. Hopea, e.g., H. nervosa and H. rudiformis), or ‘dryobalanoid’, i.e. with many fine
but arched and unequal lateral veins and obscurely reticulate intercostal venation (in sect.
Dryobalanoides subsect. Dryobalanoides), or ‘subdryobalanoid’, i.e. intermediate between
these (in sect. Dryobalanoides subsect. Sphaerocarpae); petiole rarely geniculate.
Inflorescences paniculate, slender, terminal or axillary or ramiflorous when frequently
fascicled. Flowers secund; buds small, ovoid or globose; 2 outer sepals ovate, acute or
obtuse, thickened, 3 inner ones suborbicular, thin at margins, frequently mucronate; petals
oblong, strongly contorted, connate at base forming a basal cup with rotate twisted lobes,
falling in a rosette; stamens 10, 15 or to 38 (H. plagata), attached to the base of the petals
and falling with them, filaments broad at base, tapering and filiform distally, anthers broadly
ellipsoid to globose, more or less tapering apically, connectival appendage usually at least
twice the length of anther, very slender, glabrous or minutely tuberculate; ovary glabrous or
pubescent, ovoid and without stylopodium or with stylopodium and variously pyriform,
cylindrical or hourglassshaped, style columnar, long (if without stylopodium) or more or
less short (if with stylopodium), stigma obscure. Fruits relatively small, usually glabrous; 2
outer calyx lobes generally spatulate and enlarged, the 3 inner ones remaining as broad as
long, embracing the base of the nut and generally shorter than it at maturity; frequently all
5 sepals short; all sepals expanded and thickened around the base of the nut and appressed
to it. Nuts ovoid, generally glabrous, apiculate with the distinct floral stylopodium, when
present, persisting; pericarp splitting at germination irregularly or occasionally into 3
valves.
Distribution. About 104 species, distributed from Sri Lanka and S India to S China, and
southeastward throughout Malesia (the Lesser Sunda Is. excepted), to New Guinea; 43
species occur in Sabah and Sarawak.
Ecology. Main and subcanopy trees of lowland forests at altitudes below 800 m, rarely to
1600 m. In mixed and upper dipterocarp, kerangas and mixed peat swamp forests, and a few
species on limestone karst; several are confined to river banks. Most, especially those with
short fruit sepals, occur in tightly clustered groups with their juveniles, sometimes extending
to form semigregarious populations.
Notes. Hopea is not difficult for the field worker to recognise on account of the characters
italicised in the description given above. From the related genus Shorea (see there) most
Hopea differ in being of the main canopy or subcanopy, with stilt roots and flying buttresses
(virtually absent in Shorea), more slender twigs, smaller flowers and of course the two
winged usually smaller and generally glabrous fruit. However, the great variability in ovary,
leaf, bark and mature habit make this one of the most diverse dipterocarp genera. Two
sections, each with two subsections are recognised according to the characters indicated in
139
the systematic key; but these entities have greater floral diversity than do the sections of
Shorea, and several species with intermediate characters exist; the boundaries are therefore
to an extent arbitrary. Curiously, molecular evidence (Kamiya et al., Tropics 7 (1998) 195–
207; Dayanandan et al., Amer. J. Bot. 86 (1999) 1182–1190) indicates that the genus is
indeed monophyletic, but nested between the sections of the vast genus Shorea along with
Parashorea and Neobalanocarpus (a monotypic genus of Peninsular Malaysia).
Hopea species generally have smooth, often hoopmarked, or flaky bark; but some
dryobalanoid species, viz. H. beccariana, H. cernua, H. coriacea, H. mengerawan, and H.
treubii may have deeply vsection fissured chocolate brown bark in large trees. Curiously,
this seems not to be always the case, as populations occur in H. beccariana and H. coriacea,
in which large individuals are present, in which the bark remains at most irregularly
cracked.
Key to Hopea species

1. Leaf venation hopeatype or subdryobalanoid………………………………………..…2

Leaf venation dryobalanoid (sect. Dryobalanoides subsect. Dryobalanoides)……..…4
2. Ovary and stylopodium not or only slightly constricted in between; leaf blade oblong
lanceolate, ovatelanceolate, lanceolate, ovate to narrowly elliptic or suborbicular……3
Ovary and stylopodium hourglassshaped, distinctly constricted at base of stylopodium;
leaf blade generally narrowly oblong (sect. Hopea subsect. Pierrea)……………..…37
3. Flowers remote on inflorescence; bracts subpersistent (sect. Dryobalanoides subsect.

Sphaerocarpae)…………………………………………………………………….…20
Flowers dense on inflorescence; bracts fugaceous (sect. Hopea subsect. Hopea)….…28
4. Ovary without a distinct stylopodium, ovoid with long filiform or columnar style…….5
Ovary with distinct stylopodium, style short.……………………………………….…15
5. Shorter fruit calyx lobes conceiling ripe nut.………………………………………..…6

Shorter fruit calyx lobes not or hardly conceiling ripe nut (ripe nut apex exposed)….....9
6. Leaf margin revolute………………………………………………………………….…7

Leaf margin not revolute…………………………………………………………….…8
7. Leaf blade to 5 cm wide, elliptic to ovate, thinly coriaceous; stamens 15–18……………

………………………………………………………………………………8. H. cernua
Leaf blade 6–10 cm wide, broadly ovate to suborbicular, thickly coriaceous; stamens at
most 15……………………………………………………………………9. H. coriacea
140
8. Fruit pedicel to 7 mm long; calyx lobe base prominently tuberculate…….……………...

……………………………………………………………………19. H. longirostrata
Fruit pedicel to 2 mm long; calyx lobe base incrassate but not tuberculate……………...
……………………………………………………………………………15. H. fluvialis
9. Leaf lateral veins at least 14 pairs, indistinct below.………………………………..…10

Leaf lateral veins less than 14 pairs, distinct below……………………………………11
10. Leaf base cuneate…………………………………………………21. H. mengerawan

Leaf base obtuse……………………………………………………….23. H. micrantha
11. Fruit calyx lobes short, subequal; leaf lateral veins at most 6 pairs………………………
…………………………………………………………………….17. H. kerangasensis
Fruit calyx lobes unequal, 2 larger ones spatulate; leaf lateral veins at least 8 pairs…..12
12. Nut to 1.5 cm long, cylindricaloblong………………………………...…42. H. vesquei

Nut shorter than 1.1 cm, ovoid……………………………………………………..…13
13. Fruit calyx lobes larger, 2 longer ones tapering to 5 mm above the saccate base, 3
shorter ones shorter than the nut; leaf midrib acute on both surfaces, drying black……...
…………………………………………………………………11. H. dryobalanoides
Fruit calyx lobes smaller, 2 longer ones tapering to 1.5–3 mm above the saccate base, 3
shorter ones conceiling the nut; leaf midrib terete, not drying black……………….…14
14. Leaf blade lanceolate; petiole at most 1 cm long; twig apex persistently puberulent……
………………………………………………………………………..14. H. ferruginea
Leaf blade ovate; petiole at least 1.2 cm long; twig glabrescent…………18. H. latifolia
15. Leaf midrib obscure, sunken above……………………………………16. H. griffithii

Leaf midrib evident, more or less raised above………………………………………..16
16. Stamens c. 10; leaf blade broadly ellipticobovate…………………….….40. H. treubii

Stamens c. 15; leaf blade ovate to lanceolate……………………………………….…17
17. Ovary and stylopodium pyriform or hourglassshaped…………………………….…18

Ovary and stylopodium cylindrical………………………………………………..…19
18. Leaf blade broadly ovate; petiole 1.2–1.7 cm long………………….…4. H. beccariana

Leaf blade narrowly ovatelanceolate; petiole at most 1 cm long……….….12. H. dyeri
19. Margin of leaf blade base prominently revolute…………………….…2. H. altocollina

Margin of leaf blade base not revolute..…………………………..…30. H. pedicellata
20. Leaf venation hopeatype………………………………………………………….…21

Leaf venation subdryobalanoid………………………………………………………23
21. Fruit calyx lobes short, subequal……………………………………..…1. H. aequalis

Fruit calyx lobes unequal, 2 larger ones spatulate………………………………....…22
141
22. Leaf lateral veins at most 13 pairs; blade dull below……………...…34. H. rudiformis
Leaf lateral veins 13–15 pairs; blade not dull, drying dark chocolatebrown below……..
……………………………………………………………………………25. H. nervosa
23. Fruit calyx lobes unequal, 2 larger ones spatulate……………………....24. H. montana

Fruit calyx short, subequal……………………………………………………………24
24. Leaf blade oblonglanceolate or ovatelanceolate, at least 7 cm long…………………25

Leaf blade ovate, at most 6 cm long………………………………………………..…27
25. Petiole at least 0.7 cm long; leaf blade coriaceous, drying honeybrown………………...
………………………………………………………………………..22. H. mesuoides
Petiole at most 0.6 cm long; leaf blade chartaceous, drying grey or tawny brown……26
26. Leaf blade undulate or more or less flat on drying, with obtuse base…………………….
……………………………………………………………………..38. H. sphaerocarpa
Leaf blade wrinkled on drying, base more or less cuneate…………..…35. H. rugifolia
27. Stamens c. 10; leaf blade at most 2.5 cm long…………………….…41. H. vaccinifolia

Stamens c. 15; leaf blade at least 2.5 cm long………………………...…5. H. bracteata
28. Stamens c. 10……………………………………………………………………….…29

Stamens 15 or more..……………………………………………………………….…30
29. Leaf lateral veins at most 8 pairs; domatia obscure, glabrous, or absent…………………
……………………………………………………………………..10. H. depressinerva
Leaf lateral veins at least 10 pairs; domatia evident, pubescent………..…36. H. sangal
30. Style as long as ovary……………………………………………………7. H. centipeda

Style very short (shorter than ovary)………………………………………………..…31
31. Stamens at least 32………………………………………………………32. H. plagata

Stamens at most 15………………………………………………………………….…32
32. Leaf base obtuse; domatia porelike, prominently swollen………………26. H. nutans

Leaf base cuneate; domatia not prominently swollen…………………………………33
33. Leaf lateral veins at most 5 pairs……………………………………31. H. pentanervia

Leaf lateral veins at least 6 pairs…………………………………………………….…34
34. Leaf base distinctly unequal………………………………………..…3. H. andersonii

Leaf base equal to subequal……………………………………………………………35
35. Inflorescence glabrous……………………………………………..…20. H. megacarpa

Inflorescence densely persistently buffpubescent……………………………………36
36. Ovary and stylopodium ovoid………………………………………..…28. H. ovoidea

Ovary and stylopodium cylindrical, truncate…………………….…37. H. semicuneata
142
37. Leaf blade broadly ovate, base equal; inflorescence solitary…...….43. H. wyattsmithii
Leaf blade narrowly oblong, elliptic to lanceolate, base unequal, obtuse to subcordate at
least on one side; inflorescence usually more than 1 in each leaf axil…………….....38
38. Fruit calyx lobes short, subequal; leaf blade pale, often silvery below..…………………
……………………………………………………………………..29. H. pachycarpa
Fruit calyx lobes unequal, 2 larger ones spatulate; leaf blade not silvery below..……..39
39. Leaf blade prominently bullate between the intercostal venation…...…6. H. bullatifolia
Leaf blade not bullate or only slightly so..……………………………………………40
40. Base of fruit calyx lobes auriculate………………………………...….33. H. pterygota

Base of fruit calyx lobes not auriculate……………………………………………..…41
41. Leaf lateral veins more prominent above than below; blade thickly coriaceous, drying
pale pinkish brown below..……………………………………..…27. H. obscurinerva
Leaf lateral veins more prominent below than above; blade thinly coriaceous, drying
chocolate or greenishbrown below……………………………………………….…42
42. Leaf blade (mature trees) smaller, 10–27 × 3–5.5 cm, drying chocolatebrown; lateral
veins 12–21 pairs……………………………………………….….39. H. tenuinervula
Leaf blade larger, 27–46 × 8–15 cm, drying greenish brown; lateral veins 16–30 pairs…
………………………………………………………………….13. H. enicosanthoides
Key to Hopea species

1. Leaf venation dryobalanoid or subdryobalanoid…………………………………….…2

Leaf venation hopeatype.…………………………………………………………..…29
2. Bark fissured or densely flaky; stilt roots few or none……………………………….…3

Bark smooth or cracked to patchily flaky only; stilt roots present at least on young
trees………………………………………………………………………………….….9
3. Leaf blade at least 6 cm wide, thickly coriaceous…………….…9. H. coriacea (in part)

Leaf blade at most 5.5 cm wide, not thickly coriaceous……………………………..…4
4. Bark surface dark or chocolate brown, fissured……………………………………...…5

Bark surface tawny brown, powdery flaky or hoopmarked or patchily cracked or
vsection fissured……………………………………………………………………..…7
5. Leaf lateral veins raised below, drying blackish………………..…8. H. cernua (in part)
Leaf lateral veins unraised below, not drying blackish……………………………….…6
6. Petiole at most 1.1 cm long, stout……………………………..….21. H. mengerawan

Petiole at least 1.2 cm long, slender……………………………………4. H. beccariana
143
7. Leaf blade broadly ellipticobovate, apex shortly acuminate…………..…40. H. treubii

Leaf blade narrowly ovate to lanceolate, apex not acuminate………………………......8
8. Leaf lateral veins prominent below; leaf margin revolute at base.……..2. H. altocollina
Leaf lateral veins obscure below; leaf margin not revolute at base………………………
……………………………………………………………….30. H. pedicellata (in part)
9. Midrib obscure, sunken above…………………………………………………………10

Midrib evident, more or less raised above………………………………………….…17
10. Leaf venation dryobalanoid; intercostal venation obscure below…………………..…11

Leaf venation subdryobalanoid; intercostal venation evident below (except H. montana)
………………………………………………………………………………………….12
11. Leaf lateral veins at least 9 pairs, without domatia…………………..…16. H. griffithii

Leaf lateral veins at most 6 pairs, with pubescent axillary domatia……………………...
……………………………………………………………………17. H. kerangasensis
12. Leaf larger, 8–14 cm long…………………………………………..…22. H. mesuoides

Leaf blade smaller, 1–10 cm long…………………………………………………..…13
13. Intercostal venation obscure below……………………………………24. H. montana

Intercostal venation distinct below……………………………………………………14
14. Leaf blade elliptic to narrowly or broadly ovate………………………………….……15

Leaf blade broadly lanceolate to oblonglanceolate………………………………...…16
15. Leaf blade 2.5–6 × 0.7–2 cm; petiole 0.2–0.4 cm long. In forest on clay soils…………..
…………………………………………………………………………..5. H. bracteata
Leaf blade 1–2.5 × 0.4–1.2 cm; petiole to 0.2 cm long. In forest on white sand
podsols…………………………………………………………….…41. H. vaccinifolia
16. Leaf blade obtuse at base, not wrinkled on drying……………...…38. H. sphaerocarpa

Leaf blade cuneate at base, wrinkled on drying……………………..….35. H. rugifolia
17. Leaf venation subdryobalanoid; inner bark plumred….…43. H. wyattsmithii (in part)
Leaf venation dryobalanoid; inner bark pale brown or pink…………………………18
18. Leaf blade suborbicular; main lateral veins at most 4 pairs……..9. H. coriacea (in part)
Leaf blade ovate, elliptic, lanceolate or oblonglanceolate; main lateral veins at least 5
pairs……………………………………………………………………………………19
19. Midrib sharp below, tending to dry black………………………………………….…20

Midrib terete below, not drying black…………………………………………..……22
20. Young twigs glabrous; inner bark pale brown………………….…19. H. longirostrata

Young twigs pubescent; inner bark pinkish brown………………………………….…21
144
21. Leaf blade coriaceous; main lateral veins c. 15 pairs, not drying black………………….
……………………………………………………………….30. H. pedicellata (in part)
Leaf blade chartaceous; main lateral veins at most 12 pairs, drying black……………….
………………………………………………………….11. H. dryobalanoides (in part)
22. Leaf base equal to subequal…………………………………………………….…..…23

Leaf base unequal, blade twisted slightly to one side……………………………….…25
23. Leaf base obtuse…………………………………………………..….23. H. micrantha

Leaf base cuneate………………………………………………………………………24
24. Leaf lateral veins slightly raised below, drying paler than the blade……………………..
………………………………………………………….11. H. dryobalanoides (in part)
Leaf lateral veins indistinct, hardly raised below, drying not paler than the blade………
………………………………………………………………………….18. H. latifolia
25. Basal pair of lateral veins continuing up leaf margin for one third of its length…………
…………………………………………………………………………..15. H. fluvialis
Basal pair of lateral veins short…………………………………………………….…26
26. Leaf lateral veins distinctly raised below……………………..…8. H. cernua (in part)
Leaf lateral veins somewhat or not raised below…………………………………….27
27. Leaf blade broadly ovate, thickly coriaceous, base subobtuse; bark surface with short
shallow surface fissures………………………………………………..…42. H. vesquei
Leaf blade narrowly ovatelanceolate, thinly coriaceous to coriaceous, base cuneate;
bark smooth becoming thinly flaky.………………………………………………..…28
28. Leaf lateral veins at most 12 pairs……………………………………….…12. H. dyeri

Leaf lateral veins c. 14 pairs………………………………………..…14. H. ferruginea
29. Bark scaly, vertically cracked or flaky…………………………………………….…30

Bark smooth……………………………………………………………………………39
30. Leaf lateral veins at most 5 pairs………………………………..….31. H. pentanervia

Leaf lateral veins at least 6 pairs…………………………………………………….…31
31. Leaf base markedly unequal………………………………………………………..…32

Leaf base subequal to equal………………………………………………………..…35
32. Leaf blade more than 3x as long as wide; lateral veins 12–21 pairs….…………...……..
……………………………………………………………………..39. H. tenuinervula
Leaf blade less than 2½x as long as wide; lateral veins 7–12 pairs………………..…33
33. Leaf lateral veins with pubescent axillary domatia……………………..3. H. andersonii

Leaf lateral veins without or with a few glabrous domatia…………………………...34
34. Leaf lateral veins 8–11(–12) pairs, domatia, if present, porelike; leaf bud glabrous……
………………………………………………………………………….32. H. plagata
145
Leaf lateral veins 7–8(–10) pairs, domatia canaliculate; leaf bud puberulent……………
………………………………………………………………….28. H. ovoidea (in part)
35. Leaf lateral veins 10–12 pairs; bark surface dark brown with conspicuous white
dammar coxcombs…..............................................................................…36. H. sangal
Leaf lateral veins less than 10 pairs; bark surface brown, dammar exudations not
conspicuous…………………………………………………………………………….36
36. Leaf blade broadly ovate, thickly coriaceous, greyish lepidote below…....26. H. nutans
Leaf blade lanceolate, narrowly elliptic or narrowly ovateelliptic, chartaceous to thinly
coriaceous, undulate, not pale lepidote below.………………………………………37
37. Leaf lateral veins sharply raised below, domatia obscure or absent……………………...
………………………………………………………………….10. H. depressinerva
Leaf lateral veins not sharply raised below, with prominent axillary domatia………...38
38. Domatia canaliculate………………………………………..…28. H. ovoidea (in part)

Domatia porelike…………………………………………………37. H. semicuneata
39. Leaf base equal…………………………………………………………………….….40

Leaf base markedly unequal or subequal……………………………………………....44
40. Leaf lateral veins at most 8 pairs………………………………………………………41

Leaf lateral veins at least 11 pairs………………………………………………….…42
41. Leaf blade broadly ovate, midrib prominent above; inner bark bright plumred………...
…………………………………………………………..43. H. wyattsmithii (in part)
Leaf blade narrowly ovatelanceolate, midrib flat above; inner bark yellowish brown….
……………………………………………………………………….20. H. megacarpa
42. Leaf lateral veins at least 16 pairs……………………………………..…1. H. aequalis

Leaf lateral veins at most 15 pairs………………………………………………….…43
43. Leaf blade dull below; lateral veins 11–13 pairs………………….....34. H. rudiformis
Leaf blade drying chocolatebrown below, not dull; lateral veins 13–15 pairs……….….
……………………………………………………………………………25. H. nervosa
44. Leaf lateral veins at most 9 pairs, midrib sunken above; stiltrooted river bank tree…….
…………………………………………………………………………7. H. centipeda
Leaf lateral veins at least 10 pairs, midrib raised above; not as above.......................…45
45. Leaf lateral veins and midrib more prominent above than below.....27. H. obscurinerva
Leaf lateral veins and midrib more prominent below than above………………..….46
46. Leaf blade distinctly bullate between intercostal veins…………..….6. H. bullatifolia

Leaf blade flat to somewhat convex between veins above……………………….…47
47. Leaf blade larger, 27–46 × 8–15 cm, base cordate…………..…13. H. enicosanthoides
Leaf blade smaller, 12–28 × 4–9 cm, base not cordate……………………………..…48
146
48. Leaf blade drying reddish brown, lateral veins sunken above; young parts glabrescent…
………………………………………………………………………..33. H. pterygota
Leaf blade drying greyish green, lateral veins flat above; young parts pubescent, white
pruinose or silvery lepidote……………………………………….…29. H. pachycarpa
1. Hopea aequalis P.S.Ashton

(Latin aequalis = equal; the fruit calyx lobes)
Gard. Bull. Sing. 22 (1967) 271, op. cit. (1968) 46, op. cit. (1982) 409; Anderson op. cit. (1980) 114.
Type: Ilias S 15881, Borneo, Sarawak, Bintulu district, Nyabau FR (holotype K; isotypes KEP, L).
Synonym: Hopea ‘sp. nov. aff. H. pachycarpa’ Meijer & Wood op. cit. 229.
Subcanopy tree, to 20 m tall, to 20 cm diameter, with narrow buttresses, flying buttresses

and stilt roots. Bark smooth. Living surfaces glabrous but for the puberulent midrib above.
Twigs c. 2 mm diameter apically. Leaves chartaceous, drying greyish brown above,
yellowish brown below; blade oblonglanceolate, 13–25 × 5.5–8 cm, base equal, obtuse,
apex acuminate, acumen to 1 cm long, tapering; midrib shallowly furrowed but evident
above, slender and prominent below; venation hopeatype; lateral veins 16–20 pairs,
slender but prominent below; intercostal venation densely scalariform, hardly raised; petiole
1.5–1.8 cm long, c. 2 mm diameter. Inflorescences to 9 cm long, axillary, unbranched or
singly branched; rachis c. 1 mm diameter; flowers remote; bracts subpersistent. Flowers
unknown. Fruits: calyx lobes subequal, ovate, subacute, saccate, to 1.8 × 1.6 cm. Nuts
ovoid, to 2.3 × 1.5 cm, acute, frequently resincoated.
Vernacular name. Sabah—selangan sama (preferred name).
Distribution. Endemic in Borneo; uncommon. In Sabah once recorded from Sandakan

district (SAN 38797) and in Sarawak known from Nyabau FR, Bintulu district (the type) and
from Bt. Mersing, Tatau district (e.g., S 22493). Also occurring in Brunei (Cowley 137).
Ecology. In mixed dipterocarp forest on low hills and near streams, at altitude below 100 m.
Critically endangered owing to its disappearing habitat.
2. Hopea altocollina P.S.Ashton

(Latin, altus = high, collinus = concerning hills; inhabitant of hill sites)
PROSEA op. cit. 244; Newman et al. op. cit. 145. Type: Ashton BRUN 1030, Borneo, Sarawak,
Limbang district, Bt. Antu, Ulu Sembayang (holotype K; isotypes KEP, L).
Tall main canopy or low emergent tree, to 50 m tall, to 1.8 m diameter; bole tall; buttresses
thin, convex, to 4 m tall; crown evenly hemispherical with many ascending branches. Bark
tawny brown, shallowly fissured and powdery oblongflaky. Perianth sericeous outside;
parts otherwise glabrous. Twigs c. 2 mm diameter apically. Leaves thinly coriaceous,
drying yellowish brown with dark midrib below; blade lanceolate, 7–10 × 3–4.5 cm, base
obtuse, with revolute margin and appearing cuneate, apex caudate, with tapering acumen to
1 cm long; midrib slender, slightly raised above and below; venation dryobalanoid; main
lateral veins c. 16 pairs, elevated below; petiole 1–1.3 cm long, slender, somewhat
geniculate. Inflorescences to 8 cm long, terminal or in axillary pairs, singly branched.
147
Flowers: petals oblong, cream; stamens c. 15, connectival appendage c. 2x as long as

anther; ovary and stylopodium cylindrical, with punctate obtuse apex and short style.
Fruits: pedicels to 2 mm long; calyx lobes unequal, 2 longer lobes spatulate, obtuse, to 4.5
× 0.8 cm, tapering to 3 mm wide above the saccate base, 3 shorter ones suborbicular,
obtuse, to 0.4 × 0.4 cm, shorter than the nut. Nuts ovoid, to 0.7 × 0.5 cm, apiculate.
Vernacular name. Sarawak—luis gunung (preferred name).
Distribution. Endemic in Borneo; recorded in Sabah from Ulu Mendalong, Sipitang district
(e.g., SAN 16736) and in Sarawak from Bintulu, Kapit, Limbang, Marudi, and Tatau
districts (e.g., S 21267, S 22461, S 36185, S 49026, and S 68645).
Ecology. In upper dipterocarp forest; locally common on clayrich soils, at 800–1000 m

altitude. Vulnerable owing to logging.
3. Hopea andersonii P.S.Ashton

(J.A.R. Anderson, former Forest Officer in Sarawak and Brunei)
Gard. Bull. Sing. 22 (1967) 272, op. cit. (1968) 46, op. cit. (1982) 425; PROSEA op. cit. 258. Type:
Anderson S 11096, Borneo, Sarawak, Bau district, Kuching road (holotype K; isotypes KEP, L).
Mediumsized canopy tree, to 40 m tall, to 1.5 m diameter, with even hemispherical crown;
buttresses to 4 m tall, prominent, convex, c. 4 cm thick. Bark fawnbrown or coppery
brown, vertically cracked and oblong flaky or thinly flaky. Sapwood cream; heartwood
coffeebrown. Parts of flower exposed in bud and ovary densely pale greyish puberulent;
fruit calyx caducously so; living surfaces otherwise glabrous. Twigs c. 1 mm diameter
apically, slender. Leaves coriaceous, drying tawny with venation dark reddish brown below
or pale greyish brown; blade lanceolatefalcate to elliptic or narrowly ellipticlanceolate,
5–14 × 2–6 cm, base distinctly unequal, obtuse adaxially, cuneate abaxially, apex
acuminate, acumen to 2 cm long, slender, frequently falcate; midrib evident but flat above,
prominent below; venation hopeatype; lateral veins 9–12 pairs, arched, slender, elevated
below, with prominent pored, not swollen, pubescent, axillary domatia; intercostal venation
hardly elevated; petiole 0.5–1 cm long. Inflorescences to 12 cm, terminal or in axillary
fascicles, singly branched, branches with to 9 dense secund flowers; bracts fugaceous.
Flowers: buds ellipsoid, to 3 × 2 mm; petals oblong, cream; stamens c. 15, anthers ellipsoid,
connectival appendage to 2½x the length of anther; ovary and stylopodium cylindrical,
slightly constricted medially, style short columnar. Fruits: pedicels to 3 mm long, slender;
calyx lobes unequal, 2 longer lobes oblong, obtuse, to 6 × 2 cm, tapering to 3 mm wide
above the saccate base, 3 shorter ones to 0.4 × 0.3 cm, shorter than the nut. Nuts ovoid, to
0.8 × 0.5 cm, glabrous, retaining the truncate apical stylopodium and mucronate style
remnant.
Vernacular names. Sabah—selangan (preferred name). Sarawak—luis somit (preferred

name).
Notes. Two subspecies, subsp. andersonii and subsp. basalticola, are recognised in Sabah
and Sarawak.
148
Key to subspecies
Bark becoming fawnbrown, vertically cracked and oblongflaky. Leaf blade lanceolate
falcate to elliptic, drying tawny with venation dark reddish brown below……………………
subsp. andersonii
Endemic in Sarawak and recorded from Bau, Kuching, Marudi, Serian, and Sri Aman
districts (e.g., S 10038, S 10353, S 16182, S 22876, and S 32661). Locally frequent on
limestone karst, at altitudes to 400 m. Not vulnerable.
Bark coppery brown, thinly flaky. Leaf blade frequently narrowly ellipticlanceolate, drying
pale greyish brown……….……………………………………………………….…………...
subsp. basalticola P.S.Ashton
(Latinised English, basalt (a volcanic rock), cola = growing; growing on basalt
derived soil)
Gard. Bull. Sing. 22 (1967) 273, op. cit. (1982) 426. Type: Sibat S 25005, Borneo, Sarawak,
Tatau district, Bt. Mersing, Anap (holotype K; isotypes KEP, L).
Endemic in Borneo; recorded in Sabah from Kinabatangan and Tawau districts (e.g.,
FMS 35387, SAN 23278 and SAN 28649), and in C Sarawak from Marudi, Miri and
Tatau districts (e.g., S 13798, S 23481 and S 25040). Also occurring in E Kalimantan
(e.g., bb. 13232, Kostermans 5431 and Kostermans 6006). In mixed dipterocarp forest
on deep friable clay soils, over baserich rocks, at altitudes to 400 m. Vulnerable.
4. Hopea beccariana Burck Plate 3E.

Ann. Jard. Bot. Buitenz. 6 (1887) 240; Merrill, EB (1921) 401; Masamune, EPB (1942) 489;
Symington op. cit. (1943) 122; Browne op. cit. 120; Ashton op. cit. (1964) 95, op. cit. (1968) 48, op.
cit. (1982) 407; Meijer & Wood op. cit. 207; Burgess op. cit. 128; Anderson op. cit. (1980) 115;
PROSEA op. cit. 244; Coode et al. (eds.) op. cit. 71; Newman et al. op. cit. 145. Type: Beccari PB
1177, Borneo, Sarawak (holotype BO). Synonyms: Hancea beccariana (Burck) Pierre op. cit. (1891)
t. 244; Hopea nicholsonii F.Heim op. cit. (1891) 973; Hopea intermedia King op. cit. 126, p.p.;
Balanocarpus ovalifolius Ridl., J. Fed. Mal. Str. Mus. 10 (1920) 130, p.p.
Canopy or shortly emergent tree, to 45 m tall, to 1.6 m diameter, with diffuse hemispherical
crown and short stout buttresses. Bark generally becoming deeply vsection fissured, dark
chocolatebrown. Sapwood yellowish brown; heartwood chocolatebrown, hard. Exposed
young living parts and ovary apex caducous puberulent, more or less waxy glaucescent;
parts otherwise glabrous. Twigs c. 1 mm diameter apically, slender. Leaves thinly
coriaceous, drying pale yellowish brown including venation; blade broadly ovate, 5–8 ×
2.2–4.5 cm, base cuneate, frequently subequal, margin flat, apex caudate, acumen to 1.5 cm
long; midrib slender, elevated on both surfaces; venation dryobalanoid, indistinct, hardly
raised, drying not blackish; main lateral veins c. 8 pairs; intercostal venation obscure,
reticulate; petiole 1.2–1.7 cm long, slender, somewhat geniculate. Inflorescences to 7 cm
long, generally axillary, singly branched, branches with to 5 secund flowers. Flowers: buds
small, ellipsoid, subsessile; petals linear, cream; stamens c. 15, connectival appendage c. 2x
as long as anther; ovary and stylopodium hourglassshaped, tapering into short style.
Fruits: calyx lobes unequal, 2 longer lobes spatulate, obtuse, to 3.5 × 1 cm, tapering to 2
mm above the saccate base, 3 shorter ones ovate, acute, to 0.7 cm long, shorter than nut.
Nuts broadly ovoid, to 0.9 × 0.5 cm, with 1 mm short tapering style remnant.
149
Vernacular names. Sabah—selangan penak (preferred name). Sarawak—cengal pasir

(preferred name).
Distribution. Peninsular Thailand, Peninsular Malaysia and Borneo. In Sabah widespread

and recorded from Keningau, Kinabatangan, Kota Kinabalu, Kota Merudu, Kudat, Labuk
Sugut, Lahad Datu, Ranau, Sandakan, Semporna, Sipitang, Tawau, and Tenom districts
(e.g., SAN 15483, SAN 18545, SAN 25334, SAN 38703, SAN 47353, and SAN 100048) and
in Sarawak from Bintulu, Kapit, Kuching, Lawas, Limbang, Lundu, Marudi, Miri, and
Serian districts (e.g., S 12375, S 15212, S 15786, S 25480, and S 37808). Also occurring in
Brunei (e.g., BRUN 540, FMS 35452 and S 5830).
Ecology. Locally frequent in mixed dipterocarp forest, on dry, usually deep and sandy soils
on ridges. Especially in the coastal hills but occasionally at altitudes to 1200 m. Occurring
in Lambir NP; vulnerable owing to land conversion and logging.
Notes. The leaves closely resemble those of H. latifolia which however retains smooth bark.
They are distinguished when dry from those of H. dryobalanoides by their sandy buff
brown colour and pale lateral veins and midrib below.
5. Hopea bracteata Burck

(Latin, bractea = a thin metal plate; the subpersistent bracts)
Ann. Jard. Bot. Buitenz. 6 (1887) 239; Ashton op. cit. (1964) 97, op. cit. (1968) 48, op. cit. (1978) 31,
op. cit. (1982) 414; Meijer & Wood op. cit. 229; Anderson op. cit. (1980) 115; PROSEA op. cit. 245;
Coode et al. (eds.) op. cit. 71. Type: Teysmann HB 8265, Borneo, W Kalimantan (holotype BO).
Synonyms: Balanocarpus curtisii King op. cit. 158; B. bracteatus (Burck) Merr. op. cit. (1921) 407;
Hopea minima Symington, Gard. Bull. S. S. 10 (1939) 337.
Subcanopy, occasionally low canopy tree, to 35 m tall, to 60 cm diameter, with flying

buttresses and stilt roots; crown frequently persistently monopodial, with somewhat pendent
branches in juveniles. Bark smooth. Young twig, buds and petiole shortly persistently
greyish puberulent; floral sepals fimbriate; parts otherwise glabrous. Twigs c. 0.7 mm
diameter apically, very slender. Leaves chartaceous, drying dull greyish green; blade
elliptic to narrowly ovate, 2.5–6 × 0.7–2 cm, base cuneate, subequal, apex acuminate,
acumen to 1.5 cm long; midrib obscurely sunken above, slender and elevated below;
venation subdryobalanoid; main lateral veins c. 11 pairs; intercostal venation reticulate,
distinct below; petiole 0.2–0.4 cm long, slender. Inflorescences racemose, terminal or in
axillary pairs, to 9 cm long, slender, remotely irregularly singly or doubly branched,
ascending branches zigzag bearing to 5 remote flowers; bracts subpersistent; bracteoles
deltoid, to 2 mm across, subpersistent. Flowers: buds small, broadly ovoid, subsessile;
petals oblong, deep winered, strongly contorted; stamens c. 15, anthers subglobose,
connectival appendage c. 2x as long as anther; ovary and stylopodium cylindrical, truncate,
style short. Fruits: calyx lobes subequal, ovate, obtuse, to 0.5 × 0.5 cm, conceiling all but
the apex of nut. Nuts ovoid, to 0.9 × 0.6 cm, with minutely truncate apex.
Vernacular names. Sabah—selangan (preferred name). Sarawak—luis padi (preferred

name).
150
Distribution. Peninsular Malaysia and Borneo. In Sabah known from Keningau,

Kinabatangan, Labuk Sugut, Nabawan, Ranau, Sandakan, and Tawau districts (e.g.,
Kamaruddin KMS 3433, SAN 38944, SAN 83076, SAN 83408, SAN 128811, and SAN
130025); locally abundant throughout Sarawak and recorded from Belaga, Bintulu,
Kuching, Lundu, Miri, and Serian districts (e.g., S 13347, S 18305, S 27172, S 43517, and S
49885). Also occurring in Brunei (e.g., BRUN 322, BRUN 328, BRUN 435, FMS 30618, and
S 5713).
Ecology. Very local but often abundant in mixed dipterocarp forest on both clay and sandy
soils, at altitudes to 650 m. Locally common in Lambir NP but vulnerable elsewhere.
6. Hopea bullatifolia P.S.Ashton

(Latin, bullatus= blistered, folius = a leaf; the quilted leaf blade)
Type: Mashor SA 522, Borneo, Sarawak, Belaga district, Sg. Danum (holotype K; isotype KEP).
Subcanopy tree, to 20 tall, to 20 cm diameter; buttresses low, thin. Bark smooth. Young
exposed parts including petiole and midrib below densely evenly persistently tawny
pubescent, venation and lamina below sparsely so; parts otherwise apparently glabrous.
Twigs c. 2 mm diameter apically. Leaves thinly coriaceous, prominently bullate between
the intercostal veins, drying chocolatebrown and darker above, not silvery below; blade
oblong, 16–34 × 4.5–9 cm, base markedly unequal, subcordate, apex with slender acumen
to 1 cm long; midrib prominent on both surfaces; venation hopeatype; lateral veins 17–26
pairs, slender but prominent below, sunken above, often anastomosing within margin;
intercostal venation scalariform, lax, raised below, sunken above; petiole 0.3–0.6 cm long,
stout. Inflorescence and flower unknown. Fruits subsessile; calyx lobes unequal, 2 longer
lobes spatulate, obtuse, to 8 × 1.5 cm, tapering to 3 mm above the saccate base, 3 shorter
ones to 1.5 cm long, lanceolate, enclosing and hiding the nut. Nuts ovoid, to 1 × 0.7 cm,
apiculate.
Vernacular name. Sarawak—luis melencur (preferred name).
Distribution. Endemic in Borneo. Known in Sarawak from Belaga, Kapit, Lubok Antu,
Sibu, Song, Sri Aman, and Tatau districts (e.g., S 17730, S 43453, S 45130, S 69663, and
SFN 35726). Also occurring in S Kalimantan.
Ecology. Rare, in mixed dipterocarp forest on low shale hills, at altitudes to 200 m.
Probably endangered.
7. Hopea centipeda P.S.Ashton Fig. 13.

(Latin, centi = hundred, pes = a foot; the stilt roots)
Coode et al. (eds.) op. cit. 71. Type: Murthy & Ashton S 23342, Borneo, Sarawak, Marudi district,
Ulu Dapoi, Tinjar (holotype K; isotypes KEP, L). Synonym: Hopea acuminata auct. non. Merr.:
Ashton op. cit. (1964) 94, Coode et al. (eds.) op. cit. 71.
151
Fig. 13. Hopea centipeda. A, fruiting leafy twig; B, detail of indumentum on lower leaf
surface; C, detail of indumentum on upper leaf surface; D, flower bud; E, open flower; F,
gynoecium; G, adaxial view of petal with stamens; H, abaxial view of petal; I, adaxial view
of stamens; J, abaxial view of stamens; K, fruit; L, partially exposed nut; M, fruit with the
large calyx lobes removed. (A–C and K–L from SAN 124740, D–J and M from S 23342.)
152
Canopy tree, to 30 m tall, to 60 cm diameter, often remaining monopodial with pendent

branches and diffuse foliage; bole with abundant flying buttresses and stilt roots to 3 m;
Sapwood soft, pale. Bark smooth. Young leaf shoots including leaf venation below and
petiole, inflorescence and parts of flower exposed in bud persistently shortly greyish brown
puberulent; parts otherwise glabrous. Twigs c. 1 mm diameter apically, slender. Leaves
chartaceous, drying pale yellowish grey; blade lanceolate, 5.5–9 × 1.5–3.5 cm, base
subequal, cuneate, apex caudate, acumen to 1.5 cm long; midrib prominent below, narrow
and sunken above; venation hopeatype; lateral veins 7–9 pairs, prominent below,
somewhat arched, with prominent tomentose axillary domatia; intercostal venation slender,
subscalariform; petiole 0.4–0.7 cm long, slender. Inflorescences to 2 cm long, axillary,
flowers dense; bracts fugaceous. Flowers: buds lanceolate, to 6 × 2 mm; petals cream to
pink at base, oblong; stamens c. 15, anthers subglobose, connectival appendage c. 3x the
length of anther; ovary and stylopodium pyriform, style prominent, as long as ovary and
equal to stylopodium. Fruits: pedicel c. 1 mm long, slender; calyx lobes unequal, 2 longer
lobes spatulate, obtuse, to 3 × 0.8 cm, tapering to 1.5 mm above the saccate base, 3 shorter
ones ovate, to 0.4 × 0.3 cm. Nuts ovoid, to 0.4 × 0.3 cm, apiculate.
Vernacular name. Sarawak—luis daun berbulu (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from Nabawan and Tawau districts
(e.g., SAN 62794 and SAN 124740) and in Sarawak from Belaga, Bintulu, Kapit, Marudi,
and Miri districts (e.g., S 22649, S 22682, S 23010, S 23012, and S 43577). Also occurring
in Brunei (e.g., BRUN 125, BRUN 127 and S 5750).
Ecology. Locally common along the banks of swiftflowing rivers, on shale and silt, at
altitudes to 300 m. Frequently flowering and fruiting. Vulnerable.
8. Hopea cernua Teijsm. & Binn.

(Latin, cernuus = slightly drooping; the flowers)
Nat. Tijd. Ned. Ind. 29 (1867) 252; Merrill op. cit. (1921) 402; Masamune op. cit. 490; Ashton op. cit.
Kessler & Sidiyasa op. cit. 97; Coode et al. (eds.) op. cit. 71; Newman et al. op. cit. 147. Lectotype
(designated here): Binnendijk s.n., Sumatra (hololectotype K). Synonyms: Hancea cernua (Teijsm. &
Binn.) Pierre op. cit. (1891) t. 244; Hopea argentea Meijer, Act. Bot. Neerl. 12 (1963) 348, Meijer &
Wood op. cit. 207, Burgess op. cit. 128, Ashton op. cit. (1968) 47.
Canopy tree, to 40 m tall, to 80 cm diameter, with prominent flying buttresses and stilt
roots. Bark becoming cracked and eventually vsection fissured, chocolatebrown; inner
bark pale brown. Leaf bud, young twig and petiole fugaceous greyish brown puberulent;
parts otherwise glabrous. Twigs c. 1 mm diameter apically, slender. Leaves thinly
coriaceous, often somewhat silvery lepidote below, drying greyish tawny with the veins
often darker below; blade elliptic to ovate, 5–15 × 2–5 cm, base cuneate, unequal, margin
frequently narrowly shallowly revolute, apex with acumen to 0.6 cm long; midrib terete
below, slender and elevated above, not drying black; venation dryobalanoid, slender but
generally distinctly elevated below, often with a few axillary pubescent porelike domatia;
main lateral veins 10–12 pairs, distinctly raised below, basal pairs short, drying blackish;
petiole 0.7–0.9 cm long, slender. Inflorescences terminal or axillary, singly branched, to 3
cm long. Flowers: buds spindleshaped, to 5 × 4 mm, relatively large; petals lanceolate,
cream; stamens 15–18, anthers subglobose, connectival appendage 2–3x the length of
anther; ovary ovoid, without stylopodium, style columnar, c. 1½x as long as ovary, villous
153
towards base. Fruits: pedicels c. 2 mm long; calyx lobes unequal, 2 longer lobes spatulate,
obtuse, to 6.5 × 1.2 cm, tapering to 4 mm obove the saccate but rather broad, 0.5 × 5 mm
base, 3 shorter ones lanceolate, to 1.5 cm long, conceiling the nut. Nuts ovoid, to 0.7 × 0.5
cm, apiculate.
Vernacular names. Sabah—selangan urat (preferred name). Sarawak—luis timbul

(preferred name).
Distribution. Sumatra and Borneo. Throughout Sabah and recorded from Kinabatangan,
Lahad Datu, Ranau, Sandakan, Sipitang, Tawau, and Tenom districts (e.g., SAN 16367, SAN
22090, SAN 23260, SAN 24252, SAN 35861, and SAN A 4049) and in Sarawak known from
Bau, Bintulu, Kapit, Kuching, Lundu, Miri, and Serian districts (S 15448, S 24040, S 25647,
S 32660, and S 60129). Also occurring in Brunei (e.g., BRUN 2359 and BRUN 2532) and E
Kalimantan (e.g., Kostermans 6032, Kostermans 13838, Sidiyasa 439, and Sidiyasa 539).
Ecology. In forest on organic soils over limestone in the lowlands and in upper dipterocarp
forest, at 1000–1650 m altitude; also on igneous rocks. Very local, vulnerable.
9. Hopea coriacea Burck

(Latin, coriaceus = leathery; the leaf blade)
Ann. Jard. Bot. Buitenz. 6 (1887) 237; Merrill op. cit. (1921) 402; Masamune op. cit. 490; Ashton op.
cit. (1978) 28, op. cit. (1982) 398; Anderson op. cit. (1980) 115; PROSEA op. cit. 260; Coode et al.
(eds.) op. cit. 71; Newman et al. op. cit. 148. Type: Teijsmann s.n., Borneo, Kalimantan (holotype
BO; isotype L). Synonyms: Hopea kelantanensis Symington, J. Malay. Br. Roy. As. Soc. 19 (1941)
144; H. garangbuaya P.S.Ashton, Gard. Bull. Sing. 19 (1962) 256, op. cit. (1964) 101, op. cit. (1968)
51.
Large canopy or low emergent tree, to 45 m tall, to 1.5 m diameter; crown dense
hemispherical; buttresses to 1.5 m tall, to 8 cm thick, concave or convex; stilt roots few or
none. Bark smooth to flaky or deeply vsection fissured, greyish to chocolatebrown; inner
bark pale brown or pink. Sapwood strawyellow, hard; heartwood dark chocolatebrown.
Parts glabrous but for pubescent parts of petals exposed in bud. Twigs c. 2 mm diameter
apically, stout. Leaves thickly coriaceous, drying pale chocolatebrown below, yellowish
brown above; blade broadly ovate or suborbicular, 11–16 × 6–10 cm, base equal, obtuse,
margin revolute, apex acuminate, acumen to 1.2 cm long, narrow; midrib prominently terete
below, slightly elevated though stout above; venation dryobalanoid, sharply elevated below,
strongly arched; main lateral veins 8–11 pairs or sometimes at most 4 pairs, with short
intermediates; intercostal venation scalariform, distinctly elevated below; petiole 2–2.5 cm
long, stout. Inflorescences terminal or in axillary pairs, to 9 cm long, singly branched,
branchlets bearing to 6 secund flowers. Flowers: buds ellipsoid, to 3 × 2.5 mm, large,
distinctly pedicellate; petals lanceolate, to 1.3 cm long, cream; stamens c. 15, anthers
oblong, connectival appendage 2–3x the length of anther, tuberculate towards base; ovary
ovoid, style as long as ovary, columnar, setose towards base and onto ovary apex,
stylopodium obscure. Fruits: calyx lobes unequal, 2 longer lobes to 7 × 1.5 cm, hardly
tapering to the narrowly auriculate saccate base, 3 shorter ones ovate to lanceolate, to 2 ×
1.2 cm, conceiling the nut, similar at base. Nuts narrowly ovoid, to 1.8 × 0.9 cm, with to 2
cm long filiform style remnant, often resinous.
Vernacular name. Sarawak—garang buaya (preferred name).
154
Distribution. Peninsular Malaysia and Borneo. In Borneo known in Sarawak from Bintulu,
Limbang, and Lundu districts (e.g., S 10395, S 12610, S 18340, S 18855, and S 66799). Also
occurring in Brunei (e.g., BRUN 713, BRUN 2006A, BRUN 3347, and S 5768) and
Kalimantan (e.g., the type, bb. 7699 and bb. 35251).
Ecology. In mixed dipterocarp forest and along the banks of small rivers, on yellow sandy
soils, at altitudes below 400 m. Rare and critically endangered by logging and forest
conversion.
10. Hopea depressinerva P.S.Ashton

(Latin, depressus = sunken, nervus= nerve/vein; the sunken veins on leaf upper surface)
PROSEA op. cit. 260. Type: Anderson S 15439, Borneo, Sarawak, Lundu district, Bt. Gebong
Subcanopy or canopy tree, to 25 m tall, to 50 cm diameter. Bark brown, becoming cracked

along lines of lenticels; damar exudation not conspicuous. Parts glabrescent but for densely
buffpubescent inflorescence and parts of flower exposed in bud. Twigs 1–2 mm diameter
apically, slender. Leaves thinly coriaceous, undulate, drying pale greyish brown, not pale
lepidote below; blade lanceolate to narrowly elliptic, 5–13 × 2–5 cm, base cuneate, equal,
apex with slender acumen to 1.5 cm long; venation hopeatype; lateral veins 6–8 pairs,
slender but distinctly raised below, obscure and sunken above, domatia obscure or absent;
intercostal venation densely scalariform, slender, not elevated; petiole 1.1–1.3 cm long,
slender. Inflorescences terminal or axillary, to 7 cm long, singly branched, branchlets
bearing to 5 secund dense flowers; bracts fugaceous. Flowers: buds ellipsoid, to 3 × 2 mm;
petals elliptic, pink; stamens c. 10, anthers subglobose, connectival appendage c. 2x the
length of anther; ovary and stylopodium cylindrical, truncate, style short. Mature fruits
unknown; calyx lobes unequal, 2 larger lobes aliform.
Distribution. Endemic in Borneo; known so far from G. Pueh and Bt. Gebong, Lundu
district in W Sarawak (e.g., S 395, S 15516 and the type).
Ecology. Very local, apparently uncommon in mixed dipterocarp forest on deep sandy loam
soils over granodiorite, at altitudes to 500 m. Conservation status unknown, but likely
vulnerable.
11. Hopea dryobalanoides Miq. Fig. 14.

(Greek, dryobalanoides = resembling Dryobalanops; the leaf venation)
Fl. Ned. Ind., Suppl. (1862) 492; Symington op. cit. (1939) 345; Browne op. cit. 120; Ashton op. cit.
(1963) 259, op. cit. (1964) 50, op. cit. (1968) 50, op. cit. (1982) 402; Meijer & Wood op. cit. 209;
97; Coode et al. (eds.) op. cit. 71; Newman et al. op. cit. 149. Type: Teijsmann s.n., Sumatra,
Pariaman, Padang (holotype U). Synonyms: Hancea dryobalanoides (Miq.) Pierre op. cit. t. 244;
Hopea sarawakensis F.Heim op. cit. (1891) 971; Hopea borneensis F.Heim op. cit. (1891) 972.
Large canopy or low emergent tree, to 45 m tall, to 1.3 m diameter; buttresses to 2 m tall,
thin, concave, spreading; stilt roots present on young trees. Bark becoming fawn brown,
vertically cracked and powdery thinly oblong flaky, with coxcombs of glassy dammar; inner
155
Fig. 14. Hopea dryobalanoides. A, flowering leafy twig; B, detail of venation on lower leaf
surface; C, flower bud; D, side view of open flower; E, top view of open flower; F, adaxial
view of petal with stamens; G, abaxial view of petal; H, adaxial view of stamens; I, abaxial
view of stamens; J, longitudinal section of gynoecium; K, fruit; L, fruit with the large calyx
lobes removed. (A–J from SAN 15186, K–L from SAN 100048.)
156
bark pale brown or pink. Young parts including leaf below, petiole, inflorescence and parts
of flower exposed in bud densely greyish brown fugaceouspuberulent. Twigs c. 1 mm
diameter apically, slender, drying blackish. Leaves chartaceous or thinly coriaceous, drying
dull leaden brown below; blade ovatelanceolate, 5–12 × 1.5–4.5 cm, base cuneate, equal to
subequal, apex subcaudate, acumen to 2 cm long; midrib raised and terete or sharp on both
surfaces, more so below, drying black or not; venation dryobalanoid; main lateral veins 8–
12 pairs, with a few short intermediates, slender but distinct and somewhat elevated below,
arched, drying paler or darker than the blade, sometimes with a few small glabrous pore
like axillary domatia; petiole 0.5–1 cm long, slender, drying blackish. Inflorescences
terminal or in axillary pairs, lax, singly branched, branchlets to 5 cm long, bearing to 6
flowers. Flowers: buds small, broadly ovoid; petals lanceolate, creamyellow; stamens c.
15, anther subglobose, connectival appendage c. 2x the length of anther; ovary ovoid,
stylopodium obscure, style filiform, as long as ovary, villous towards base. Fruits: pedicels
to 2 mm long; calyx lobes unequal, 2 longer lobes strongly twisted, spatulate, subacute, to
6.5 × 1.5 cm, tapering to 5 mm above the saccate base, 3 shorter ones broadly ovate, obtuse
or subacute, to 0.8 × 0.6 cm, shorter than the nut. Nuts broadly ovoid, to 1 × 0.8 cm, with to
1.5 mm filiform style remnant.
Vernacular names. Sabah—selangan daun kapur (preferred name). Sarawak—mata

kucing hitam (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah widespread and recorded
from Beaufort, Kinabatangan, Kota Kinabalu, Kudat, Labuk Sugut, Lahad Datu, Ranau,
Semporna, Tawau, and Tenom districts (e.g., SAN 16837, SAN 19335, SAN 22654, SAN
22849, and SAN 24514) and in Sarawak from Belaga, Bintulu, Kapit, Kuching, Lawas,
Limbang, Lundu, Miri, Serian, Sibu, and Tatau districts (e.g., S 7972, S 15249, S 22476, S
28172, and S 43827). Also occurring in Brunei (e.g., BRUN 3376, BRUN 5230 and BRUN
5376) and E Kalimantan (e.g., Kostermans 13518, Kostermans 13867 and Sidiyasa 707).
Ecology. Locally abundant in mixed dipterocarp forest on clayrich soils, especially on

intermediate to basic igneous and volcanic rocks but also sedimentaries, at altitudes to 600
m. Well represented in Kinabalu NP, Sepilok FR and Danum Valley Conservation Area in
Sabah and Lambir and Mulu NPs in Sarawak; not vulnerable.
Uses. Formerly a major source of damar mata kucing, yielding a high quality resin.
Notes. One of the largest of the dryobalanoid Hopea species, with flaky bark and prominent
thin buttresses. The leaf dries a distinctive greenish grey with distinct curved lateral veins,
black drying midrib below and long black petiole (cf. H. beccariana).
12. Hopea dyeri F.Heim

(J.T. ThiseltonDyer, 1824–1928, a British botanist, one time Director of the Royal Botanic
Gardens, Kew)
Bull. Mens. Soc. Linn. Paris 2 (1891) 972; Merrill op. cit. (1921) 402; Symington op. cit. (1939) 353;
(1982) 407; Meijer & Wood op. cit. 210; Burgess op. cit. 129; Anderson op. cit. (1980) 116; PROSEA
op. cit. 246; Coode et al. (eds.) op. cit. 71. Type: Beccari PB 2962, Borneo, Sarawak, Matang
(holotype P). Synonym: Hopea intermedia King op. cit. 126, p.p.
157
Mediumsized canopy tree, to 40 m tall, to 65 cm diameter; crown narrowly hemispherical;

bole frequently misshapen, with low narrow buttresses and a few stilt roots. Bark smooth,
becoming patchily flaky, sometimes with yellowish dammar exudations; inner bark pale
brown or pink. Sapwood yellowish. Bud, stipules, inflorescence, outside of petals and
domatia sparsely or densely greyish brown puberulent; twig, petiole and parts of calyx
exposed in bud caducously so. Twigs c. 1 mm diameter apically, slender, muchbranched.
Leaves coriaceous, frequently lepidote below, drying greyish brown; blade ovate
lanceolate, 2.7–7 × 1.2–2.5 cm, base unequal, cuneate, margin often slightly revolute, apex
caudate, acumen to 1.5 cm long; midrib terete and raised on both surfaces, more so above,
not drying black; venation dryobalanoid; main lateral veins 8–12 pairs with many
intermediates, distinct and somewhat elevated below, the longest subequal, basal pairs
short; petiole 0.5–0.8 cm long, slender. Inflorescences terminal or axillary, to 3 cm long,
singly or doubly shortly branched, branchlets bearing to 4 flowers. Flowers: buds small,
ovoid; petals narrowly lanceolate, cream; stamens c. 15, anthers subglobose, connectival
appendage c. 3x the length of anther; ovary and stylopodium pyriform, villous towards apex,
style short. Fruits: calyx lobes unequal, 2 longer lobes spatulate, narrowly obtuse, to 2.5 × 1
cm, tapering to 2.5 mm above the saccate base, 3 shorter ones ovate, acute to subacuminate,
to 0.7 × 0.4 cm, shorter than the nut. Nuts ovoid, to 0.9 × 0.4 cm, with vestigial stylopodium
and subacute style remnant.
Vernacular names. Sabah—selangan daun halus (preferred name). Sarawak—luis palit

(preferred name).
Distribution. Peninsular Malaysia and Borneo. In Sabah recorded from Keningau, Labuk
Sugut, Ranau, Sandakan, Sipitang, and Tawau districts (e.g., SAN 19452, SAN 21506, SAN
41082, SAN 66241, and SAN 99285) and in Sarawak from Bintulu, Kapit, Kuching, Lawas,
Lubok Antu, Miri, Samarahan, and Serian districts (e.g., S 15219, S 23962, S 24655, S
3180) and C and E Kalimantan (e.g., bb. 24781 and Ridsdale PBU 593).
Ecology. Locally common in mixed dipterocarp forest on sandy and sandy clay soils, on
low hills and along sandstone ridges, occasionally at altitudes to 1000 m but usually lower.
Occurring in Kubah NP, Sarawak; regeneration abundant; probably not vulnerable.
Notes. This species one of a group of smallleaved and smallfruited dryobalanoidveined

Hopea species, also notebly including H. ferruginea, H. micrantha and H. vesquei, which
can be difficult to identify on field characters alone. The leaf of H. micrantha is obtuse at
base, but that of the others is cuneate, and the leaf dries pink not coppery brown. The twigs
and petioles of H. vesquei are densely persistently puberulent, but the leaf is shorter and
broader with unequal base, and more coriaceous than the other species. Variability in leaf
size and texture of H. dyeri as recognised here suggests that fertile material of H. vesquei
may yet to be found east of Batang Lupar (Sarawak).
13. Hopea enicosanthoides P.S.Ashton

(Greek, resembling Enicosanthum, Annonaceae; the leaf)
Type: Ashton S 18116, Borneo, Sarawak, Bintulu district, Ulu Labang (holotype K; isotypes KEP, L).
158
Small tree, to 18 m tall, to 20 cm diameter; crown monopodial, with pendent branches and
leaves; buttresses low, thin; stilt roots few. Bark smooth. Young parts including petiole and
base of midrib above more or less caducous tawny pubescent. Twigs c. 3 mm diameter
apically, often somewhat compressed, ribbed below the petiole insertion. Leaves thinly
coriaceous, somewhat convex between the veins above, drying greenish to rustbrown;
blade very large, oblong, 27–46 × 8–15 cm, base unequal, cordate, apex with slender
acumen to 2.5 cm long; midrib evident but furrowed above, prominent below; venation
hopeatype; lateral veins 16–30 pairs, slender but prominent below; intercostal venation
densely scalariform, evident but unraised below; petiole 0.5–0.8 cm long, stout.
Inflorescences in axillary pairs or ramiflorous, slender, lax, to 12 cm long, singly
branched, branchlets bearing to 5 flowers. Flowers: unknown. Fruits subsessile; calyx
lobes unequal, 2 longer lobes broadly spatulate, obtuse, to 13 × 3 cm, tapering to 6 mm
above the saccate base, 3 shorter ones lanceolate, acute, to 2 cm long, similar at base,
conceiling the nut. Nuts ovoid, to 1 × 0.6 cm, with 2 mm long filiform style remnant.
Vernacular name. Sarawak—luis selukai (preferred name).
Distribution. Endemic in Borneo and known only from Bintulu, Kapit and Miri districts in
Sarawak (e.g., S 1467, S 1470, S 13200, S 18304, S 37686, and S 69631).
Ecology. On the banks of sluggish eutrophic rivers and moist slopes by streams, at altitudes
below 100 m, locally abundant; but killed by prolonged immersion in turbid water. Probably
endangered.
14. Hopea ferruginea Parijs

(Latin, ferrugineus = rustcoloured; the indumentum)
In Fedde, Rep. 33 (1933) 243; Symington op. cit. (1939) 349, op. cit. (1943) 125; Masamune op. cit.
490; Meijer & Wood op. cit. 211; Burgess op. cit. 129; Ashton op. cit. (1982) 404; PROSEA op. cit.
247; Coode et al. (eds.) op. cit. 71. Type: bb. 14459, Sumatra, Jambi (holotype BO; isotype L).
Mediumsized canopy tree, to 40 m tall, to 70 cm diameter, with thin buttresses and stilt
roots. Bark smooth, becoming thinly flaky, with clear globular dammar exudations; inner
bark pale brown or pink. Young twigs, petiole, midrib above, inflorescence, and parts of
flower exposed in bud sparsely but persistently greyish to rustbrown puberulent, or
glabrescent. Twigs c. 1 mm diameter apically, slender, blackish. Leaves thinly coriaceous,
drying greyish brown; blade narrowly ovatelanceolate, 4.5–7.5 × 1.5–4 cm, base cuneate,
unequal, shortly decurrent, apex caudate, acumen to 1.5 cm long; midrib terete, slender,
elevated on both surfaces, not drying black; venation dryobalanoid; main lateral veins c. 14
pairs, basal pairs short, with many subequal intermediates, somewhat raised below, with
prominent axillary domatia in juveniles; petiole 0.6–1 cm long, slender. Inflorescences
axillary, to 2 cm long, with short branchlets bearing to 4 secund flowers. Flowers: buds
ellipsoid, to 3 × 2 mm; petals lanceolate, pale yellow; stamens c. 15, anther subglobose,
connectival appendage c. 2x the length of anther; ovary ovoid, stylopodium obscure, style
columnar, as long as ovary, villous towards base or glabrous. Fruits: pedicels to 2 mm
long, very slender; calyx lobes unequal, 2 longer lobes spatulate, narrowly obtuse, to 3 ×
0.5 cm, tapering to 1.5 mm above the saccate base, 3 shorter ones ovateacuminate, to 0.8 ×
0.4 cm, closely clasping and conceiling the ripe nut. Nuts ovoid, to 0.7 × 0.4 cm, apiculate.
Vernacular name. Sabah—selangan mata kucing (preferred name).
159
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo common and

widespread. In Sabah recorded from Kinabatangan, Kota Belud, Kudat, Lahad Datu,
Sandakan, Sipitang, and Tawau districts (e.g., SAN 15358, SAN 19380, SAN 26590, and
SAN 67737) and in Sarawak from Lawas, Limbang and Miri districts (e.g., Nooteboom
2332, S 31527 and S 39619). Also occurring in Brunei (e.g., BRUN 3375) and E Kalimantan
(e.g., bb. 14974).
Ecology. In mixed dipterocarp forest on clayrich soils and along ridges in upper
dipterocarp forest, at altitudes to 750 m. Locally common and occurring in Kinabalu NP, but
vulnerable.
Notes. Sterile specimens of this species is not easily distinguishable from those of H. dyerii
(see there).
15. Hopea fluvialis P.S.Ashton

(Latin, fluviatilis = pertaining to rivers; the natural habitat)
Gard. Bull. Sing. 19 (1962) 254, op. cit. (1964) 100, op. cit. (1968) 51, op. cit. (1982) 399; Anderson
op. cit. (1980) 116; PROSEA op. cit. 247; Coode et al. (eds.) op. cit. 71. Type: Flemmich FMS 34523,
Brunei, Belait district, Sg. Melayan (holotype KEP).
Mediumsized leaning riverbank tree, to 25 m tall, to 80 cm diameter, with or without low

buttresses and stilt roots. Bark smooth; inner bark pale brown or pink. Sapwood whitish,
relatively soft. Young parts shortly densely greyish brown puberulent; indumentum
persistent on twig, bud, stipule, inflorescence and petiole, caducous elsewhere. Twigs c. 1.5
mm diameter apically. Leaves chartaceous to thinly coriaceous, drying greyish brown;
blade ovatelanceolate, somewhat falcate, 7–12 × 2.8–4.8 cm, base unequal, cuneate,
margin not revolute, apex subcaudate, acumen to 1.5 cm long; midrib stout, terete, hardly
raised below, more prominently so above, not drying black; venation dryobalanoid; main
lateral veins c. 10 pairs with many long intermediates, the basal pair extending to one third
the length of the blade margin; intercostal venation evident below, reticulate; petiole 0.7–1
cm long. Inflorescences in axillary clusters of 3, or rarely terminal, to 6 cm long, singly or
doubly branched, branchlets bearing to 7 secund flowers. Flowers: buds to 5 × 3 mm; petals
lanceolate, cream; stamens c. 15, anthers subglobose, connectival appendage c. 2x the
length of anther; ovary ovoid, stylopodium obscure, style columnar tapering, as long as
ovary. Fruits erect; pedicels to 2 mm long, slender; calyx lobes unequal, 2 longer lobes
broadly spatulate, obtuse, to 5 × 1 cm, subauriculate and hardly tapering into the saccate
base, 3 shorter ones unequal, lanceolate, acute, 1–2.5 cm long and conceiling the nut. Nuts
narrowly ovoid, to 1.1 × 0.6 cm, tapering to a short apiculus.
Vernacular name. Sarawak—luis air (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from Kinabatangan, Labuk Sugut and
Sandakan districts (e.g., SAN 37886, SAN 94100 and SAN 99580) and in Sarawak from
Belaga, Bintulu, Kapit, Lawas, Marudi, Miri, and Tatau districts (e.g., S 3433, S 15577, S
23489, S 25024, and S 41429). Also occurring in Brunei (e.g., BRUN 3394, BRUN 5668,
FMS 34492, Forman 1165, and Prance 30711) and Kalimantan (e.g., bb. 20448).
Ecology. Locally common in forest on clayrich river banks, both fast and sluggish, at
altitudes to 200 m. Fruiting frequently. Common in Mulu NP; probably not vulnerable.
160
16. Hopea griffithii Kurz

(W. Griffith, 1810–1845, surgeon in the British East India Company at Malacca, Peninsular
Malaysia)
J. As. Soc. Beng. 42, 2 (1873) 60; Symington op. cit. (1939) 343, op. cit. (1943) 127; Ashton op. cit.
(1968) 51, op. cit. (1982) 406; Anderson op. cit. (1980) 116; PROSEA op. cit. 248; Coode et al. (eds.)
op. cit. 72; Newman et al. op. cit. 150. Type: Griffith 717, Myanmar, Mergui, Tennaserim (holotype
CAL; isotype K). Synonym: Hancea griffithii (Kurz) Pierre op. cit. t. 248.
Canopy tree, to 40 m tall, to 60 cm diameter, with even hemispherical crown and thin, flying
buttresses and stilt roots to 1 m tall. Bark smooth becoming cracked in older trees. Parts
glabrous but for pubescent parts of petals exposed in bud. Twigs c. 1 mm diameter apically,
slender, muchbranched. Leaves coriaceous, shiny and drying purplish brown above, tawny
brown below; blade ovate to lanceolate, 4–9 × 1.7–4.5 cm, base cuneate, margin frequently
subrevolute, apex subcaudate, acumen to 1.5 cm long; midrib obscure, sunken above,
prominent below; venation dryobalanoid, with the veins hardly evident; main lateral veins
c. 9 pairs, with shorter intermediates, without domatia; intercostal venation obscure,
reticulate; petiole 0.8–1.5 cm long, slender. Inflorescences terminal or axillary, to 2.5 cm
long, singly branched, branchlets bearing to 5 secund flowers. Flowers: buds ellipsoid, to 2
mm long; petals lanceolate, dark red; stamens c. 15, anthers broadly ellipsoid, connectival
appendage c. 2x the length of anther; ovary and stylopodium stoutly pyriform, papillose
towards apex, tapering abruptly to the short columnar style. Fruits: pedicels c. 1 mm long;
calyx lobes unequal, 2 longer lobes spatulate, obtuse, to 3 × 0.5 cm, tapering to 2 mm above
the saccate base; 3 shorter ones linear, to 0.8 × 0.1 cm. Nuts ovoid, to 0.7 × 0.5 cm,
apiculate.
Vernacular name. Sarawak—luis jantan (preferred name).
Distribution. Peninsular Myanmar, Peninsular Thailand, Peninsular Malaysia, and Borneo.

In Borneo, rare in Sabah and known from Kota Marudu and Ranau districts (e.g., SAN
25334 and SAN 100048) but widespread in Sarawak with most records in the west, from
Kapit, Kuching, Lundu, Serian, and Simunjan districts (e.g., S 9338, S 10052, S 22267, S
5249) and W Kutei in NE Kalimantan (e.g., bb. 8205, bb. 20133 and Hallier 2225).
Ecology. In mixed dipterocarp forests on leached yellow clay soils, at altitudes to 500 m.
Locally common. Vulnerable.
17. Hopea kerangasensis P.S.Ashton

(from Iban word—kerangas = a type of soil not suitable for cultivating rice; the natural
habitat)
Type: Yakup S 8944, Borneo, Sarawak, Kuching district, Semengoh FR (holotype K; isotype L).
161
Subcanopy or small canopy tree, to 30 m tall, to 35 cm diameter; crown diffuse, at first

remaining monopodial; buttresses, flying buttresses and a few stilt roots to 1 m tall. Bark
smooth. Twig, petiole, midrib above, domatia below, bud, and inflorescence more or less
densely persistently tawny puberulent, more or less caducous. Twigs c. 0.5 mm diameter
apically, very slender. Leaves chartaceous, drying purplish brown above, deep tawny
below; blade ovate, 1.5–4.5 × 1–3 cm, base broadly cuneate, apex caudate, acumen to 1 cm
long; midrib shallowly sunken above, slender but prominent below; venation dryobalanoid;
main lateral veins c. 6 pairs with short intermediates, slightly sunken above, slender but
distinctly raised below, with prominent pubescent domatia; intercostal venation obscure,
reticulate; petiole 0.3–0.5 cm long, slender. Inflorescences axillary, to 1.2 cm long, singly
branched, branchlets bearing to 3 distichous flowers. Flowers: buds ovoid, to 1.5 × 1 mm;
petals lanceolate, cream; stamens c. 15, anthers subglobose, connectival appendage 2–3x the
length of anther; ovary ovoid, stylopodium obscure, style columnar, tapering, as long as
ovary. Fruits: pedicels c. 1 mm long; calyx lobes subequal, ovate, to 0.6 × 0.5 cm, acute,
saccate, appressed against the nut. Nuts exposed above calyx towards apex, ovoid, to 0.8 ×
0.5 cm, subacute.
Vernacular name. Sarawak—luis kerangas (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah recorded from Bt. Tawai,
Kinabatangan district (e.g., SAN 39349) and in Sarawak from Bau, Kuching, Lundu, and
Simunjan districts (e.g., S 2274, S 14926, S 15530, S 21299, and S 29030). Also occurring in
C Kalimantan (e.g., bb. 14959 and bb. 16033).
Ecology. Locally abundant in mixed dipterocarp forest on leached clayrich soils and in
kerangas on podsols, at altitudes below 300 m. Endangered.
18. Hopea latifolia Symington

(Latin, latus = wide, folius = leaf; with broad leaves)
Gard. Bull. S. S. 10 (1939) 360, op. cit. (1943) 131; Ashton op. cit. (1964) 102, op. cit. (1968) 52, op.
cit. (1982) 404; Anderson op. cit. (1980) 116; PROSEA op. cit. 249. Type: Mohamad FMS 8149,
Peninsular Malaysia, Pahang, Kuantan, Telok Siseh (holotype KEP).
Main canopy tree, to 40 m tall, to 70 cm diameter, with small thin buttresses and stilt roots.
Bark smooth; inner bark pale brown or pink. Twigs c. 1 mm diameter apically, slender,
glabrescent. Leaves thinly coriaceous, drying pale brown including venation; blade ovate,
5–8 × 2.2–4.5 cm, base cuneate, frequently subequal, margin not revolute, apex caudate,
acumen to 1.5 cm long; midrib slender, terete, elevated on both surfaces; venation
dryobalanoid, veins indistinct and hardly raised, drying not blackish; main lateral veins c. 8
pairs; intercostal venation obscure, reticulate; petiole 1.2–1.7 cm long, slender, somewhat
geniculate. Inflorescences axillary or rarely terminal, greyish caducous puberulent, to 4 cm
long, singly or doubly branched, branchlets bearing to 5 distichous flowers. Flowers: buds
small, ovoid; petals ovate, cream; stamens c. 15, anthers globose, connectival appendage as
long as anther; ovary ovoid, stylopodium obscure, style columnar, villous towards base, as
long as ovary. Fruits subsessile; calyx lobes unequal, 2 longer lobes spatulate, to 6 × 1.4
cm, narrowly obtuse, tapering to 3 mm above the saccate base, 3 shorter ones ovate, acute,
to 0.9 × 0.7 cm, hardly conceiling the nut. Nuts broadly ovoid, to 0.8 × 0.7 cm, with c. 2
mm long filiform style remnant.
162
Vernacular name. Sarawak—luis daun bulat (preferred name).
Distribution. Peninsular Malaysia and Borneo. In Borneo known in Sabah from Labuk
Sugut district (e.g., SAN 90497) and in Sarawak from Kuching, Serian and Tatau districts
(e.g., S 7614, S 9340, S 16631, S 44906, and S 64962). Also occurring in Brunei (e.g.,
BRUN 3179 and BRUN 3338) and E Kalimantan (e.g., Kostermans 6752).
Ecology. Locally common in mixed dipterocarp forest on leached sandy clay soils, on low
hills at altitudes to 400 m. Well represented in Bako NP but elsewhere vulnerable.
Notes. For differences from H. beccariana see there.
19. Hopea longirostrata P.S.Ashton

(Latin, longus = long, rostratus = beaked; the tapering fruit base and long pedicel)
Type: Ashton S 17742, Borneo, Sarawak, Belaga district, Ulu Belaga (holotype K; isotypes KEP, L).
Subcanopy tree, to 14 m tall; buttresses thin, low. Bark smooth, becoming patchily thinly
flaky; inner bark pale brown. All parts glabrous. Twigs c. 2 mm diameter apically. Leaves
coriaceous, drying greyish brown; blade ovateelliptic, 7–9 × 3–5 cm, base obtuse, margin
not revolute, apex subcaudate, acumen to 1.5 cm long; midrib slender but prominent above,
sharply elevated below, tending to dry black; venation dryobalanoid, unraised and more or
less obscure, arched; main lateral veins c. 12 pairs, with subequal intermediates; petiole
0.7–1 cm long, slender, geniculate, drying black. Flowers unknown. Infructescences
terminal or in axillary clusters of 3, to 4 cm long, singly branched, branchlets bearing to 5
secund fruits. Fruits: base tapering; pedicels to 7 mm long; calyx lobes unequal, 2 longer
lobes spatulate, to 2.4 × 0.6 cm, tapering to 3 mm and ending abruptly in a small incrassate
central tubercle, 3 shorter ones linear to spatulate, to 1.5 cm long, similar at base,
conceiling the nut. Nuts ovoid, to 0.6 × 0.4 cm, with to 2 mm long slender style remnant.
Distribution. Endemic in Borneo and confined to Ulu Tubau, Bintulu district and Ulu
Belaga, Belaga district in Sarawak (e.g., S 18191 and the type).
Ecology. Rare and endangered, in small groups in mixed dipterocarp forest on clayrich
soils, at c. 300 m altitude.
20. Hopea megacarpa P.S.Ashton

(Greek, mega = huge, karpos = fruit; the large nut)
Type: Daud & Tachun SFN 35625, Borneo, Sarawak, Kapit district, Nanga Pelagus (holotype K;
isotype KEP).
163
Subcanopy tree, to 15 m tall, to 20 cm diameter. Bark smooth; inner bark yellowish brown.
Young twig, petiole and venation below sparsely caducous puberulent; leaf bud persistently
so; parts otherwise glabrous. Twigs c. 1 mm diameter apically, slender, sparingly branched,
dark brown. Leaves thinly coriaceous, undulate, drying pale brown; blade narrowly ovate
lanceolate, 6–12 × 1.5–5 cm, base equal, cuneate, apex caudate, acumen to 2 cm long,
slender; midrib prominent and terete below, flat to somewhat raised and slender above;
venation hopeatype; lateral veins 6–7 pairs, slender, raised below, ascending and arched;
domatia not prominently swollen; intercostal venation densely scalariform, slender and
unraised; petiole c. 0.6 cm long, slender. Inflorescences axillary, lax, to 3 cm long, singly
branched, branchlets bearing to 3 dense flowers; bracts fugaceous. Flowers: buds
subglobose, to 4 × 3 mm; petals ellipticoblong, pale pink; stamens c. 15, anthers oblong,
connectival appendage c. 3x the length of anther; ovary and stylopodium cylindrical,
subtruncate, style short. Fruits: pedicels c. 3 mm long, broadening into receptacle; calyx
lobes unequal, 2 longer lobes narrowly spatulate, acute, to 10 × 1.3 cm, tapering to c. 5 mm
above the subauriculate saccate base, 3 shorter ones ovate, to 2 × 0.9 cm, similarly
subauriculate, conceiling the nut. Nuts ovoid, to 1.2 × 1 cm, with mucronate minutely
truncate apex.
Distribution. Endemic in Borneo. Known in Sarawak only from Kapit district (e.g., S 639,
S 14743, S 29085, S 29490, and S 41546). Also occurring in SE Kalimantan (e.g., Suzuki K
9754).
Ecology. Locally gregarious understorey trees of mixed dipterocarp forest on sandy clay
soils, at altitudes below 500 m. Vulnerable, probably endangered.
21. Hopea mengerawan Miq.

(Sumatran vernacular name)
Fl. Ned. Ind., Suppl. (1862) 491; Ridley op. cit. (1922) 238; Foxworthy op. cit. 137; Symington op.
cit. (1939) 361, op. cit. (1943) 132; Ashton op. cit. (1968) 53, op. cit. (1982) 400; Anderson op. cit.
(1980) 116; PROSEA op. cit. 249; Kessler & Sidiyasa op. cit. 98; Newman et al. op. cit. 151. Type:
Teijsmann s.n. (= U sub. No. 035930), Sumatra, Palembang, Muara Enim (holotype U; isotypes BO,
L). Synonym: Hancea mengerawan (Miq.) Pierre op. cit. t. 243.
Large main canopy or low emergent tree, to 45 m tall, to 1 m diameter; crown

hemispherical; buttresses to 5 cm thick, flying buttresses to 1.5 m tall; stilt roots sometimes
present. Bark becoming deeply narrowly vsection fissured, dark chocolatebrown and
greymottled. Sapwood honeyyellow, hard; heartwood dark brown. Twig, petiole and leaf
below caducous lepidote; parts of petal exposed in bud sericeous; parts otherwise glabrous.
Twigs c. 2 mm diameter apically. Leaves coriaceous, drying warm chocolatebrown below,
goldenbrown above; blade oblonglanceolate, 6–12 × 2.5–5 cm, base cuneate, apex with
tapering acumen to 1.5 cm long; midrib stout, prominent on both surfaces; venation
dryobalanoid, veins evident but hardly elevated below, drying not blackish; main lateral
veins c. 14 pairs, with many short to subequal intermediates; petiole 0.9–1.1 cm long, stout.
Inflorescences terminal or axillary, stout, to 3 cm long, singly branched, branchlets bearing
to 6 secund flowers. Flowers: buds ovoid, to 3 × 2 mm; petals oblonglanceolate, pale
yellow; stamens c. 15, anthers subglobose, connectival appendage 2–3x the length of anther;
ovary ovoid, stylopodium obscure, style columnar, villous in basal half, c. 2x the length of
ovary. Fruits: pedicels c. 2 mm long; calyx lobes unequal, 2 longer lobes narrowly
spatulate, narrowly obtuse, to 7 × 1.2 cm, tapering to c. 3 mm above the saccate base, 3
164
shorter ones ovate, acute, to 0.6 × 0.5 cm, appressed to but not conceiling the nut apex.
Nuts narrowly ovoid, to 1 × 0.5 cm, with prominent terminal filiform style remnant.
Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo. In Sabah recorded

from Lahad Datu, Sandakan and Tawau districts (e.g., SAN 16981, SAN 23267, SAN 32508,
and SAN 39302) and in Sarawak from Bintulu and Mukah districts (e.g., S 15577 and S
23247). Also occurring in W, E and S Kalimantan (e.g., bb. 24523, Kostermans 6657,
Kostermans 6745, Kostermans 12513, and Sidiyasa 605).
Ecology. In mixed dipterocarp forest near the base of low hills, on sandy soils often with
impeded drainage, at altitudes below 200 m. Inexplicably rare and critically endangered
throughout Borneo, with records based on few collections from isolated localities.
22. Hopea mesuoides P.S.Ashton

(Latin = resembling Mesua, Guttiferae)
Coode et al. (eds.) op. cit. 72. Type: Ilias S 15551, Borneo, Sarawak, Bintulu district, Segan FR
(holotype K; isotypes KEP, L). Synonym: Hopea subalata auct. non Symington: Ashton op. cit.
(1964) 110.
Subcanopy to low canopy tree, to 30 m tall, to 50 cm diameter, with persistently

monopodial diffuse crown and pendent branches; buttresses thin and low, flying buttresses
and stilt roots present. Bark smooth. Leaf bud, flower parts exposed in bud and stipule pale
yellowish brown puberulent, indumentum persistent except on calyx; other parts glabrous.
Twigs c. 1.5 cm diameter apically, slender, sparingly branched. Leaves coriaceous, drying
pale chocolatebrown below, yellowish brown above; blade ovatelanceolate, 8–14 × 2.5–5
cm, base obtuse, equal, margin more or less narrowly revolute, apex caudate, acumen to 1.5
cm long; midrib slender but prominent below, obscure and sunken above; venation
subdryobalanoid; main lateral veins c. 11 pairs, slender and distinctly raised below, arched,
with short intermediates; intercostal venation evident below, reticulate; petiole 0.7–1 cm
long, slender. Inflorescences axillary, slender, to 4 cm long, lax, singly branched; bracts c.
1 mm long, subpersistent. Flowers: buds small, ovoid; petals oblong, dark red; stamens c.
15, anthers subglobose, connectival appendage 2–3x the length of anther; ovary and
stylopodium cylindrical, truncate, papillose at apex, style short. Fruits: pedicels c. 1 mm
long; base impressed; calyx lobes subequal, ovate, chartaceous, erose near the subacute
apex, to 1.4 × 1.2 cm, appressed to the nut. Nuts subglobose, apex protruding above calyx,
crowned by persistent truncate stylopodium.
Distribution. Endemic in Borneo. Recorded in Sarawak from Bintulu, Limbang, Marudi,

Miri, and Tatau districts (e.g., S 11247, S 25058, S 27123, S 31839, and S 32314) and in
Brunei from Amo, Belait and Tutong districts (e.g., BRUN 879, BRUN 5674 and FMS
34451).
Ecology. In mixed dipterocarp forest on yellow sandy clay soil and kerangas forest on
podsols, on low hillslopes, at altitudes to 200 m. Locally abundant, forming dense groups
with abundant juveniles. Vulnerable owing to its accessable habitat.
165
23. Hopea micrantha Hook.f.

(Greek, mikro = small, anthos = a flower; with small flowers)
Trans. Linn. Soc. 23 (1860) 161; Merrill op. cit. (1921) 402; Symington op. cit. (1939) 355;
Masamune op. cit. 490; Browne op. cit. 121, p.p.; Ashton op. cit. (1964) 103, op. cit. (1968) 54, op.
Coode et al. (eds.) op. cit. 72. Lectotype (Symington, 1939): Motley 215, Borneo, Labuan
(hololectotype K). Synonym: Hancea micrantha (Hook.f.) Pierre op. cit. t. 243.
Canopy tree, to 30 m tall, to 30 cm diameter, rarely to 40 m tall, to 80 cm diameter;

buttresses small, narrow; stilt roots many. Bark smooth; inner bark pale brown or pink.
Young parts, leaf blade excepted, pale brown fugaceous puberulent. Twigs c. 1 mm
diameter apically, slender. Leaves thinly coriaceous, drying reddish brown below with the
midrib darker, greyish brown above; blade oblonglanceolate, 6–8 × 2.5–3 cm, base equal,
obtuse, apex caudate, acumen to 1.5 cm long; midrib evident, slightly raised above, terete
and prominent below, not drying black; venation dryobalanoid, hardly raised below; main
lateral veins c. 14 pairs, with shorter intermediates, arched; petiole 0.7–1 cm long, slender.
Inflorescences terminal or axillary, to 1 cm long, singly branched, branchlets bearing to 5
secund flowers. Flowers: buds small, ovoid; petals linear, pale pink; stamens c. 15, anthers
broadly oblong, connectival appendage c. 1½x the length of anther; ovary ovoid,
stylopodium obscure, style columnar, c. 1½x the length of ovary. Fruits subsessile; calyx
lobes unequal, 2 longer lobes spatulate, obtuse, to 5 × 1.2 cm, tapering to 2 mm above the
saccate base, 3 shorter ones broadly ovate to suborbicular, subacute or obtuse, to 0.5 × 0.5
cm, shorter than the nut. Nuts ovoid, to 1 × 0.6 cm, with slender style remnant to 1.5 mm
long.
Vernacular names. Sabah—selangan lunas (preferred name). Sarawak—luis kerangas

(preferred name).
Distribution. Endemic in Borneo. In Sabah known from Sipitang district (e.g., SAN 16264),
formerly also in Labuan (e.g., the type); and in Sarawak from Bintulu, Limbang, and Marudi
districts (e.g., S 951, S 1113, S 8267, and S 15893). Also occurring in Brunei (e.g., BRUN
512, BRUN 5526 and Kirkup DK 748).
Ecology. Very local, but there common, in kerangas forest on podsols, at altitudes below
100 m. Critically endangered.
24. Hopea montana Symington

(Latin, montanus = growing on mountains; the natural habitat)
J. Mal. Br. Roy. As. Soc. 19, 2 (1941) 141, op. cit. (1943) 133; Meijer & Wood op. cit. 214; Burgess
op. cit. 129, 136; Ashton op. cit. (1982) 413; PROSEA op. cit. 250. Type: Symington FMS 32257,
Peninsular Malaysia, Perak, G. Korbu (holotype KEP).
Subcanopy or canopy tree, to 35 m tall, to 1 m diameter; bole often crooked; buttresses thin,
flying buttresses and stilt roots present. Bark smooth or patchily cracked. Parts entirely
glabrous. Twigs c. 1 mm diameter apically, slender, blackish. Leaves thinly coriaceous,
drying pale chocolatebrown; blade ovatelanceolate, 6.5–9 × 2.2–4 cm, base equal,
abruptly cuneate, apex cuspidate, acumen to 1 cm long; midrib obscure, sunken above,
slender but raised below as also the veins; venation subdryobalanoid; main lateral veins c.
14 pairs, with many short intermediates; intercostal venation obscure; petiole 0.9–1.1 cm
166
long, slender. Inflorescences terminal or axillary, to 2 cm long, lax, slender, hardly

branched, with small deltoid subpersistent bracts. Flowers (mature) unknown. Fruits:
pedicels c. 2 mm long, expanding into receptacle; calyx lobes unequal, 2 longer lobes
spatulate, subacute, to 5 × 1.2 cm, tapering to 2 mm above the saccate thickened base, 3
shorter ones ovate to lanceolate, to 1 × 0.3 cm, conceiling the nut. Nuts ovoid, to 0.7 × 0.5
cm, with minutely truncate apex bearing a central apiculus.
Vernacular name. Sabah—selangan bukit (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo, recorded in Sabah from
Kota Belud, Sipitang and Tambunan districts (e.g., KEP 80317, SAN 16280, SAN 17002,
and SAN 40258).
Ecology. Rare but locally frequent in upper dipterocarp forest on hillslopes on clayrich
soils, at 900–1200 m altitude. Probably vulnerable as the habitat is mostly outside the
National Park boundary.
25. Hopea nervosa King

(Latin, nervosus = bearing nerves; the many prominent leaf lateral veins)
J. As. Soc. Beng. 62, 2 (1893) 124; Ridley op. cit. (1922) 236; Foxworthy op. cit. 129; Symington op.
cit. (1943) 185; Ashton op. cit. (1964) 104, op. cit. (1968) 54, op. cit. (1982) 410; Meijer & Wood op.
cit. 215; Burgess op. cit. 129; Anderson op. cit. (1980) 116; PROSEA op. cit. 250; Coode et al. (eds.)
op. cit. 72. Type: King’s Collector 3690, Peninsular Malaysia, Perak, Larut (holotype CAL; isotype
K).
Mediumsized subcanopy or low canopy tree, to 30 m tall, to 1 m diameter; buttresses thin,

flying buttresses to 2 m tall; stilt roots present. Bark smooth, greyish brown. Sapwood pale,
soft. Young parts including midrib throughout caducously greyish puberulent. Twigs c. 1
mm diameter apically, slender, sparsely branched. Leaves thinly coriaceous, drying warm
chocolatebrown below; blade narrowly ovate to lanceolate, 9–18 × 4–7 cm, base equal,
broadly cuneate, apex with tapering acumen to 1.4 cm long; midrib slender but prominent
and terete below, narrowly sunken above; venation hopeatype; lateral veins 13–15 pairs,
slender but prominent below; intercostal venation very slender, dense, sinuatescalariform,
unraised; petiole 1–1.2 cm long. Inflorescences terminal or in axillary pairs, lax, to 9 cm
long, singly branched, branchlets bearing to 5 remote flowers; bracts deltoid, subpersistent.
Flowers: buds subglobose, to 3 mm diameter; petals narrowly oblong, purplish with paler
tips; stamens c. 15, anthers subglobose, connectival appendage c. 2x the length of anther;
ovary and stylopodium broadly cylindrical, truncate, style short. Fruits subsessile; calyx
lobes unequal, 2 longer lobes spatulate, to 12 × 1.8 cm, narrowly obtuse, thinly coriaceous,
strongly twisted, tapering to 3 mm above the saccate base, 3 shorter ones lanceolate, acute,
to 1.5 cm long, more or less conceiling the nut. Nuts broadly ovoid, to 1 × 1 cm, with
apiculate minutely truncate apex.
Vernacular names. Sabah—selangan jangkang (preferred name). Sarawak—luis jangkang

(preferred name).
Distribution. Peninsular Malaysia and Borneo. In Borneo recorded in Sabah from

Kinabatangan and Sandakan districts (e.g., FMS 35404, SAN 23305, SAN 34254, SAN
38954, and SAN 39016) and in Sarawak from Belaga, Kapit, Lundu, and Tatau districts
167
BRUN 790, FMS 35671 and Wong WKM 1922) and E Kalimantan (e.g., Kostermans 4109,
Kostermans 4342, Kostermans 4394, and Kostermans 13981). Particularly common in W
Sarawak and E Sabah, elsewhere scattered.
Ecology. In mixed dipterocarp forest, locally common on wellstructured deep clay soils on
slopes and low hills on intermediate and basic igneous rocks, more scattered on
sedimentaries, at altitudes to 400 m. Occurring in G. Gading and Mulu NPs; elsewhere
vulnerable owing to forest conversion.
26. Hopea nutans Ridl.

(Latin, nutans = nodding; the flowers)
FMP 1 (1922) 235; Foxworthy op. cit. 123; Symington op. cit. (1943) 136; Browne op. cit. 125;
Ashton op. cit. (1964) 104, op. cit. (1968) 54, op. cit. (1982) 424; Meijer & Wood op. cit. 217;
72; Newman et al. op. cit. 153. Lectotype (designated here): Henbrey KEP 417, Peninsular Malaysia,
Pahang, Kuantan (hololectotype K; isolectotype KEP).
Canopy tree, to 40 m tall, to 1.2 m diameter; crown diffuseirregular; bole often misshapen;
buttresses low. Bark greyish brown, becoming coarsely irregularly flaky; dammar
exudations not conspicuous. Sapwood yellowish brown; heartwood warm brown, hard.
Young parts, leaf blade excepted, sparsely pale brown puberulent; leaf blade below more or
less sparsely greyish lepidote. Twigs to 2 mm diameter apically, relatively stout, much
branched. Leaves thickly coriaceous, drying greyish; blade broadly ovate, 8–13 × 4.5–8.5
cm, base obtuse, subequal, margin more or less revolute, apex broadly tapering, with
acumen to 1 cm long; midrib stout, somewhat raised on both surfaces; venation hopeatype;
lateral veins 7–10 pairs, relatively stout, somewhat raised below, arched, usually with a few
large glabrous swollen porelike domatia; intercostal venation slender but distinct,
subscalariform; petiole 1–1.5 cm long, fairly stout. Inflorescences terminal or axillary, to 7
cm long, branchlets bearing to 5 secund dense flowers; bracts fugaceous. Flowers: buds
broadly ellipsoid, to 4 × 2 mm; petals linear, cream; stamens c. 15, anthers subglobose,
connectival appendage c. 2x the length of anther, scabrous towards apex; ovary and
stylopodium subcylindrical, style short. Fruits: subsessile; calyx lobes unequal, 2 longer
lobes oblong, broad, obtuse, thinly coriaceous, to 8 × 1.5 cm, tapering to 2.5 cm above the
saccate base, 3 shorter ones ovate, acute, to 1 cm long. Nuts ovoid to 1.5 × 0.8 cm, acute.
Vernacular name. Sabah and Sarawak—giam (preferred name).
Distribution. Peninsular Malaysia and Borneo. In Sabah known from Lahad Datu and
Semporna districts (e.g., FMS 44588 and SAN 26123) and in Sarawak from Kuching and
Brunei (e.g., S 1668, S 1953 and S 5801) and NE Kalimantan.
Ecology. Local, uncommon in coastal hill mixed dipterocarp forest on sandy soils,
frequently with impeded drainage, ultrabasic soil and limestone outcrops. Occurring in Bako
and Mulu NPs but endangered elsewhere.
168
27. Hopea obscurinerva P.S.Ashton

(Latin, obscurus = indistinct, nervus = nerve; the indistinct venation on the leaf
undersurface)
TFSS 5 (2004) 479. Type: Mohtar S 51417, Borneo, Sarawak, Bintulu district, Ulu Tubau (holotype
KEP; isotype SAR).
Small subcanopy tree, to 10 m tall, c. 15 cm diameter, with low thin buttresses. Bark
smooth. All parts glabrous. Twigs c. 2 mm diameter apically, sparsely branched, warm
brown, somewhat shiny. Stipules linear, to 3 mm long, not at first caducous. Leaves thickly
coriaceous, not bullate, drying warm pinkish brown below, yellowish brown and somewhat
shiny above; blade oblonglanceolate, 15–26 × 5–8 cm, base markedly unequal, obtuse to
subcordate, apex with slender acumen to 1.5 cm long; midrib prominent above, less so
below; venation hopeatype; lateral veins 11–14 pairs, with a few intermediates, ascending,
arched, prominently raised above, hardly elevated below; intercostal venation obscure,
densely scalariform; petiole 0.4–0.6 cm long, c. 0.3 cm diameter, stout, drying black.
Inflorescences and flowers unknown. Fruits (young): pedicels 1–2 mm long; base of fruit
impressed; calyx lobes unequal, 2 longer lobes narrowly spatulate, at least 4 × 0.8 cm, not
auriculate, 3 shorter ones narrowly deltoid, to 1.2 × 0.5 cm. Nuts ovoid, glabrous,
surmounted by a c. 3 mm long, spindleshaped stylopodium remnant.
Distribution. Endemic in Borneo and known to date only from the lower Tubau drainage in
Sarawak (e.g., S 18195 and the type).
Ecology. In mixed dipterocarp forest on clay soil, at c. 300 m altitude. Endangered.
28. Hopea ovoidea P.S.Ashton

(Latin, ovoideus = eggshaped; the ovary)
Gard. Bull. Sing. 31 (1978) 34, op. cit. (1982) 426. Type: Elmer 21428, Borneo, Sabah, Tawau
district (holotype K; isotype L).
Canopy tree, to 1 m diameter; buttresses prominent, thin. Bark flaky, brown. Sapwood
hard. Leaf bud, inflorescence, and parts of flower exposed in bud densely persistently pale
buffpubescent; parts otherwise glabrescent. Twigs c. 1 mm diameter apically, slender.
Leaves chartaceous and undulate, drying pale greyish brown; blade elliptic to narrowly
ovate, 9–13 × 3–6.5 cm, base unequal or subequal, cuneate, shortly decurrent, apex with
slender, tapering acumen to 2 cm long; midrib slender but distinctly raised on both surfaces;
venation hopeatype; lateral veins 7–8(–10) pairs, slender but distinctly elevated below,
arched, often with a few glabrous porelike canaliculate domatia; intercostal venation
scalariform, distinctly elevated below; petiole 1–1.5 cm long, slender. Inflorescences in
axillary pairs or terminal, pendent, to 13 cm long, singly branched, lax, branchlets bearing
to 7 secund flowers; bracts fugaceous. Flowers: buds ovoid, to 3 × 2 mm; petals lanceolate,
cream; stamens c. 15, anthers narrowly elliptic, connectival appendage 1½–2x the length of
anther; ovary and stylopodium ovoid, tomentose, style c. ½x the length of ovary, glabrous.
Fruits unknown.
Distribution. Endemic in Borneo; known only from Lahad Datu, Sandakan and Tawau
districts in Sabah (e.g., SAN 21690, SAN A 2489 and SAN A 4829).
169
Ecology. Rare and endangered (possibly extinct), in mixed dipterocarp forest on low coastal
hills.
29. Hopea pachycarpa (F.Heim) Symington Fig. 15.

(Greek, pachy = thick, karpos = fruit; the large and thick fruit)
Gard. Bull. S. S. 8 (1934) 30; Masamune op. cit. 491; Browne op. cit. 125; Ashton op. cit. (1964) 105,
op. cit. (1967) 271, op. cit. (1968) 54, op. cit. (1978) 35, op. cit. (1982) 432; Anderson op. cit. (1980)
117; Coode et al. (eds.) op. cit. 72. Basionym: Pierrea pachycarpa F.Heim op. cit. (1891) 958. Type:
Beccari PB 3314, Borneo, Sarawak, Marop (holotype P). Synonyms: Balanocarpus pubescens Ridl.
op. cit. (1922) 247; Hopea laxa Symington op. cit. (1934) 33; H. resinosa Symington, Gard. Bull. S.
S. 8 (1935) 278.
Subcanopy tree, to 40 m tall, to 80 cm diameter, with pendent leaves and twigs; crown dark,
irregular; buttresses low, thin; stilt roots present. Bark smooth. Young twig, leaf bud,
stipules outside, foliar domatia, and petiole densely persistently pale fawnpubescent or
whitishpruinose or silvery lepidote; inflorescence fugaceously so; floral sepals fimbriate;
parts otherwise glabrous. Twigs c. 2 mm apically, sparingly branched. Leaves thinly
coriaceous, drying greyish green, more or less silvery lepidote below; blade flat, elliptic to
lanceolate, 13–22 × 4–7 cm, base unequal, cuneate to obtuse, margin often narrowly
revolute, apex with a slender acumen to 1 cm long; midrib stout, terete, raised on both
surfaces; venation hopeatype; lateral veins (10–)13–17 pairs, slender but raised below, flat
above, arched and running concurrently up margin, with small tomentose axillary domatia;
intercostal venation slender, densely scalariform, sinuate; petiole 0.7–1 cm long, stout.
Inflorescences in axillary pairs or ramiflorous, rarely branched. Flowers: buds broadly
ellipsoid, to 4 × 2.5 mm; petals cream; stamens c. 15, anthers subglobose, connectival
appendage 2–3x the length of anther; ovary small, ovoid, style and stylopodium spindle
shaped, glandularpapillose towards tapering apex. Fruits large, impressed at base; calyx
lobes subequal, ovate, subacute, to 2 × 1.5 cm, thickened, saccate, appressed to the nut.
Nuts ovoidglobose, to 1.5 cm diameter, acute, conceiled in calyx except for the apex,
usually resincoated.
Vernacular name. Sarawak—merkoyong (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo recorded in Sabah from
Keningau and Pensiangan districts (e.g., SAN 128204, SAN 136792, SAN 139662, and SAN
139667) and in Sarawak from Belaga, Bintulu, Kapit, Kuching, Lawas, Lundu, Miri, Serian,
Simunjan, Sri Aman, and Tatau districts (e.g., S 13572, S 15408, S 22406, S 36881, S
5647) and Kalimantan (e.g., Burley & Tukirin 762, Hansen 1374 and Kessler et al. Berau
19).
Ecology. Locally abundant in mixed dipterocarp forest on moist lower slopes and
streamsides, on clayrich soils, on both sedimentary and intermediate and basic igneous
rocks, at altitudes below 400 m. Occurring in G. Gading and Mulu NPs but vulnerable
elsewhere.
30. Hopea pedicellata (Brandis) Symington

(Latin, pedicellatus = with prominent stalk; the flowers)
170
Fig. 15. Hopea pachycarpa. A, flowering leafy twig; B, detail of indumentum on lower leaf
surface; C, detail of indumentum on upper leaf surface; D, open flower; E, longitudinal
section of opening flower; F, adaxial view of outer sepal; G, adaxial view of inner sepal; H,
adaxial view petal with stamens; I, abaxial view of petal; J, adaxial view of stamens; K,
abaxial view of stamens; L, fruit; M, longitudinal section of fruit. (A–K from S 21300, L–M
from S 36881.)
171
Gard. Bull. S. S. 9 (1938) 327, op. cit. (1943) 138; Ashton op. cit. (1968) 54, op. cit. (1978) 30, op. cit.
(1982) 408; Anderson op. cit. (1980) 117; PROSEA op. cit. 252; Coode et al. (eds.) op. cit. 72.
Basionym: Hopea griffithii Kurz var. pedicellata Brandis, J. Linn. Soc. Bot. 31 (1895) 69, p.p.
Lectotype (Ashton, 1978): Curtis 167, Peninsular Malaysia, Penang (hololectotype K). Synonym:
Hopea siamensis F.Heim, Bot. Tidsskr. 25 (1902) 46.
Small canopy tree, to 40 m tall, to 60 cm diameter; bole often crooked; buttresses low, thin;
stilt roots present. Bark dark brown, smooth, hoopmarked or sometimes patchily cracked;
inner bark pinkish brown; dammar exudations glassy. Young twig and inflorescence
caducously, petiole, leaf bud, stipules and domatia persistently greyish brown puberulent.
Twigs c. 1 mm diameter apically, slender, muchbranched. Leaves thinly coriaceous to
coriaceous; blade flat, ovatelanceolate, 4–9 × 1–3.5 cm, base cuneate, margin flat, apex
subcaudate, acumen to 1.5 cm long; midrib slender but raised on both surfaces or sharp
below and tending to dry black; venation dryobalanoid, hardly elevated, indistinct; main
lateral veins c. 8–12(–15) pairs, not drying black, with many subequal intermediates,
obscure below; petiole 0.6–0.8 cm long. Inflorescences terminal or axillary, to 4 cm long,
singly branched, branchlets bearing to 7 secund flowers. Flowers: buds ovoid, to 2.5 × 2
mm; petals oblonglanceolate, pale yellow; stamens c. 15, anthers oblong, connectival
appendage c. 2x the length of anther; ovary and stylopodium cylindricconical, attenuate,
truncate, punctate in the distal half, style short. Fruits: pedicels 2 mm long; calyx lobes
unequal, 2 longer lobes spatulate, to 3 × 0.5 cm, tapering to 2 mm above the saccate base, 3
shorter ones ovate, to 0.3 × 0.3 cm, appressed to the nut. Nuts ovoid, to 0.6 × 0.4 cm,
abruptly acute, exceeding calyx.
Vernacular name. Sarawak—mata kucing bukit (preferred name).
Distribution. Peninsular Thailand, Peninsular Malaysia and Borneo. In Sabah recorded

from Kota Belud district (e.g., SAN 76269, young buds, tentative) and in Sarawak from
Belaga, Bintulu, Miri, Serian, Simunjan, and Tatau districts (e.g., S 15213, S 17023, S
18351, S 25017, and S 64525). Also occurring in Nunukan Is. in Kalimantan.
Ecology. Uncommon and scattered in distant localities in mixed dipterocarp forest on

undulating land and ridges, at altitudes to 600 m. Vulnerable.
31. Hopea pentanervia Symington ex Wood

(Greek, penta = five, nervius = nerved; the leaf blade)
Gard. Bull. Sing. 17 (1960) 495; Browne op. cit. 126, Anderson, Gard. Bull. Sing. 20 (1963) 157, op.
cit. (1980) 117; Ashton op. cit. (1964) 106, op. cit. (1968) 55, op. cit. (1982) 425; Meijer & Wood op.
cit. 219, Burgess op. cit. 135; PROSEA op. cit. 263, Coode et al. (eds.) op. cit. 72, Newman et al. op.
cit. 154. Type: Tready S 1257, Borneo, Sarawak, Miri district, Baram (holotype KEP; isotype SAR).
Canopy tree, to 35 m tall, to 1 m diameter; crown irregular, with large twisted branches;
buttresses to 1 m tall, c. 4 cm thick. Bark vertically cracked and oblongflaked, warm dark
brown and greymottled. Sapwood straw yellow, hard; heartwood chocolatebrown. Young
parts puberulent; all parts glabrescent but for the fimbriate flower sepals and pubescent
parts of petals exposed in bud. Leaves coriaceous, drying yellowish brown; blade ovate, 5–
10 × 3.2–5 cm, base equal, obtuse or broadly cuneate, margin somewhat revolute, apex
with slender, tapering acumen to 1.5 cm long; midrib slender, raised below, flat above;
venation hopeatype; lateral veins c. 5 pairs, slender but prominent below, arched, with
small glabrous porelike domatia; intercostal venation densely scalariform, evident; petiole
172
0.6–1.1 cm long. Inflorescences terminal or in axillary pairs, lax, to 8 cm long, singly

branched, branchlets bearing to 6 dense secund flowers; bracts fugaceous. Flowers: buds
ellipsoid, c. 1.5 mm long; petals lanceolate, cream; stamens c. 15, anthers subglobose,
connectival appendage c. 3x the length of anther; ovary and stylopodium cylindrical,
truncate, style short. Fruits subsessile; calyx lobes unequal, 2 longer lobes spatulate,
obtuse, chartaceous, to 5 × 1.2 cm, tapering to 3 mm above the saccate base, 3 shorter ones
ovate, acute, appressed to the nut. Nuts ovoid, to 0.4 × 0.35 cm, acute.
Vernacular names. Sabah—selangan lima urat (preferred name). Sarawak—cengal paya

(preferred name).
Distribution. Endemic in Borneo, occurring throughout SW and E Sabah and recorded

from Beaufort, Keningau, Kinabatangan, Labuk Sugut, Papar, Sandakan, and Sipitang
districts (e.g., SAN 25467, SAN 38766, SAN 96930, SAN 130223, and SAN 138822) and in
Sarawak from Betong, Bintulu, Kuching, Lawas, Limbang, Marudi, Miri, and Sibu districts
(e.g., S 260, S 1530, S 18387, S 23288, and S 24651). Also occurring in Brunei (e.g., BRUN
5007, Coode 7649 and FMS 35467).
Ecology. Locally frequent on shallow peat, usually over sand, in mixed peat swamp forest
and the ecotone of kerangas, and in kerangas on podsols on terraces and low hills, at
altitudes to 400 m; on ultrabasic rocks in E Sabah; usually near the coast. Occurring in Mulu
NP, elsewhere endangered.
32. Hopea plagata (Blanco) Vidal

(Latin, plagata, wounded; the woundlike domatia)
Synopsis (1883) t.15A; Merrill, Spec. Blancoan. (1918) 2691; Ashton op. cit. (1978) 32, op. cit.
(1982) 423, PROSEA op. cit. 264; Newman et al. op. cit. 155. Basionym: Mocanera plagata Blanco,
Fl. Filip. ed. 1 (1837) 447. Neotype (designated here): Merrill Sp. Blancoan. 109 (= US 903784), the
Philippines, Luzon, Rizal Province, Langhaya (K, US). Synonyms: Dipterocarpus plagatus (Blanco)
Blanco, Fl. Filip. ed. 2 (1845) 311; Anisoptera plagata (Blanco) Blume, Mus. Bot. Lugd. Bat. 2 (1852)
42; Hopea dasyrrhachis auct. non Slooten: Ashton op. cit. (1968) 49.
Large canopy tree, to 35 m tall, to 1 m diameter; crown irregular; buttresses to c. 1 m tall,

thin. Bark becoming dark greyish brown flaky. Sapwood hard, yellow; heartwood brown.
Parts entirely glabrous. Twigs c. 1 mm diameter apically, muchbranched, dark brown.
Leaves coriaceous, sometimes sparsely stellatelepidote below, drying dark greyish brown;
blade ellipticlanceolate to ovatefalcate, 6–12 × 2.5–7 cm, base markedly unequal, cuneate
to obtuse, apex with tapering acumen to 1.5 cm long; midrib stoutly prominent on both
surfaces; venation hopeatype; lateral veins 8–11(–12) pairs, slender but elevated below,
usually with glabrous porelike domatia; intercostal venation densely scalariform, hardly
elevated; petiole 0.6–1.6 cm long. Inflorescences solitary, axillary or terminal, to 3 cm
long, singly branched, branches bearing to 6 dense secund flowers; bracts fugaceous.
Flowers: buds ellipsoid, to 3 × 2 mm; petals lanceolate, cream; stamens c. 35, anthers
elongate, tapering, connectival appendage equal in length to anther, acicular; ovary ovoid,
stylopodium obscure, style short, broad. Fruits: pedicels c. 2 mm long; calyx lobes unequal,
2 longer lobes broadly oblongspatulate to suborbicular, to 4.5 × 2 cm, tapering to 3 mm
above the saccate base, 3 shorter ones ovate, to 0.7 × 0.4 cm, shorter than the nut. Nuts
narrowly ovoid, to 1 × 0.7 cm, acute.
173
Distribution. Borneo and the Philippines. In Sabah recorded from Kinabatangan, Kota
Kinabalu, Lahad Datu, Sandakan, Semporna, Tawau, and Tenom districts (e.g., KEP 80469,
SAN 9414, SAN 15042, SAN 20178, and SAN 43093) and in Sarawak from Marudi district
(e.g., BRUN 3203, S 22568, S 24037 and S 30767).
Ecology. Scattered in mixed dipterocarp forest, apparantly mostly on coastal hills in Sabah;
also very local on humic soils over limestone, at low altitudes and in lower montane forest
at 900–1100 m altitude. Well represented in Mulu NP but vulnerable in Sabah.
33. Hopea pterygota P.S.Ashton Fig. 16.

(Greek, pterygotos = winged; the fruit)
Coode et al. (eds.) op. cit. 73. Type: Smythies S 15206, Borneo, Sarawak, Simunjan district, G.
Gaharu (holotype K; isotypes KEP, L).
Small monopodial hardly buttressed tree, to 10 m tall, to 10 cm diameter, with pendent

twigs and leaves. Bark smooth, soft, whitish. Young vegetative parts densely pale tawny
pubescent; indumentum caducous on leaf below, petiole and twig, persistent on buds,
stipules and also parts of petal exposed in bud; reproductive parts otherwise glabrous.
Twigs c. 2 mm diameter apically, sparingly branched. Leaves thinly coriaceous, drying
reddish brown below; blade flat, oblonglanceolate to oblanceolate, 12–28 × 5–9 cm, base
markedly unequal, obtuse to subcordate, apex subcaudate, acumen to 2 cm long; midrib
prominent on both surfaces, more so below; venation hopeatype; lateral veins 12–21 pairs,
slender, prominent below, sunken above, dense; intercostal venation scalariform, not dense,
slightly elevated; petiole 0.3–0.8 cm long, stout. Inflorescences in axillary clusters of 4 or
ramiflorous, slender, lax, singly branched, branches bearing to 8 secund flowers. Flowers:
buds ellipsoid, to 3 × 2 mm; petals linear; stamens c. 15, anthers subglobose, connectival
appendage very slender, 3–4x the length of anther; ovary narrowly ovoid, stylopodium and
style spindleshaped, somewhat shorter than ovary. Fruits: pedicels c. 1 mm long; calyx
lobes unequal, 2 longer lobes spatulate, to 10 × 1.5 cm, chartaceous, tapering to c. 4 mm
then expanding to 7 mm wide paired auricles at the saccate base, 3 shorter ones lanceolate
deltoid, acute, to 1.2 cm long, similarly auriculate. Nuts ovoid, to 0.7 × 0.5 cm, acute,
conceiled within auricles.
Distribution. Endemic in Borneo; found throughout Sarawak and recorded from Belaga,
Kapit, Marudi, Miri, Mukah, Serian, Sibu, Song, Sri Aman, and Tatau districts (e.g., S
23612, S 25022, S 29630, S 36567, S 57666, and S 64844). Also occurring in Brunei.
Ecology. Locally gregarious, on a wide range of soils over acid and basic volcanic rocks,
shale, and also white sand podsol terrace; on welldrained sites, in mixed dipterocarp forest,
kerangas, and on ridges in the lower levels of upper dipterocarp forest, at 700–800 m
altitude. Locally abundant in Lambir and Mulu NPs; not yet vulnerable.
174
Fig. 16. Hopea pterygota. A, fruiting leafy twig; B, open flower; C, longitudinal section of
open flower; D, adaxial view of outer sepal; E, adaxial view of inner sepal; F, adaxial view
of petal with stamens; G, abaxial view of petal; H, adaxial view of stamens; I, abaxial view
of stamens; J, fruit; K, exposed nut. (A, J and K from S 29630, B–I from S 66783.)
175
34. Hopea rudiformis P.S.Ashton Fig. 17.

(Latin, rudis = the gladiatorial sword, formis = shape; the leaf shape)
Gard. Bull. Sing. 31 (1978) 30, op. cit. (1982) 409; Kessler & Sidiyasa op. cit. 99. Type: Kostermans
4394, Borneo, Kalimantan, Balikpapan, Sg. Wain (holotype L; isotypes BO, K).
Subcanopy to low canopy tree, to 35 m tall, to 40 cm diameter; crown more or less

persistently monopodial; buttresses to 1 m tall, thin; flying buttresses and stilt roots present.
Bark greyish, smooth. Sapwood pale, soft. Fleshy exposed parts pale tawny puberulent;
indumentum persistent on young twigs, buds and petals; sparse on inflorescence; caducous
on leaf venation below and calyx. Twigs c. 2 mm diameter apically, ribbed, sparingly
branched. Leaves thinly coriaceous, more or less sparsely greyish stellatelepidote below
thereby appearing dull; blade ovate to broadly lanceolate, 6.5–14 × 3.5–7.5 cm, base equal,
broadly cuneate, margin narrowly revolute, apex tapering, downcurved, acumen to 1.5 cm
long; midrib and veins obscure and sunken above, slender but prominent below; venation
hopeatype; lateral veins 11–13 pairs, arched; intercostal venation densely scalariform,
obscure; petiole 0.8–1.3 cm long. Inflorescences in axillary pairs, to 3.5 cm long, singly
branched, branchlets bearing to 3 remote flowers; bracts subpersistent. Flowers: buds
ovoid, to 3 × 2 mm; petals oblong, deep purplish red; stamens c. 15, anthers broadly oblong,
connectival appendage slightly longer than anther; ovary ovoid, surmounted by indistinct
tapering stylopodium and short columnar style. Fruits: pedicels to 2 mm long; calyx lobes
unequal, 2 longer lobes broadly spatulate, to 9 × 2 cm obtuse, tapering to 3 mm broad
above the saccate base, 3 shorter ones ovate, to 0.8 × 0.8 cm, subacute, appressed to nut and
enclosing it. Nuts ovoidglobose, to 0.8 × 0.8 cm, terminating in a minutely truncate
stylopodium remnant.
Vernacular name. Sabah—selangan jangkang (Dusun).
Distribution. Endemic in Borneo. In Sabah, recorded from Kinabatangan, Labuk Sugut,

Lahad Datu, Sandakan, Tenom, and Tawau districts (e.g., SAN 18563, SAN 25471, SAN
29672, SAN 53121, and SAN 124401). Also known in E Kalimantan (e.g., bb. 25152,
Kostermans 4109, Kostermans 4549, and Kostermans 13981).
Ecology. In lowland forest on undulating land on clayrich soil, sometimes in freshwater

swamps, at altitudes to 500 m; locally common. Vulnerable owing to forest conversion.
35. Hopea rugifolia P.S.Ashton

(Latin, rugosus = wrinkled, folius = leaf; the wrinkled dried leaf)
Gard. Bull. Sing. 54, 2 (2002) 213. Type: Wong WKM 1414, Borneo, Brunei, Temburong district, Bt.
Belalong south ridge (holotype K; isotype BRUN). Synonym: Hopea sphaerocarpa auct. non
(F.Heim) P.S.Ashton: Coode et al. (eds.) op. cit. 73.
Small monopodial understorey tree, to 20 m tall, to 20 cm diameter, with low buttresses and
stilt roots. Bark smooth. Young parts densely evenly minutely greyish brown puberulent;
indumentum persisting sparsely on leaf venation below, midrib above at base, and petiole;
elsewhere glabrescent. Leaves chartaceous, drying greyish brown and wrinkled; blade
broadly lanceolate, 5–9 × 2–3.5 cm, base broadly cuneate, apex with slender acumen to 1.5
cm long; midrib slender but elevated below, obscure and sunken above; venation
subdryobalanoid; main lateral veins c. 10 pairs, slender but sharply raised below; intercostal
176
Fig. 17. Hopea rudiformis. A, flowering leafy twig; B, detail of indumentum on lower leaf
surface; C, detail of indumentum on upper leaf surface; D, flower bud; E, gynoecium; F,
abaxial view of petal; G, adaxial view of petal with stamens; H, adaxial view of stamens; I,
abaxial view of stamens; J, fruit; K, exposed nut. (A–I from FMS 38775, J–K from S
73781.)
177
venation densely scalariform, evident below; petiole to 0.4 cm long, slender. Inflorescences
in axillary pairs, slender, lax, to 4 cm long, singly branched, branchlets bearing to 6 remote
distichous flowers; bracts deltoid, to 1 mm long, subpersistent. Flowers: buds broadly
ovoid, to 2 × 1.5 mm; petals oblong, purple; stamens c. 15, anther ellipsoid, connectival
appendage c. 2x the length of anther; ovary and stylopodium narrowly cylindrical, truncate,
medially constricted, style short conical. Fruits: pedicels to 5 mm long, slender; calyx lobes
subequal, ovate, to 0.8 × 0.7 cm, acute, appressed to nut. Nuts broadly ovoid, to 1 × 0.8 cm,
with to 2 mm prominent stylopodium remnant.
Distribution. Endemic in Borneo; so far known only from Miri (Lambir NP) and Tatau (Bt.
Mersing) districts in Sarawak (e.g., S 25028 and S 46439), and Temburong district in Brunei
(e.g., the type).
Ecology. In mixed dipterocarp and upper dipterocarp forest on friable fertile clay soils, at
100–800 m altitude. Well represented in Lambir NP but vulnerable elsewhere.
36. Hopea sangal Korth.

(from Dayak name in Kalimantan)
Kruidk. (1841) 75; Symington op. cit. (1934) 18, op. cit. (1943) 141; Keith op. cit. 41; Masamune op.
cit. 491; Browne op. cit. 121; Ashton op. cit. (1963) 260, op. cit. (1964) 108, op. cit. (1968) 56, op. cit.
(1982) 420, Meijer & Wood op. cit. 222; Burgess op. cit. 129; Anderson op. cit. (1980) 117; PROSEA
op. cit. 253; Coode et al. (eds.) op. cit. 73; Newman et al. op. cit. 156. Type: Korthals s.n. (= RHL
Sheet No. 933193), Borneo (holotype L; isotypes BO, U). Synonyms: Dryobalanops sericea Korth.
op. cit. 72; Hopea sericea (Korth.) Blume op. cit. (1852) 35; Petalandra micrantha Hassk., Hort. Bog.
Desc. (1852) 105; H. fagifolia Miq. op. cit. 490; Doona micrantha (Hassk.) Burck op. cit. 233; Doona
javanica Burck op. cit. 235; H. hasskarliana F.Heim, Rech. Dipt. (1892) 64; H. javanica (Burck)
F.Heim op. cit. (1892) 64; H. curtisii King op. cit. 124; H. globosa Brandis op. cit. 61; H. lowii Dyer
ex Brandis op. cit. 63; H. minutiflora C.E.C. Fischer, Bull. Misc. Inform. Kew (1927) 207.
Mediumsized canopy tree, to 40 m tall, to 1.3 m diameter; bole often misshapen; buttresses
low, thin. Bark vertically cracked and scaly, becoming conspicuous dark brown with
contrasting creamwhite opaque dammar coxcombs. Sapwood pale, soft. Inflorescence,
parts of perianth exposed in bud and domatia densely whitish buffpuberulent; indumentum
persistent on inflorescence and domatia; parts otherwise glabrous. Twigs c. 1 mm diameter
apically, slender, muchbranched. Leaves chartaceous, drying wrinkled, dull sooty greyish
brown; blade ovate, 5.5–10 × 3.5–5 cm, base equal, broadly cuneate, apex caudate, acumen
to 1.2 cm long; midrib flat, evident above, slender and prominent below; venation hopea
type; lateral veins 10–12 pairs, slender but prominent below, often with small pale
pubescent domatia; intercostal venation densely scalariform, indistinct; petiole 0.5–1 cm
long, slender. Inflorescences terminal or axillary, lax, to 7 cm long, singly or doubly
branched, branchlets bearing to 7 dense secund flowers; bracts fugaceous. Flowers: buds
ellipsoidovoid, to 2 × 1.5 mm; petals oblonglanceolate, cream; stamens c. 10, anthers
oblong, connectival appendage as long as anther; ovary and stylopodium shortly broadly
cylindrical, truncate, more or less puberulent, style c. 2/3x the length of ovary and
stylopodium. Fruits: pedicels c. 1 mm long; calyx lobes unequal, 2 longer lobes oblong
spatulate, obtuse, to 7 × 1.5 cm, tapering to 4 mm above the saccate base, 3 shorter ones
ovate, obtuse, to 0.5 × 0.4 cm, shorter than the nut. Nuts ovoid, to 0.7 × 0.4 cm, with
minutely subtruncate apiculate apex, often sparsely puberulent.
Vernacular name. Sabah and Sarawak—gagil (preferred name).
178
Distribution. Myanmar, Peninsular Thailand, Sumatra, Peninsular Malaysia, Java, Bali, and
Borneo. Widespread throughout Sabah and recorded from Kinabatangan, Kudat (including
Banggi Is.), Labuk Sugut, Lahad Datu, Ranau, Sandakan, Semporna, Tawau, and Tenom
Sarawak from Bau, Belaga, Bintulu, Kuching, Lundu and Tatau districts (e.g., S 922, S
13190, S 18440, S 43527, and S 49872). Also occurring in Brunei (e.g., BRUN 3378, FMS
30544 and FMS 39619) and Kalimantan (e.g., bb. 34337 and Endert 2117).
Ecology. Scattered in mixed dipterocarp forest on friable fertile clayrich soils, on

intermediate and basic igneous rocks and also shales and clays, at altitudes to 500 m.
Vulnerable though occurring in G. Gading and Mulu NPs.
37. Hopea semicuneata Symington

(Latin, semi = half, cuneatus = wedgeshaped; the leaf base)
Gard. Bull. S. S. 8 (1934) 24, op. cit. (1943) 143; Masamune op. cit. 491; Meijer & Wood op. cit. 224;
PROSEA op. cit. 264; Newman et al. op. cit. 157. Type: Awang FMS 4526, Peninsular Malaysia,
Pahang, Temerloh (holotype KEP). Synonym: Hopea diversifolia Miq., op. cit. 451, p.p.
Large canopy or low emergent tree, to 50 m tall, to 2 m diameter; bole tall, frequently
misshapen; crown diffuse, small, hemispherical; buttresses to 7 m tall, 5 m long, c. 10 cm
thick. Bark becoming vertically cracked and oblongflaked, fawnbrown, with opaque white
dammar coxcombs; inner bark pinkish brown. Sapwood straw yellow, hard; heartwood dark
brown. Inflorescences and parts of flower exposed in bud densely pale buffpubescent;
indumentum caducous on calyx, otherwise persistent; young twigs and petioles fugaceous
puberulent; parts otherwise glabrous. Twigs c. 1 mm diameter apically. Leaves
chartaceous, frequently undulate, drying greyish brown below, purplish brown above,
curling up; blade elliptic to ovatelanceolate, 6.5–14 × 2–7 cm, base narrowly or broadly
cuneate, subequal, apex tapering, acumen slender, to 2.5 cm long; midrib prominent, stout
below, evident but hardly raised or somewhat sunken above; venation hopeatype; lateral
veins 6–9 pairs, ascending, arched, slender but prominent below, narrowly sunken above,
often with prominent pustular domatia; intercostal venation densely scalariform; petiole
0.6–1.2 cm long, slender. Inflorescences terminal or axillary, to 7 cm long, singly branched,
branchlets bearing to 7 dense secund flowers; bracts fugaceous. Flowers: buds ellipsoid, to
2 × 1 mm; petals lanceolate, cream; stamens c. 15, anthers ellipsoid, connectival appendage
c. 3x the length of anther; ovary and stylopodium cylindrical, truncate, more or less
medially constricted, apical platform papillose, style short, columnar. Fruits: pedicels c. 2
mm long; calyx lobes unequal, 2 longer lobes broadly lorate, 9.5 × 2.2 cm, obtuse, tapering
to c. 4 mm above the saccate base, 3 shorter ones broadly ovate, to 0.4 × 0.6 cm, subacute,
shorter than the nut. Nuts subglobose, to 0.6 × 0.5 cm, shortly apiculate.
Vernacular name. Sabah—selangan batu jantan (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo, very local and
generally uncommon. In Sabah recorded from Beaufort, Keningau, Kinabatangan, Kudat,
Lahad Datu, Sandakan, and Tawau districts (e.g., SAN 15205, SAN 16422, SAN 17718, SAN
36463, and SAN 96656) and in Sarawak from Bintulu, Kapit and Lundu districts (e.g., S
1815, S 9643 and S 17722).
179
Ecology. Uncommon in mixed dipterocarp forest, on clayrich periodically flooded

alluvium, at altitudes below 200 m. Endangered.
Notes. Material from Kudat, Lahad Datu and Sandakan districts differs in the smaller
somewhat chartaceous leaf, wrinkling on drying but otherwise conforms with the species.
38. Hopea sphaerocarpa (F.Heim) P.S.Ashton

(Greek, sphaero = round, karpos = fruit)
Basionym: Balanocarpus sphaerocarpus F.Heim op. cit. (1892) 77, Merrill op. cit. (1921) 407,
Masamune op. cit. 483. Type: Beccari PB 3021, Borneo, ‘Sarawak’ (holotype P).
Small tree, to 10 m, to 10 cm diameter; crown remaining monopodial; with low, thin

buttresses and flying buttresses and stilt roots. Bark smooth. Young parts, including
sometimes leaf venation but excluding inflorescence, tawny puberulent; indumentum
caducous on calyx, more or less persistent elsewhere. Twigs c. 0.5 mm diameter apically,
slender, sparsely branched. Leaves chartaceous, undulate, more or less flat, drying tawny
brown below, dark purplish above; blade ovate to oblonglanceolate, 3.5–10 × 1.4–4.5 cm,
base obtuse to subcordate, apex with slender acumen to 1 cm long; midrib slender, sunken
above, elevated below; venation subdryobalanoid; lateral veins 9–11 pairs, slender but
elevated below, with short indistinct intermediates; intercostal venation densely scalariform,
slender but evident below; petiole 0.3–0.6 cm long, slender. Inflorescences in axillary pairs
or occasionally terminal, very slender, lax, to 10 cm long, singly branched, branchlets zig
zag, bearing to 6 remote distichous flowers; bracts linear, to 2 mm long, subpersistent.
Flowers: buds subglobose, to 2 mm diameter; petals ellipticoblong, dark crimsonred;
stamens c. 15, anthers subglobose, connectival appendage 3–5x the length of anther; ovary
and stylopodium cylindrical, truncate, somewhat constricted medially, style short, narrowly
conical. Fruits: pedicels to 4 mm, long; calyx lobes subequal, ovate, to 0.8 × 0.7 cm, acute,
appressed to the nut. Nuts broadly ovoid, to 1 × 0.8 cm, apex exposed with prominent
truncate stylopodium remnant.
Distribution. Endemic in Borneo. In Sarawak recorded from Bau, Kuching, Lundu, and Sri
Aman districts (e.g., S 15004, S 29455, S 37849, S 49928, and S 68433).
Ecology. Local, but often in dense populations; in mixed dipterocarp forest on leached
sandy clay soils, at altitudes below 200 m. Critically endangered.
Notes. An apparantly related entity has been collected in young fruit on G. Gading, Lundu
district (S 12601 and S 15570). In differs in its 12–15 pairs of steeply arching lateral veins.
Confirmation of its status must await flowering material.
39. Hopea tenuinervula P.S.Ashton

(Latin, tenuis, slender, nervulus, a little nerve; the slender intercostal veins)
Coode et al. (eds.) op. cit. 73. Type: Ariffin S 9616, Borneo, Sarawak, Lundu district (holotype K;
isotype KEP). Synonym: Hopea philippinensis auct. non Dyer: Ashton op. cit. (1964) 107.
180
Subcanopy tree, to 30 m tall, to 35 cm diameter; buttresses and stilt roots to 1 m tall, thin.
Bark thinly powdery reddish brown mottled, flaky. Young vegetative parts and perianth
outside densely pale tawny puberulent; indumentum persistent on buds, stipules, petals, and
somewhat so on veins and midrib on both surfaces, elsewhere caducous. Twigs to 2 mm
diameter apically, dark, sparingly branched. Leaves thinly coriaceous, not bullate, drying
dark chocolatebrown below, paler above; blade narrowly ovate to lanceolate, 10–27 × 3–
5.5 cm, base markedly unequal, obtuse, margin narrowly revolute, apex with slender
acumen to 1 cm long; midrib stout and raised on both surfaces; venation hopeatype; lateral
veins dense, 12–21 pairs, slender but prominent below, arched and running concurrent to
margin before terminating; intercostal venation very slender and not elevated, densely
sinuatescalariform; petiole 0.3–0.7 cm long, stout. Inflorescences in axillary pairs, rarely
terminal, lax, to 8 cm long, singly branched, branchlets bearing to 4 secund flowers; bracts
deltoid, to 1 mm long, subpersistent. Flowers: buds ellipsoid, to 4 × 2 mm; petals oblong
lanceolate, pale yellow; stamens c. 15, anthers oblongellipsoid, connectival appendage c.
2x the length of anther; ovary ovoid, stylopodium and style as long as ovary, hourglass
shaped. Fruits subsessile; calyx lobes unequal, 2 longer lobes spatulate, narrowly obtuse, to
10 × 1.7 cm, tapering to 2 mm above the saccate base, not auriculate, 3 shorter ones
lanceolate, to 3 × 0.7 cm, slightly flanged along margin, conceiling the nut. Nuts ovoid, to
1.2 × 0.8 cm, shortly apiculate.
Vernacular names. Sabah—selangan daun serong (preferred name). Sarawak—luis daun

serong (preferred name).
Distribution. Endemic in Borneo. In Sabah only known from Gaya Is., Kota Kinabalu
district (e.g., SAN 38708, SAN 46596 and SAN 135353) and in Sarawak from Kuching,
Lundu and Sibu districts (e.g., S 11091, S 12614, S 14812, S 25452, and S 37782). Also
occurring in Brunei (e.g., BRUN 2624, BRUN 3290 and S 1187) and E and C Kalimantan
(e.g., bb. 17854 and Ridsdale PBU 49).
Ecology. Very local, but there often in dense populations, in mixed dipterocarp forest on
leached sandy and sandy clay soils, at altitudes to 400 m. Endangered.
40. Hopea treubii F.Heim

(M. Treub, 1851–1910, sometime Director of the Bogor Botanic Gardens, Indonesia)
Bull. Mens. Soc. Linn. Paris 2 (1891) 955; Merrill op. cit. (1921) 403, Masamune op. cit. 492; Ashton
op. cit. (1964) 111, op. cit. (1968) 57, op. cit. (1982) 406; PROSEA op. cit. 254; Coode et al. (eds.)
op. cit. 73. Type: Beccari PB 2895, Borneo, Sarawak, Matang (holotype P).
Canopy tree, to 40 m tall, to 80 cm diameter, with irregular crown supported by a few large
ascending twisted branches; buttresses and flying buttresses low. Bark becoming densely v
section fissured, tawny brown. Sapwood straw yellow, hard; heartwood dark warm brown.
Parts of petals exposed in bud pubescent; parts otherwise glabrous. Twigs to 1.5 mm
diameter apically. Leaves coriaceous, drying yellowish brown; blade broadly elliptic to
obovate, 5–8 × 3–5.5 cm, base cuneate, margin more or less revolute, apex shortly broadly
bluntly acuminate; midrib stout, somewhat raised on both surfaces; venation dryobalanoid;
main lateral veins elevated below, prominently arched, with long intermediates; petiole c. 1
cm long, rather stout. Inflorescences terminal or in axillary pairs or ramiflorous, rigid,
slender, ascending, to 8 cm long, singly branched, branchlets bearing to 7 secund flowers.
Flowers: buds ellipsoid, to 2.5 mm long, on prominent pedicels; petals narrowly lanceolate,
181
lemonyellow; stamens c. 10, anthers broadly oblong, connectival appendage c. 2x the

length of anther; ovary and stylopodium cylindrical, subtruncate, papillose at apex, style
short. Fruits: pedicels to 3 mm long, slender; calyx lobes unequal, 2 longer lobes spatulate,
obtuse, to 3.5 × 0.7 cm, tapering to 2 mm wide above the saccate base, 3 shorter ones ovate,
to 0.8 × 0.2 cm, appressed to but shorter than the nut. Nuts narrowly ovoid, to 1.1 × 0.5 cm,
acute.
Vernacular name. Sarawak—luis daun tebal (preferred name).
Distribution. Endemic in Borneo. Known in Sarawak from Bau, Belaga, Kuching, Lundu,
and Simunjan districts (e.g., Chew CWL 1163, S 9482, S 10134, S 28979, S 43972, and S
68718). Also occurring in Brunei (e.g., BRUN 620, BRUN 3262, BRUN 3283, BRUN 5432,
and Coode et al. MC 7103).
Ecology. Scattered in mixed dipterocarp forest on leached yellow sands, on both sandstone
and the Arip, Mukah, rhyolite, at altitudes to 400 m. Very local, highly vulnerable though
frequent in Bako NP and recorded from Mulu NP.
41. Hopea vaccinifolia Ridl. ex P.S.Ashton

(Latin, vaccinium = the bilberry, folius = a leaf; the tiny Vacciniumlike leaves)
op. cit. (1980) 117; Coode et al. (eds.) op. cit. 73. Type: Hose 583, Borneo, Sarawak, Marudi district
(holotype K).
Understorey monopodial tree, to 15 m tall, to 15 cm diameter, with flying buttresses and

stilt roots. Bark smooth. Young twig, leaf bud, stipules and petiole densely shortly
persistently greyish brown puberulent; parts of petals exposed in bud sericeous; flower
sepals fimbriate; other parts glabrous. Twigs to 0.5 mm diameter apically, very slender,
much horizontally branched, with persisting distichous leaf arrangement. Leaves
chartaceous, drying greenish grey; blade elliptic to broadly ovate, 1–2.5 × 0.4–1.2 cm, base
equal; midrib slender more or less slightly elevated below, obscure and sunken above;
venation subdryobalanoid; lateral veins 6 pairs, indistinct, hardly elevated, with short
slender intermediates, arched; intercostal venation reticulate, evident below; petiole to 0.2
cm long, slender. Inflorescences in axillary pairs, slender, to 1.3 cm long, singly branched,
branchlets bearing to 3 remote flowers; bracts deltoid, to 1 mm long, subpersistent.
Flowers: buds subsessile; petals oblong, strongly contorted, rotate distally, dark red;
stamens c. 10, anthers subglobose, connectival appendage c. 2x the length of anther; ovary
and stylopodium cylindrical, truncate, puberulent on apical platform, style short. Fruits:
calyx lobes subequal, ovate, to 0.4 × 0.35 cm, obtuse or acute, appressed round the base of
the nut. Nuts ovoid, to 0.8 × 0.6 cm, minutely truncate.
Vernacular names. Sabah—selangan (preferred name). Sarawak—luis ribu (preferred

name).
Distribution. Endemic in Borneo. Known in Sarawak from Marudi district (e.g., BRUN
5664 and S 33). Also occurring in Brunei (e.g., BRUN 140, BRUN 3035, FMS 30614, and
FMS 35567).
182
Ecology. Locally abundant, in dense clusters, on the giant podsols of the Pleistocene sea
beach, in tall kerangas forest, at low altitudes. Critically endangered throughout its range.
42. Hopea vesquei F.Heim

(J. Vesque, 1848–1895, a Luxemburg botanist at the Paris Herbarium)
Bull. Mens. Soc. Linn. Paris 2 (1891) 971; Merrill op. cit. (1921) 403; Masamune op. cit. 492; Browne
op. cit. 122; Ashton op. cit. (1968) 57, op. cit. (1982) 401; Anderson op. cit. (1980) 118; PROSEA op.
cit. 254. Type: Beccari PB 2550, Borneo, ‘Sarawak’ (holotype P).
Subcanopy to low canopy tree, to 30 m tall, to 30 cm diameter; buttresses to 80 cm tall, thin;

stilt roots present. Bark becoming patchily cracked; inner bark pale brown or pink. Twigs,
leaf bud, stipules, petiole, inflorescence, and parts of flower exposed in bud densely greyish
brown puberulent; parts otherwise glabrous. Twigs c. 1 mm diameter apically, much
branched. Leaves thickly coriaceous; blade broadly ovate, 3.5–6 × 1.5–3.5 cm, base
subobtuse, unequal, apex with slender acumen to 1 cm long; midrib stout, raised on both
surfaces, not drying black; venation dryobalanoid; main lateral veins 10–13 pairs, slender,
hardly elevated but distinct below, with shorter intermediates, basal pair short; intercostal
venation obscure, reticulate; petiole 0.6–0.7 cm long, slender. Inflorescences terminal or
axillary, solitary, to 3 cm long, singly branched, branchlets bearing to 5 secund flowers.
Flowers: buds ellipsoid, to 3 × 2 mm; petals lanceolate, pale creamyellow; stamens c. 15,
anthers subglobose, connectival appendage slightly longer than anther; ovary ovoid,
stylopodium obscure, style columnar, c. 1½x the length of ovary. Fruits: pedicels to 2 mm
long; calyx lobes unequal, 2 longer lobes spatulate, obtuse, to 3.4 × 0.8 cm, tapering to c. 2
mm above the saccate base, 3 shorter ones ovate, acute, to 0.4 × 0.4 cm, clasping the base
of the nut. Nuts cylindrical, to 1.5 × 0.3 cm, tapering to c. 1 mm filiform style remnant.
Vernacular name. Sarawak—luis tebal (preferred name).
Distribution. Endemic in Borneo. Known only in W Sarawak from Kuching, Lundu and
Miri districts (e.g., S 3021, S 6363, S 9306, S 12498, and S 15171).
Ecology. Local, but there common in mixed dipterocarp forest on usually shallow yellow
sandy soil, at altitudes below 200 m. Locally common in Bako and Lambir NPs but
elsewhere highly vulnerable.
Notes. Difficult to distinguish from H. dyeri without flowers or ripe fruit (cf. Meijer &
Wood op. cit. 226).
43. Hopea wyattsmithii Wood ex P.S.Ashton

(J. WyattSmith, 1917–2002, former Silviculturist at the Forest Research Institute,
Malaysia)
Wood op. cit. 227; Burgess op. cit. 129; Anderson op. cit. (1980) 118; Coode et al. (eds.) op. cit. 73.
Type: G.H.S. Wood SAN 15061, Borneo, Sabah, Beaufort district (holotype K; isotypes KEP, SAN).
183
Subcanopy tree, to 25 m tall, to 30 cm diameter, with prominent flying buttresses and stilt
roots. Bark smooth; inner bark plumred. Parts glabrous but for sericeous parts of petals
exposed in bud and stylopodium. Twigs c. 1 mm diameter apically, slender, sparsely
branched. Leaves thinly coriaceous, sparsely greyish lepidote below, drying greyish below,
pale tawnybrown above; blade broadly ovate to elliptic, 9–14 × 5.5–9 cm, base equal,
broadly cuneate or sometimes obtuse, apex caudate, acumen to 2 cm long; midrib slender
but evident and raised above, acutely elevated below; venation hopea or subdryobalanoid
type; lateral veins 4–8 pairs, with the main ones irregularly spaced owing to occasional
prominent intermediates, slender, elevated below, arched but the basal 2–3 pairs at first
decurrent with midrib, straight; intercostal venation evident but unraised, scalariform, lax;
petiole 1.2–1.7 cm long, slender. Inflorescences solitary, terminal or axillary, to 6 cm long,
singly branched, branchlets bearing to 6 secund flowers; bracts deltoid, to 1 mm long, not at
first caducous. Flowers: buds subglobose, to 3 mm diameter; petals oblonglanceolate, dark
red; stamens c. 15, anthers broadly oblong, connectival appendage c. 2x the length of
anther; ovary and stylopodium hourglassshaped, tapering into short style. Fruits
subsessile; calyx lobes subequal, ovate, subacute, to 1 × 0.8 cm, clasping the nut. Nuts
broadly ovoid, to 1.2 × 1.2 cm, acute, the apical ¼ exposed.
Vernacular names. Sabah—selangan daun bulat (preferred name). Sarawak—luis putih

(preferred name).
Distribution. Endemic in Borneo. Recorded in Sabah from Beaufort, Kinabatangan, Labuk

Sugut, Lahad Datu, Sandakan, and Tawau districts (e.g., SAN 16467, SAN 22634, SAN
28633, and SAN 44228) and in Sarawak from Kapit, Limbang and Lubok Antu districts
(e.g., S 692, S 32275 and S 32288). Also occurring in Brunei (e.g., BRUN 885, FMS 35475
and S 5589).
Ecology. In small groups of individuals, on clayrich soils in mixed dipterocarp forest on

moist slopes and low hills, at altitudes to 200 m. Rare and critically endangered by forest
conversion.
6. PARASHOREA Kurz
(Greek, para = similar to; resembling Shorea)
urat mata (preferred name)
J. As. Soc. Beng. 39, 2 (1870) 65; Slooten, Bull. Jard. Bot. Buitenz. 3, 8 (1927) 370; Symington,
Malay. For. Rec. 16 (1943) 97; Masamune, EPB (1942) 492; Browne, FTSB (1955) 126; Ashton,
MDB (1964) 82, MDBS (1968) 38, FM I, 9 (1982) 379; Meijer & Wood, Sabah For. Rec. 5 (1964)
190; Burgess, TBS (1966) 140; Anderson, CLTS (1980) 118; PROSEA 5, 1 (1993) 325; Newman et
al., MDFBLHW (1996) 66.
Emergent trees, with stout slightly concave hardly branched buttresses; crown dense,
hemispherical or domeshaped, the leaves crowded towards the surface (except P.
macrophylla). Bark mauvegrey to purplish brown, becoming narrowly shallowly fissured
and flaky, the fissures and buttress ridges densely whitish corky lenticellate; inner bark
fibrous, warm brown; dammar exudations opaque pale yellowish cream. Sapwood pale
whitish to yellowish; heartwood pale to chocolatebrown. Leaf buds often prominent, ovoid
or falcate. Stipules linear to hastate, falling early. Leaves drying mauvegrey, undersurface
184
greyish to silvery, persistently so in juveniles (except P. parvifolia and P. tomentella);

juvenile leaves often peltate; blade oblongovate, often asymmetric; midrib evident above;
lateral veins without intermediates, rather straight, not joining into intramarginal veins,
dense with plicate folds remaining evident in the open leaf (except P. parvifolia); intercostal
venation scalariform (except P. parvifolia); petiole mostly weakly geniculate.
Inflorescences paniculate, terminal or axillary. Flowers: buds lanceolate; sepals narrow,
hardly imbricate; petals falling separately; stamens 15 in 3 whorls, exceeding ovary in bud,
filaments short, dilated at base, abruptly tapering, anthers with 4 pollen sacs, narrowly
oblong, connectival appendage short or stout columnar; ovary small, ovoid, pubescent, with
or without stylopodium, style long, filiform. Fruits: pedicels short; mature calyx lobes
valvate, all long, spatulate, but unequal with 3 longer and broader than the other 2, pushed
apart by the ripening nut. Nuts globose to ellipsoid, more or less pubescent and white corky
lenticellate; style caducous. Germination as in Shorea but seedling leaves linear.
Distribution. About 14 species; S Myanmar, Thailand, IndoChina and southernmost parts

of China to Sumatra, Borneo and the Philippines. Six species occurring in Sabah and
Sarawak; throughout but absent in Sarawak west of Sri Aman district.
Ecology. In mixed (and in IndoBurmese seasonal wet evergreen) dipterocarp forest on clay
soils, especially fertile and moist places, at altitudes to 1400 m. Usually scattered but some
species in some areas locally abundant.
Uses. Parashorea macrophylla, P. malaanonan and P. tomentella are light hardwoods

traded in Sabah as white seraya; the last two were among the most important export timbers
from E Sabah, now largely exhausted. The other species are medium hardwoods. All are
generally mixed with red meranti consignments.
Key to Parashorea species

1. Leaf blade at least 30 cm long; lateral veins at least 23 pairs; twigs flattened; crown
largeleaved, long remaining monopodial……………………………2. P. macrophylla
Leaf blade at most 20 cm long; lateral veins at most 20 pairs; twigs round; crown dense,
domeshaped……………………………………………………………………………2
2. Mature leaf blade without visible folds between the at most 9 curved pairs of lateral
veins, not silvery lepidote below……………………………………..…4. P. parvifolia
Leaf blade with visible corrugations between the at least 9 pairs of straight lateral veins,
more or less greyish silvery lepidote or tawny velutinous below…………………..…3
3. Midrib glabrous below; petiole glabrescent……………………..…3. P. malaanonan

Midrib below and petiole tomentose or velutinous…………………………………...…4
4. Leaf undersurface persistently pale tawny velutinous………………..…6. P. tomentella

Leaf undersurface more or less sparsely caducous yellowish brown scabridpubescent
on the veins………………………………………………………………………….…5
185
5. Leaf lateral veins 9–12 pairs; petiole not geniculate.……………………..…1. P. lucida

Leaf lateral veins 8–10 pairs; petiole geniculate……………………..…5. P. smythiesii
1. Parashorea lucida (Miq.) Kurz

(Latin, lucidus = clear; the venation)
J. As. Soc. Beng. 39, 2 (1870) 66; Slooten op. cit. (1927) 372, p.p; Symington op. cit. (1943) 102;
Ashton op. cit. (1968) 38, op. cit. (1982) 387; PROSEA op. cit. 331; Newman et al. op. cit. 70.
Basionym: Shorea lucida Miq., Fl. Ned. Ind., Suppl. (1862) 487. Lectotype (designated here):
Diepenhorst HB 2074, Sumatra, Pariaman (hololectotype U). Synonym: Shorea subpeltata Miq., op.
cit. 488.
Tree to 60 m tall, to 1.5 m diameter; crown dense and domeshaped. Young parts and leaf
venation below more or less caducously greyish brown scabridpubescent; petiole,
inflorescence, parts of flower exposed in bud, and ovary persistently so. Twigs terete;
stipule scars short, pale, horizontal. Leaf buds ovoid, to 4 × 2 mm. Leaves thinly
coriaceous, dull and more or less silvery stellate pubescent below, more or less persistently
corrugated in between the lateral veins; blade ovatelanceolate to elliptic, 6–14 × 2.5–6.5
cm, base unequal, broadly cuneate to subcordate, apex acuminate, acumen to 1 cm long;
lateral veins 9–12 pairs, slender but prominent below, straight; intercostal venation slender
but distinct below; petiole 1–2 cm long, tomentose, not geniculate. Inflorescences to 12 cm
long, with secund flowers. Flowers: buds to 7 × 4 mm; connectival appendage acicular,
prominent and longer than anthers. Fruits: calyx lobes to 8 × 1.7 cm. Nuts to 2.5 cm
diameter.
Distribution. Sumatra and Borneo. In Borneo, known only in C and E Sarawak from
Belaga, Bintulu, Kapit, and Sarikei districts (e.g., S 18059, S 22238, S 22316, S 27990, and
S 48164).
Ecology. In Sarawak, local and uncommon and scattered in mixed dipterocarp forest on clay
and sandy clay soils, at altitudes below 700 m. Vulnerable owing to land conversion.
2. Parashorea macrophylla Wyatt Smith ex P.S.Ashton

(Greek, makros = big, phullon = leaf; with large leaves)
op. cit. (1980) 118; PROSEA op. cit. 331; Newman et al. op. cit. 71; Coode et al. (eds.), CLBD (1996)
73. Type: Ladi BRUN 2002, Brunei, Kuala Belalong (holotype K; isotypes KEP, L).
Large tree, to 50 m tall, to 1 m diameter, remaining monopodial into maturity; crown diffuse
and adorned with the giant silvery leaves. Young parts, buds and inflorescence densely
ochreous puberulent. Twigs c. 12 × 5 mm apically, compressed at first; stipule scars
amplexicaul. Leaf buds linearfalcate, to 9 × 0.8 cm. Stipules to 15 × 2.5 cm. Leaves
distichous, subchartaceous, silvery below, prominently corrugated between lateral veins;
blade oblongelliptic, 30–50 × 16–24 cm, base subcordate, apex obtuse to shortly
acuminate; lateral veins 28–36 pairs, straight, prominent below; intercostal venation very
slender, dense; petiole 3–5 cm long. Inflorescences to 16 cm long, 2–3branched, terminal
branches cymose; bracts ovate, to 40 × 25 mm, acute, amplexicaul. Flowers: buds to 20 × 8
mm; connectival appendage prominent, c. ½x the length of anther. Fruits: calyx lobes to 22
× 2 cm. Nuts ellipsoid, to 2.5 × 1.5 cm.
186
Vernacular names. Sarawak—bilat (Iban), peran (preferred name).
Distribution. Endemic in Borneo. In Sarawak known from Bintulu, Kapit, Limbang,

Marudi, Miri, and Tatau districts (e.g., S 13791, S 18176, S 23049, S 23471, and S 75391).
Also occurring in Brunei (e.g., BRUN 901, BRUN 3136, FMS 30388, and SAN 17377) and
W Kalimantan.
Ecology. In mixed dipterocarp forest on moist lower slopes and periodically flooded
alluvium, along the inland rivers, on clay soils, at altitudes to 300 m. The gigantic silvery
fallen leaves on the forest floor look like crashed model zeppelins. Occurring in Mulu NP
but vulnerable elsewhere.
3. Parashorea malaanonan (Blanco) Merr.

(Tagalog, mala = false, anonang = the custard apple; a putative vernacular name)
Sp. Blancoan. (1918) 271, PEB (1929) 400; Slooten op. cit. (1927) 375; Symington, Gard. Bull. S. S.
9 (1938) 334; Keith op. cit. 19; Masamune op. cit. 192; Browne op. cit. 128; Ashton op. cit. (1964) 84,
cit. (1980) 118; PROSEA op. cit. 332; Newman et al. op. cit. 72; Coode et al. (eds.) op. cit. 73.
Basionym: Mocanera malaanonan Blanco, Fl. Filip. ed. 1 (1837) 858. Neotype (designated here):
Merrill Sp. Blancoan. 1053 (= US 874771), the Philippines, Luzon, Laguna Province, Mt. Maquiling
(A, US). Synonyms: Dipterocarpus malaanonan (Blanco) Blanco, Fl. Filip. ed. 2 (1845) 312; Shorea
malaanonan (Blanco) Blume Mus. Bot. Lugd. Bat. 2 (1852) 34; Parashorea plicata Brandis, J. Linn.
Soc. Bot. 31 (1895) 104.
Large tree to 60 m tall, to 2 m diameter, with dense domeshaped crown. Bark dark,
eventually blackish purple, fissured, thinly flaky. Young parts spasely greyish brown
pubescent, glabrescent except on bud, inflorescence and nut. Twigs terete, with amplexicaul
stipule scars. Leaf buds lanceolatefalcate, to 6 × 2 mm. Stipules hastate, to 15 × 6 mm.
Leaves thinly coriaceous, with visible corrugations between lateral veins, greyish silvery
lepidote below; blade broadly ellipticovate, 9–15 × 3.5–7.5 cm, base unequal, obtuse to
broadly cuneate, margin wavy distally, apex acuminate, acumen to 1 cm long; midrib
prominent and glabrous below; lateral veins 9–14 pairs, prominent below; intercostal
venation slender, sinuate; petiole 1.2–2 cm long, somewhat geniculate, glabrescent.
Inflorescences to 18 cm long, doubly branched. Flowers: buds to 14 × 8 mm; connectival
appendage somewhat longer than anther. Fruits: calyx lobes to 16 × 1.7 cm. Nut to 1.7 cm
diameter.
Vernacular names. Sabah—urat mata daun licin (preferred name). Sarawak—urat mata
(preferred name).
Distribution. Borneo and the Philippines. In Sabah widespread and recorded from Beaufort,
Kinabatangan, Kota Belud, Kota Merudu, Kudat, Lahad Datu, Papar, Ranau, Sandakan,
Semporna, Sipitang, Tambunan, Tawau, and Tenom districts (e.g., SAN 15225, SAN 17229,
SAN 28928, SAN 30811, and SAN 76053) and in Sarawak from Lawas and Miri districts
(e.g., S 3431, S 24976, and S 25329). Also occurring in Brunei (e.g., FMS 30531, KEP
80138, S 5728, and SAN 17065) and E Kalimantan (e.g., Endert 5165).
Ecology. In mixed dipterocarp forest on deep friable clay soils overlying shale, basic and
intermediate igneous rocks, at altitudes to 1300 m. Once abundant in the eastern parts of
187
Sabah lowlands and still locally so in surviving forests; rare in areas west of the Crocker
Range (Sabah) and in Sarawak. Vulnerable.
Uses. In the past, the species supplied timber export from Sabah.
4. Parashorea parvifolia Wyatt Smith ex P.S.Ashton

(Latin, parvus = small, folius = leaf; smallleaved)
Wood op. cit. 195; Burgess op. cit. 140; Anderson op. cit. 118; PROSEA op. cit. 332; Newman et al.
op. cit. 73; Coode et al. (eds.) op. cit. 73. Type: Haviland 2810/2331, Sarawak, Belaga (holotype K).
Tree to 60 m tall, to 1.5 m diameter; crown dense, domeshaped. Bark at first blackish
purple. Young parts pale yellowish brown puberulent, indumentum persistent only on bud,
inflorescence, and ovary. Twigs terete, slender; stipule scar short. Leaf buds to 3 × 1 mm.
Stipules narrowly hastate, to 4 mm long, fugaceous. Leaves thinly coriaceous, not
corrugated, not silvery lepidote below; blade elliptic to narrowly ovate, 6–9 × 3–4.5 cm,
base broadly cuneate, equal, apex with slender acumen to 1.5 cm long; midrib slender but
elevated below; lateral veins 8–10 pairs, slender and hardly raised below, arching, well
spaced; intercostal venation subreticulate, wellspaced; petiole 1–1.8 cm long, somewhat
geniculate. Inflorescence to 14 cm long, singly branched. Flowers: buds to 4.5 × 3 mm;
connectival appendage short, slightly extruded above anther. Fruits: calyx lobes to 8.5 ×
1.7 cm. Nuts to 1.5 cm diameter.
Vernacular names. Sabah—urat mata daun kecil (preferred name). Sarawak—urat mata
bukit (preferred name).
Distribution. Endemic in Borneo. In Sabah known from Kota Belud, Labuk Sugut,
Pandawan, Penampang, Pensiangan, Ranau, and Tenom districts (e.g., SAN 37782, SAN
76191, SAN 95283, and SAN 119902) and in Sarawak from Belaga, Bintulu, Lawas,
Limbang and Miri districts (e.g., KEP 79350, S 18194 and S 43169). Also occurring in
Brunei (e.g., BRUN 2604, BRUN 3013, BRUN 3381, and FMS 34567) and E Kalimantan
(e.g., bb. 11772 and bb. 18165).
Ecology. Scattered, usually uncommon or rare in mixed dipterocarp forest on clay soils.
Especially on upper slopes and ridges, and at altitudes to 1350 m in upper dipterocarp forest.
Occurring in Lambir NP but vulnerable elsewhere.
5. Parashorea smythiesii Wyatt Smith ex P.S.Ashton

(B.E. Smythies, Conservator of Forests, Sarawak 1959–1964; ornithologist and botanist)
Wood op. cit. 197; Burgess op. cit. 140; Anderson op. cit. 118; PROSEA op. cit. 332; Newman et al.
op. cit. 74; Coode et al. (eds.) op. cit. 74. Type: Ladi BRUN 2000a, Brunei, Kuala Belalong (holotype
K; isotypes KEP, L).
Large tree, to 55 m tall, to 1.2 m diameter; crown dense, domeshaped. Young parts pale
yellowish brown scabridpuberulent, indumentum persistent on inflorescence, bud and
ovary. Twigs terete, c. 3 mm diameter apically; stipule scars to 1.5 mm at first, horizontal.
188
Leaf buds to 5 × 3 mm. Leaves coriaceous, silvery lepidote below, drying greyish mauve
above, evidently corrugated between lateral veins; blade ellipticoblong, 10–18 × 4–7 cm,
base unequal, obtuse, apex with short slender acumen to 0.5 cm long; midrib prominent and
tomentose below, narrowly furrowed above; lateral veins 8–10 pairs, prominent below,
somewhat arched; intercostal venation distinct, dense; petiole 1.5–2.2 cm long, stout,
geniculate, tomentose. Inflorescences to 10 cm long, doubly branched. Flowers: buds
ovoid, to 7 × 4 mm; connectival appendage short, not exceeding anther. Fruits: calyx lobes
to 10 × 1.7 cm. Nuts to 1.3 × 0.9 cm.
Vernacular names. Sabah—urat mata batu (preferred name). Sarawak—meruyun (Iban),

urat mata daun putih (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from Beaufort, Sipitang and Tawau
districts (e.g., SAN 18720, SAN 25272, SAN 44595, and SAN 72180) and in Sarawak from
Belaga, Bintulu, Kapit, Limbang, Miri, and Tatau districts (e.g., S 29151, S 32551, S 41302,
S 41329, and S 56857). Also occurring in Brunei (e.g., BRUN 3016, BRUN 3169, FMS
35448, and KEP 80138), and E Kalimantan (e.g., bb. 17890 and bb. 18320).
Ecology. Locally frequent in mixed and occasionally upper dipterocarp forests on clay soils,
at altitudes to 1000 m. Occurring in Lambir and Mulu NPs, probably not yet vulnerable.
6. Parashorea tomentella (Symington) Meijer Fig. 18.

(Latin, tomentellus = with small stuffing (e.g., of a pillow); the indumentum)
(1982) 385; PROSEA op. cit. 333; Newman et al. op. cit. 75. Basionym: Parashorea malaanonan
(Blanco) Merr. var. tomentella Symington op. cit. (1938) 338. Type: Otik FMS 38745, Borneo, Sabah,
Sandakan district, KabiliSepilok FR (holotype KEP).
Large emergent tree, to 65 m tall, to 2 m diameter; crown dense, domeshaped. Young twigs,
buds, leaf undersurface, petiole, inflorescence, bracts outside, parts of calyx and corolla
exposed in buds, and ovary densely evenly more or less persistently pale tawny velutinous.
Twigs c. 3 mm diameter apically, muchbranched, ribbed, becoming terete, smooth, dark
brown; stipule scars slender, horizontal, amplexicaul. Buds lanceolate, acute, to 10 × 4 mm.
Stipules narrowly lanceolate, to 16 × 6 mm. Leaves subcoriaceous, subpersistently
corrugated between veins; blade ellipticoblong, 10–20 × 5–10 cm, base obtuse to
subcordate, subequal (peltate in young trees and saplings), margin frequently narrowly
subrevolute, apex subacute or broadly acuminate, acumen to 1 cm long; midrib stout and
prominent below, elevated above; lateral veins 11–13 pairs, ascending, prominent below,
somewhat arched; intercostal venation densely scalariform, evident and slightly elevated
below; petiole 1.5–2.5 cm long, c. 3 mm diameter, stout, hardly geniculate. Inflorescences
to 13 cm long, singly or doubly branched, branches bearing to 3 flowers; bracts lanceolate,
to 10 × 4 mm. Flowers: buds broadly lanceolate, 15 × 10 mm; sepals narrowly deltoid;
stamens somewhat shorter than style, filament compressed, tapering, anthers oblonglinear,
connectival appendage shorter than anthers, stoutly acicular; ovary small, ovoid, style
columnar, c. 5x the length of ovary, stout, pubescent but for the apical 1/5. Fruits: pedicels
c. 3 mm long; calyx lobes aliform, 3 larger lobes spatulate, to 20 × 2.3 cm, obtuse, 2 shorter
ones lorate, to 10 × 0.8 cm, acute. Nuts subglobose, to 2 cm diameter, verrucose, apiculate.
Vernacular name. Sabah—urat mata beludu (preferred name).
189
Fig. 18. Parashorea tomentella. A, fruiting leafy twig; B, detail of indumentum on lower
leaf surface; C, flower bud; D, longitudinal section of open flower; E, adaxial view of petal;
F, abaxial view of petal; G, adaxial view of stamens; H, abaxial view of stamens; I, fruit
with one of the smaller calyx lobes cutoff; J, longitudinal section of fruit. (A–B from SAN
18703, C–H from FMS 55170, I–J from SAN A 1464.)
190
Distribution. Endemic in Borneo. In Sabah recorded from Kinabatangan, Labuk Sugut,

Lahad Datu, Papar, Sandakan, and Tawau districts (e.g., SAN 18785, SAN 31098, SAN
38163, SAN 96625, and SAN 134924). Also occurring in E Kalimantan (e.g., Ambriansyah
& Arifin Berau 804, Ambriansyah & Arifin Berau 1050 and Kessler et al. Berau 851).
Ecology. Locally abundant in mixed dipterocarp forest on fertile clay soils, on undulating
land and alluvium, at altitudes to 200 m. Vulnerable owing to forest conversion.
7. SHOREA Roxb. ex Gaertn.f.

(Sir John Shore, GovernorGeneral for the British East India Company, 1793–1798)
lun, meranti, selangan batu (preferred names), seraya
Fruct. 3 (1805) 48; Brandis, J. Linn. Soc. Bot. 31 (1895) 73; Masamune, EPB (1942) 492; Symington,
Malay. For. Rec. 16 (1943) 1; Ashton, MDB (1964) 115, MDBS (1968) 60, FM 1, 9 (1982) 436;
Meijer & Wood, Sabah For. Rec. 5 (1964) 48; Burgess, TBS (1966) 134; Anderson, CLTS (1980)
118; PROSEA 5, 1 (1993) 384; Coode et al. (eds.), CLBD (1996) 74; Newman et al., MDFLHW
(1996) 77, MDFMHHW (1998) 159. Synonyms: Doona Thwaites in Hooker, Kew J. 3 (1851) t. 14;
Pentacme A.DC., Prod. 16, 2 (1868) 626; Isoptera Scheff. ex Burck, Med. Lands Pl. Tuin 3 (1886) 27;
Ridleyinda Kuntze, Rev. Gen. Pl. 1 (1891) 65; Richetia F.Heim, Bull. Mens. Soc. Linn. Paris (1891)
975; Parahopea F.Heim, Rech. Dipt. (1892) 66; Pachychlamys (Dyer ex King) Dyer ex Ridl., FMP 1
(1922) 233.
Emergent, sometimes main canopy trees; buttresses stout or thin (in most species of sect.
Shorea), somewhat concave, usually more or less as wide as tall; stilt roots or flying
buttresses rarely present; mature crown sympodial, hemispherical, spreading. Bark various.
Stipule scars more or less short. Leaves: blade pinnately veined, rarely with intermediates;
apex with tapering acumen, rarely caudate; intercostal venation generally scalariform
(excepting, notably, many species of sect. Richetioides); petiole frequently but not generally
geniculate. Inflorescences paniculate, terminal or generally 1axillary, singly if axillary,
doubly if terminal, branched, flowers secund. Flowers: buds lanceolate to spindleshaped,
occasionally subglobose (always so in sect. Shorea subsect. Barbata, sect. Isoptera, and
sect. Ovalis); sepals free down to receptacle, unequal or sometimes subequal, if unequal the
3 outer lobes longer than the 2 inner ones; stamens 10–105, in 1, 2 or 3 whorls, or irregular;
ovary usually pubescent, occasionally glabrous. Fruits: calyx lobes unequal or subequal,
expanded into a broad, saccate, spoonshaped and distinctly imbricate base appressed to
the nut; if unequal then all expanded into spatulate aliform lobes, with the 3 outermost lobes
longer and broader than the 2 inner ones, if subequal then all 5 lobes are short but the
outer 3 ones still somewhat thicker and longer than the inner 2 (all 5 equal, aliform, in S.
isoptera). Nuts relatively large, most usually pubescent, acute, free from fruit calyx base,
splitting irregularly at germination.
Distribution. About 196 species from S Asia through IndoBurma and Malesia to the
Philippines, Java and the Moluccas. One hundred thirty eight (138) species are known in
Borneo, of which 91 are endemic, and 130 occur in Sabah and Sarawak. There are few local
endemics, for instance between two major rivers, or confined to the Kinabalu Massif or the
Bau limestone; but 65 species, not all of them endemic in Borneo Is. are restricted within
Borneo to north of a line between Pontianak and Tidung in Kalimantan. Most of those are
191
soils specialists, especially of white or yellow sands. Of these, many do not extend into
Sabah beyond Sipitang and Beaufort districts in the southwest.
Ecology. The most frequently dominant genus in the emergent stratum of the zonal mixed
and upper dipterocarp forests on yellow/red soils, at altitudes below 1200 m in W Malesia
and the Philippines. A few, especially among the yellow meranti field group (sect.
Richetioides) reach maximum height in the main canopy. In Sarawak, and Brunei still, most
of the vast coastal peat swamp was dominated by the single species Shorea albida, the alan.
Most species have narrow edaphic ranges within the dipterocarp forests. A few in our
region, such as S. retusa, S. revoluta and S. stenoptera are confined to kerangas forest,
others such as S. inaequilateralis, S. macrantha, S. platycarpa, S. teysmanniana and S.
uliginosa to the peat swamp forests, and yet others including S. platyclados, S. flaviflora and
S. monticola to upper dipterocarp forests; while S. seminis and several others are exclusively
found on river banks and alluvium. Few species of dipterocarps are confined to limestone
habitat, though S. calcicola appears to be a good example in our area. Other species found
on limestone habitat either grow on acid organic soils overlying it on plateaux and are
kerangas species, e.g., S. multiflora, or are species of the seasonal tropics which penetrate
our area on dry limestone karst hills, e.g., S. guiso and H. plagata, or are species of mesic
lower limestone slopes that occur elsewhere on clay soils over basic rocks, e.g., S. isoptera
and S. virescens.
Uses. The most important timber genus in Sabah and Sarawak, albeit largely exhausted. The
four field groups produce distinctively different timbers (cf. PROSEA op. cit. 1993).
Selangan batu species yield heavy hard yellowbrown closegrained timbers suitable for
decking, garden furniture and construction. White meranti timbers are pale, of medium
density, and siliceous, suitable for veneer; yellow merantis yield a light hardwood pale
yellowbrown in colour, and red merantis a pink to crimson hardwood of light to moderately
heavy density that was until recently the most favoured tropical general utility hardwood on
international markets. Several largefruited species, mostly in botanical sect. Pachycarpae
of the red meranti field group, continue to be harvested, on account of the high content of
cocoa butter with an unusually high melting point in their fleshy cotyledons. This is mixed
with true cocoa butter in chocolates, and for lipsticks and polishes.
Notes. Shorea is a very large genus, but fairly clearly divided into subgroups/sections based
on characters of the stamens, and in many or some cases also the ovary, wood anatomy,
bark anatomy and morphology, bud and leaf morphology, and resin chemistry. Six rather
distinct field groups are also recognised, four of which occur in Sabah and Sarawak. Four of
these six field groups exactly correlate with four of the botanical groupings, one includes
two of them, but the largest field group, the red merantis, includes five botanical groups
each with its distinct floral and bark characters. Meijer (Act. Bot. Neerl. 12 (1963) 322)
recognised the red merantis as a botanical subgenus, Rubroshorea, solely on the basis of the
pink to crimson red colour of the wood and inner bark. There are, however, exceptions even
to this tenuous definition: Shorea guiso, in the selangan batu field group and botanical sect.
Shorea, has red inner bark and heartwood, while S. scaberrima, whose wood anatomy and
botanical grouping places it as a red meranti, has yellowbrown inner bark and heartwood.
There is no single qualitative character which binds the red merantis as a natural botanical
group.
A more serious problem has been revealed by molecular phylogenetic studies (Kamiya et al.
op. cit. 1998; Dayanandan et al. op. cit. 1999), which indicate that three botanical groups
within Shorea, namely Anthoshorea, Doona (of Sri Lanka) and Pentacme (IndoBurma and
192
the Philippines), are more closely related to Hopea and Neobalanocarpus than to other
Shorea; and that Parashorea is closer to the ancestral line of those other Shorea than they
are to that of Shorea sect. Anthoshorea, Doona, and Pentacme, and Hopea and
Neobalanocarpus. Yet no character has been identified which characterises each of these
two groups of Shorea sections separately. The two natural, phylogenetically separate major
groupings within Shorea, cannot therefore be given generic status, recognizable in the
herbarium or in the field. The alternative might be to give full generic status to the eleven
currently recognised botanical sections. This would be difficult in some cases, such as sect.
Rubella (red meranti), some of whose species share some characters with sect. Brachyptera
(red meranti) or sect. Anthoshorea (white meranti). Perhaps more importantly, noone who
is interested in these trees who is not a botanist would welcome the division of the red
merantis into 11 generic names. We, therefore, retain the commonly accepted definition and
name of Shorea, perhaps the best known indigenous tree genus in Sabah and Sarawak,
notwithstanding its paraphyletic origins.
Classification. In Borneo (Sabah and Sarawak), the genus Shorea as delimited in the
present study can be subdivided into 9 sections base on the following combinations of
vegetative and floral characters:
Key to Sections and Subsections of Shorea

1. Connectival appendage setose, stout; anthers 4loculed, apices of anterior locules often
also setose………………………………………………………………………..……2
sect. Shorea (selangan batu group)
Flowers: petals cream, often pink at base; stamens 20–60, not distinctly whorled,
filaments broad at base, gradually tapering, anthers 4loculed, each locule oblong,
often setose distally, earlike with protruding base and apex, connectival
appendage hardly exceeding the length of anther, more or less setose; ovary
shortly conical, pubescent, with short columnar style. Leaf blade with conspicuous,
flat or raised, or sometimes furrowed midrib above; intercostal venation densely
scalariform, usually sinuate and hardly elevated.
Connectival appendage generally sericeous or glabrous, if setose, then anthers glabrous
and 2loculed……………………………………………………………………………3
2. Flower buds ovoid to spindleshaped; petals narrow, falling separately; connectival

appendage sparsely setose…………………………….…sect. Shorea, subsect. Shorea
Flower buds globose; petals short, broadly oblong, connate at base on falling;
connectival appendage and anther locule apices densely setose………………………….
…………………………………………………………..sect. Shorea, subsect. Barbata
3. Fruit calyx lobes far exceeding ripe nut, spatulate, winglike but equal, patent; stamens
15, anthers subglobose, connectival appendage shorter than anther apices, stout, not
recurved…………………………………………………………………………………...
sect. Neohopea (selangan batu group)
Flowers small; buds globose; petals cream, broadly elliptic, falling separately;
stamens to 15, in 3 whorls, filaments short, stout, hardly tapering compressed,
anthers 4loculed, locules subglobose, connectival appendage short, glabrous;
ovary and stylopodium conical, style very short. Leaf blade with conspicuous,
raised midrib on both surfaces; intercostal venation densely scalariform, slender,
hardly raised below. Sapling leaves narrowly peltate.
193
Fruit calyx lobes unequal and spatulate, or equal and shorter than nut; stamens at least
30, anthers of other shapes, connectival appendage longer than anther apices or, if
shorter, then recurved………………………………………………………………..…4
4. Anthers 2loculed………………………………………………………………………..5
sect. Richetioides (yellow meranti group)
Flowers usually small; buds usually at most 5 × 3 mm, small; petals usually a
shade of yellow, narrow, strongly twisted at anthesis forming a distinct albeit small
cup at base enclosing the anthers, falling in rosette. Leaf blade with conspicuous,
flat or somewhat furrowed midrib above; intercostal venation more or less
reticulate or subscalariform with lesser crossconnecting veinlets. Opening leaves
violet or magenta; sapling leaves often peltate.
Anthers 4loculed.........................................................................................................…6
5. Flower buds larger, broadly ovoid; stamens at least 100, not whorled, filaments
gradually tapering, anthers lorate, connectival appendage shorter than anther apice,
densely setose; ovary without stylopodium, style short broad, distinctly truncate, stigma
………………………………………trifid……sect. Richetioides, subsect. Polyandrae
Flower buds usually smaller, ellipsoid or spindleshaped; stamens (10–)15, in (2–)3
whorls, filaments tapering abruptly medially, anthers subglobose to broadly ellipsoid,
connectival appendage aristate, much exceeding anther apice, sericeous or glabrous;
ovary with prominent stylopodium, style long, slender, stigma not trifid………………..
……………………………………………….sect. Richetioides, subsect. Richetioides
6. Anther locules linear to oblong; filaments lorate or tapering gradually; ovary without
stylopodium, style at least as long as ovary………………………………………….…7
Anther locules broadly oblong to subglobose; filaments tapering abruptly medially;
ovary usually with distinct stylopodium, style usually short………………………..…8
7. Connectival appendage at least 3x the length of anther, frequently sericeous; wood pale
yellow, vessel arrangement reticulate in TS, with silica…………………….……………
sect. Anthoshoreae (white meranti group)
Flowers usually large; buds ovoid; petals usually white, sometimes with pink
median suffusion, broadly elliptic to ovatelanceolate, contorted and imbricate at
anthesis forming a broad goblet enclosing the anthers, loosely connate on falling;
stamens 15–30, in 3 or more whorls, filaments broad at base, tapering gradually,
anthers 4loculed, locules narrowly oblong to lorate, connectival appendage
aristate, erect, at least 3x longer than anther; ovary ovoid, without distinct
stylopodium, style longer than ovary, filiform, stigma more or less distinctly trifid.
Leaf blade more or less bent backward along midrib; midrib generally obscure,
sunken above. Woodray cells containing silica crystals; vessels reticulately
arranged in TS of the wood.
Connectival appendage less than ¾x the length of anther, usually stout and glabrous;
wood pink, vessel arrangement loosely oblique in TS of wood, without silica..
sect. Rubella (red meranti group)
Flowers mediumsized; buds spindleshaped; petals cream suffused with pink,
lanceolate, contorted, imbricate at base forming a goblet enclosing the anthers,
connate on falling; stamens 15–50, in 3 (if 15) or indistinct whorls, filaments
lorate, gradually or abruptly tapering below anthers, anthers 4loculed, connectival
appendage short or exceeding the anther but of shorter length, tapering, glabrous,
becoming recurved at anthesis; ovary with or without stylopodium, style columnar,
194
stigma obscure. Leaf blade (in mature tree) more or less cream lepidote below;
midrib conspicuous (except in S. albida); intercostal venation densely scalariform,
hardly conspicuous. Wood without silica; vessels loosely diagonal to rays in TS.
8. Stamens 50–70, not in distinct whorls whorls, filaments very long, filiform, connectival
appendage vestigial…………………………………………………………………..…...
sect. Ovalis (red meranti group)
Stipules and bracts not early caducous. Flower buds broadly ovoid; petals as in
sect. Rubella; anthers subglobose, 4loculed; ovary and stylopodium narrowly
conical, style short. Leaf blade with obscure, sunken midrib above; intercostal
venation scalariform.
Stamens less than 30, in 3 whorls, filaments compressed and broad at base, connectival
appendage aristate……………………………………..……………………………..…9
9. Connectival appendage at most as long as anthers, becoming more or less reflexed at

anthesis; filaments applanate, tapering from the base to apex………………………....10
sect. Mutica (red meranti group)
Stipules and bracts early caducous (except in S. quadrinervis). Flower buds ovoid;
petals as in sect. Rubella; stamens to 15, in 3 whorls, anthers 4loculed, locules
globose; ovary with distinct stylopodium, style shorter or only slightly longer than
ovary. Leaf blade generally with obscure or very slender midrib above; intercostal
venation scalariform.
Connectival appendage at least 1½x the length of anther, not becoming reflexed at
anthesis, filaments broad and flat at base, tapering abruptly medially and filiform below
anthers……………………………………………………………………………..…11
10. Bark surface mostly becoming deeply and persistently vsection fissured, rotting off
only in large trees. Fruit calyx lobes not auriculate………sect. Mutica, subsect. Mutica
Bark surface generally at first remaining smooth, then, after passing through a generally
ephemeral shallowly vsection period, becoming chunkily flaky. Fruit calyx lobes
auriculate at base……………………………..………sect. Mutica, subsect. Auriculatae
11. Base of filaments connate at base but otherwise free; ovary ovoid without, or pyriform
with, a stylopodium, style filiform, prominent…………………………………………...
sect. Brachypterae (red meranti group)
Inflorescences lax, with long branches; stipules and bracts at times somewhat
persistent. Flower buds ovoid; petals as in sect. Rubella; stamens 15 or 24–28, in 3
whorls or indistinctly so, anthers 4loculed, locules globose or broadly ellipsoid,
connectival appendage 1½–3½x as long as anthers, slender, erect; ovary with
distinct stylopodium, together pyriform, or ovoid without stylopodium but with
puberulent style base. Leaf blade with conspicuous or obscure and shallowly
sunken midrib above; intercostal venation scalariform. Bark cracking and
becoming flaky in clean more or less flat thin pieces.
195
Margin of flattened bases of outer filaments united to form a tube around ovary; ovary
ovoid, small, stylopodium tapering into style, style and stylopodium spindleshaped,
tapering at base and apex, or stylopodium indistinct and style filiform, more than 2½x
the length of ovary………………………………………………………………………..
sect. Pachycarpae (red meranti group)
Stipules and bracts subpersistent, usually relatively large. Flower buds ovoid to
fusiform; petals as in sect. Rubella; stamens to 15, in 3 whorls, filaments lorate,
abruptly tapering and filiform below anthers; anthers subglobose to broadly
ellipsoid, connectival appendage at least 2x as long as anther locules; ovary small,
ovoid, with glabrous or glabrescent stylopodium. Bark remaining smooth, hoop
marked, pale greyish brown, eventually often becoming irregularly flaked and
scrollmarked.
Key to Shorea species

1. Connectival appendage of stamen setose, stout; anthers 4loculed, apices of anterior

locules often also setose…………………………………………………………..…2
Connectival appendage generally sericeous or glabrous but, if setose, then anthers
glabrous and 2loculed...........................................................................................…27
2. Flower buds ovoid to spindleshaped, petals narrow, falling separately, connective

appendage sparsely setose……………………………………………………………3
Flower buds globose, petals short, broadly oblong, connate at base on falling,
connective appendages and apices of anther sacs densely setose………………..…22
3. Connectival appendage typically with 1–4 (a few with to 8) bristles, stamens

otherwise glabrous……………………………………………………………..……4
Connectival appendage with at least 5 bristles, shoulder of anther apices setose…....8
4. Lateral veins at least 15 pairs………………………………………….…49. S. guiso

Lateral veins at most 14 pairs…………………………………………………...……5
5. Fruit calyx lobes short, subequal………………………………..…50. S. havilandii

Fruit calyx lobes unequal, 3 larger ones winglike………………………………...…6
6. Stamens at least 40………………………………………………18. S. brunnescens

Stamens 20–30…………………………………………………………………….…7
7. Lateral veins 10–12 pairs, dense. In mixed dipterocarp forest…..114. S. scrobiculata

Lateral veins to 9 pairs, distant. In forest on limestone karst hills…….20. S. calcicola
8. Fruit calyx lobes subequal, shorter than nut…………………………………….…9

Fruit calyx lobes unequal, winglike, longer than the nut……………………….…10
9. Stamens c. 55; ripe nut to 5 × 5 cm…………………………….….46. S. geniculata

Stamens 30–40; nut at most 2 cm diameter……………………….…115. S. seminis
196
10. Staminal filaments setose along their distal margins……………………………..…11

Staminal filaments glabrous…………………………………………………………18
11. Leaf blade not eventually pale lepidote below.............................…45. S. foxworthyi

Leaf blade of mature tree pale lepidote below....................................................…12
12. Twigs at first compressed…………………………………………………………..13

Twigs entirely terete…………………………………………………………….…14
13. Tomentum on twig and petiole yellowish brown scabrid; petiole less than 1.5 cm
long ................................................................................................…35. S. exelliptica
Tomentum on twig and petiole buff, even; petiole at least 1.5 cm long……………….
……………………………………………………………….56. S. inappendiculata
14. Leaf blade falcate, base asymmetric; lateral veins slender below..........................…15
Leaf blade more or less symmetric, base equal; lateral veins prominent below…….16
15. Stamens 33–44…………………………………………………….…38. S. falcifera

Stamens 25–32…………………………………………………….…74. S. materialis
16. Stamens 25–33…………………………………………………....12. S. atrinervosa

Stamens 35–46…………………………………………………………………..…17
17. Petiole at least 3.5 cm long…………………………………………..…27. S. crassa

Petiole at most 2 cm long…………………………………………..…83. S. obscura
18. Stamens at least 40…………………………………………………………………19

Stamens at most 37……………………………………………………………….…20
19. Leaf with more or less equal base, not cream lepidote below; lateral veins 11–15
pairs, stout below…………………………………………………69. S. lunduensis
Leaf falcate, cream lepidote below (mature trees); lateral veins 8–12 pairs, slender
below……………………………………....39. S. falciferoides (subsp. glaucescens)
20. Lateral veins 16–24 pairs, puberulent below…………………………121. S. superba

Lateral veins at most 16 pairs, not puberulent below………………………….…..21
21. Lateral veins 11–16 pairs; domatia minute…………………………53. S. hypoleuca

Lateral veins at most 12 pairs; domatia prominent, porelike………33. S. domatiosa
22. Stamens at least 45………………………………………………………63. S. laevis

Stamens at most 35…………………………………………………………………23
23. Leaf blade less than 8 cm long; lateral veins 6–7 pairs, not sunken above…………24
Leaf blade at least 10 cm long; lateral veins at least 8 pairs, sunken above or not….25
24. Fruit calyx lobes subequal, shorter than nut…………………………..…11. S. asahi

Fruit calyx lobes unequal, winglike, longer than nut……………….…77. S. micans
197
25. Fruit calyx lobes unequal, winglike, longer than nut…………...75. S. maxwelliana
Fruit calyx lobes subequal, shorter than nut……………………………………..…26
26. Leaf blade coriaceous; lateral veins 5–6 pairs, not sunken above……62. S. ladiana
Leaf blade thintextured; lateral veins at least 8 pairs, slightly sunken above…………
…………………………………………………………………………16. S. biawak
27. Fruit calyx lobes far exceeding the length of ripe nut, spatulate, winglike but equal,
patent; stamens 15, anthers subglobose, connectival appendage not exceeding the
anther apice, stout……………..………………………………………58. S. isoptera
Fruit calyx lobes either unequal and spathulate, or equal and shorter than nut;
connectival appendage as long as or longer than the anther apice or, if shorter, either
recurved or stamens at least 20………………………………………………….…28
28. Anthers 2loculed..................................................................................................…29

Anthers 4loculed..................................................................................................…57
29. Flower buds large, broadly ovoid; stamens at least 100, filaments gradually tapering,
anthers lorate, connectival appendage shorter than anther apice, densely setose; ovary
ovoid, without stylopodium, style short, broad, distinctly trifurcate..99. S. polyandra
Flower buds usually small, ellipsoid or spindleshaped; stamens (10–)15, in (2–)3
whorls; anther locules subglobose to broadly ellipsoid, tapering abruptly medially,
connectival appendage aristate, much exceeding anther apice, sericeous or glabrous;
ovary with prominent stylopodium, with long slender style, stigma not trifid……30
30. Fruit calyx lobes shorter than nut, subequal……………………………………...…31

Fruit calyx lobes longer than nut, winglike, unequal………………………………47
31. Nut glabrous, shiny, drying black…………………………………………………32

Nut pubescent, drying dull, pale………………………………………………..…33
32. Petiole at most 1.2 cm long; ovary glabrous; stamens c. 15…………67. S. longiflora
Petiole at least 1.8 cm long; ovary puberulent; stamens c. 10…..71. S. macrobalanos
33. Leaf blade less than twice as long as wide…………………………………………34

Leaf blade more than twice as long as wide.…………………………………….…39
34. Stamens 16–17…………………………………………………….…106. S. richetia

Stamens 10–15………………………………………………………………………35
35. Leaf venation persistently cream puberulent below................................…65. S. laxa

Leaf entirely glabrous………………………………………………………………36
36. Mature tree leaf deeply peltate; understorey tree with smooth bark……94. S. peltata
Juvenile leaf peltate only; canopy tree with flaky bark…………………………..…37
37. Leaf blade 7–12 cm long; lateral veins 5–7 pairs; nut to 3 × 1.3 cm, drying mauve
puberulent...........................................................................…14. S. balanocarpoides
198
Leaf blade at most 8 cm long; lateral veins at least 7 pairs; nut to 2 × 1.2 cm, cream
buff tomentose (rarely glabrous)………………………………………………….…38
38. Lateral veins 8–10 pairs, hardly raised below; blade base not decurrent; petiole
drying rugulose; ovary without distinct stylopodium…………….…80. S. multiflora
Lateral veins 7–9 pairs, slender but distinctly raised below; blade base shortly
decurrent; petiole drying smooth, black; ovary with distinct stylopodium…………….
………………………………………………………………………92. S. patoiensis
39. Leaf blade greyish brown scabridpubescent below..…………..…57. S. induplicata

Leaf glabrous, or venation alone pubescent below.............................................…40
40. Midrib and lateral veins shallowly furrowed above……….….120. S. subcylindrica

Midrib and lateral veins not furrowed above…………………………………….…41
41. Leaf venation puberulent below...…………………………………………………42

Leaf venation entirely glabrous……………………………………………………43
42. Blade base equal; midrib prominently narrowly furrowed above…...82. S. obovoidea
Blade base unequal; midrib not narrowly furrowed above………….…22. S. chaiana
43. Petiole persistently pale cream, puberulent or glabrous......................................…44

Petiole drying greyish brown or black, glabrous……………………………………45
44. Petiole at least 1.9 cm long; blade greyish matte below; stamens c. 10; fruit calyx
lobes subpatent…………………………………………………….…23. S. collaris
Petiole less than 1.5 cm long; blade pale tawny below; stamens c. 15; fruit calyx
lobes clasping the nut…………………………………………….…9. S. angustifolia
45. Twig slender, pale; leaf blade dull below; distal end of petiole drying cream………...
………………………………………………………………123. S. tenuiramulosa
Twig stout, drying dark; leaf blade shiny below; petiole drying entirely black…….46
46. Leaf blade coriaceous; midrib flat above………………………..…13. S. bakoensis

Leaf blade thintextured; midrib raised above……………….…130. S. xanthophylla
47. Petiole at least 1.9 cm long; lateral veins at least 10 pairs……………………….…48

Petiole at most 1.8 cm long or, if longer; lateral veins less than 9 pairs……….……49
48. Petiole 3–4 mm diameter, stout; blade thickly coriaceous………………54. S. iliasii

Petiole less than 2 mm diameter; blade chartaceous…………..….37. S. faguetioides
49. Lateral veins at least 14 pairs………………………………………...…6. S. alutacea

Lateral veins at most 12 pairs…………………………………………………..…50
50. Leaf venation persistently tomentose below…………………………………….…51

Leaf venation glabrescent or glabrous below………………………………………53
51. Leaf blade more or less densely pale greyish green puberulent below………………...
……………………………………………………………………68. S. longisperma
199
Leaf blade glabrous; venation scabridpuberulent below……………………….….52
52. Leaf margin revolute; lateral veins 9–12 pairs; tomentum scabrid…………………….
………………………………………………………………..1. S. acuminatissima
Leaf margin not revolute; lateral veins 7–9 pairs; tomentum even…….47. S. gibbosa
53. Lateral veins very slender, hardly raised, with distinct shorter intermediates; midrib
drying reddish or blackish, frequently with paired glabrous porelike domatia at base
…………………………………………………………………….…52. S. hopeifolia
Lateral veins stout, without intermediates; midrib not drying darker than blade,
distinctly raised, without domatia…………………………………………………54
54. Petiole 1.7–2.2 cm long……………………………………………60. S. kudatensis

Petiole at most 1.6 cm long……………………………………………………….…55
55. Lateral veins 5–7 pairs…………………………………………..…29. S. cuspidata

Lateral veins at least 8 pairs……………………………………………………..…56
56. Lateral veins c. 8 pairs, prominently raised below; leaf margin narrowly revolute…...
…………………………………………………………………..79. S. mujongensis
Lateral veins 9–12 pairs, raised but not prominent below; leaf margin not revolute….
……………………………………………………………………36. S. faguetiana
57. Anther locules linear to oblong; filaments lorate or tapering gradually; ovary without
stylopodium, style at least as long as ovary…………………………………….…58
Anther locules broadly oblong to subglobose; filaments tapering abruptly medially;
ovary usually with distinct stylopodium………………………………………….…72
58. Connectival appendage at least 3x the length of anther, frequently sericeous; wood
pale yellow, vessel arrangement reticulate in TS, with silica…………………….…59
Connectival appendage less than ¾x the length of anther, usually stout, glabrous;
wood pink, vessel arrangement loosely oblique in TS; silica absent……………….69
59. Stamens at least 17……………………………………………………………..…60

Stamens to 15………………………………………………………………………61
60. Stamens c. 17; blade undersurface pale pink lepidote (mature trees); lateral veins 20–
24 pairs.………………………………………………………………31. S. dealbata
Stamens c. 25; blade not pale lepidote below; lateral veins at most 14 pairs………….
……………………………………………………………………..48. S. gratissima
61. Leaf venation persistently tomentose below.........................................................…62

Leaf venation early glabrescent below...................................................................…64
62. Tomentum red to creambrown; leaf blade oblongovate, bright yellow lepidote
below………………………………………………………………....84. S. ochracea
Tomentum dull greyish to tawny brown; blade obovate or ovateoblong not pale
lepidote below………………………………………………………………..……63
200
63. Leaf blade obovate; lateral veins at most 17 pairs; twig at first compressed…………..
……………………………………………………………………127. S. virescens
Leaf blade ovateoblong; lateral veins 17–26 pairs; twig terete……64. S. lamellata
64. Lateral veins at least 15 pairs; twigs at first compressed…………………………65

Lateral veins at most 15 pairs or, if more, then twigs terete………………………66
65. Fruit pedicel c. 1 cm long, c. 5 mm diameter; receptacle obconical; longer fruit calyx
lobes to 18 cm long…………………………………………..…122. S. symingtonii
Fruit pedicel at most 2 mm long and diameter; receptacle cupshaped, abruptly
constricted at base; longer fruit calyx lobes at most 13 cm long…..….25. S. cordata
66. Twig apices more or less compressed; leaf blade subchartaceous, drying chocolate
brown………………………………………………………….…17. S. bracteolata
Twig terete; leaf blade coriaceous, drying tawny to pale yellowish brown……..…67
67. Base of leaf blade cuneate………………………………………….103. S. resinosa

Base of leaf blade obtuse to subcordate………………………………………….…68
68. Base of fruit impressed; flower bud to 5 × 2.5 mm…………………..…3. S. agamii

Base of fruit tapering; flower bud to 9 × 5 mm…………………...……24. S. confusa
69. Midrib obscure above; twigs compressed……………………………..…4. S. albida

Midrib conspicuous above; twigs terete……………………………………………70
70. Stamens 25………………………………………………………….…32. S. dispar

Stamens 15–20……………………………………………………………………....71
71. Stamens 15…………………………………………………………..…108. S. rubella

Stamens 19–20……………………………………………………..…34. S. elliptica
72. Stamens 50–70, connectival appendage vestigial, filaments very long, filiform……...
…………………………………………………………………………85. S. ovalis
Stamens less than 30, connectival appendage aristate, filaments compressed and
broad at base……………………………………………………………………..…73
73. Connectival appendage aristate, at least 1½x the length of anther, not reflexed,
filaments broad and flat at base, tapering abruptly and filiform below anthers……..74
Connectival appendage at most of the same length as anther, generally becoming
reflexed at anthesis; filaments flat, tapering from base to apex……………………106
74. Filaments broad at base, tapering abruptly medially and filiform below anthers,
connate at base but otherwise free; ovary ovoid without, or pyriform with, a
stylopodium, style filiform, prominent…………………………………………..…75
Margins of flattened bases of outer filaments united to form a tube round the ovary;
stylopodium tapering into style, style and stylopodium spindleshaped, tapering at
base and apex, or stylopodium indistinct and style more than 2½x the length of
ovary, filiform…………………………………………………………………….…96
201
75. Stamens 24–28………………………………………………….…117. S. smithiana

Stamens 15……………………………………………………………………..…76
76. Ovary without distinct stylopodium, ovoid, style at least as long as ovary, generally
puberulent in the basal half…………………………………………………………77
Ovary and stylopodium distinctly pyriform, style shorter than both, glabrous…….89
77. Leaf blade falcate, base distinctly unequal……………….…55. S. inaequilateralis

Leaf and base more or less symmetrical……………………………………………78
78. Leaf blade below, petiole and young twig scabridtomentose………………….…79

Leaf blade below, petiole and young twig shortly evenly tomentose or glabrous…..81
79. Leaf blade coriaceous, margin revolute, creamlepidote below (mature trees); longer
fruit calyx lobes to 6.5 × 2.5 cm……………………………………43. S. flemmichii
Leaf blade more or less chartaceous, margin not revolute, not lepidote below; longer
fruit calyx lobes to 14 × 2.5 cm……………………………………………………80
80. Leaf blade concave; tomentum drying paler than blade undersurface….…5. S. almon
Leaf blade flat; tomentum drying same colour or darker than blade undersurface……
…………………………………………………………………91. S. parvistipulata
81. Lateral veins at least 13 pairs, very slender, hardly raised below; intercostal venation
not raised, densely scalariform…………………………………………………..…82
Lateral veins, if more than 13 pairs, prominent below; intercostal venation not as
above………………………………………………………………………………83
82. Leaf blade 10–15 × 5–8 cm, base subpeltate discernable as a rib over the base of the
midrib (mature trees); stipule scars cuneate………………………..…26. S. coriacea
Leaf blade 6–10 × 3–5 cm, base not subpeltate; stipule scars long, often amplexicaul
…………………………………………………………………..…126. S. venulosa
83. Lateral veins at least 16 pairs; blade pinkish cream lepidote below…..128. S. waltoni
Lateral veins at most 16 pairs; blade not pale lepidote below…………………..…84
84. Petiole 4–6 cm long………………………………………………87. S. pachyphylla

Petiole at most 3.5 cm long…………………………………………………………85
85. Connectival appendage scarious……………………………………93. S. pauciflora

Connectival appendage glabrous…………………………………………………....86
86. Leaf blade with to 3 pairs of pale scalelike domatia at base.…….…59. S. johorensis
Leaves without scalelike domatia…………………………………………………87
87. Leaf thickly coriaceous; style glabrous. In karapa forest, at 800–1200 m altitudes…...
…………………………………………………………………………21. S. carapae
Leaf thinly coriaceous; style densely pubescent at least in basal ¼. In flood plain and
mixed dipterocarp forest, at altitudes to 600 m…………………………………..…88
202
88. Fruit calyx lobes shorter than nut………………………………...88. S. palembanica

Fruit calyx lobes winglike, much longer than nut………………..…8. S. andulensis
89. Twigs compressed; midrib sharply acute below; lateral veins hardly raised below…...
……………………………………………………………………..98. S. platyclados
Twigs terete; midrib terete but prominent below; lateral veins distinctly raised…....90
90. Fruit calyx lobes less than 2x the length of nut……………………………………...91

Fruit calyx lobes at least 3x the length of nut…………………………………….…92
91. Ovary and stylopodium hardly distinguishable, filiform towards apex, stouter below,
frequently further swollen in the basal half, style short; ripe nut to 5 × 2.5 cm……….
…………………………………………………………………..112. S. scaberrima
Ovary and stylopodium ovoid, crowned by a filiform style equal in length; ripe nut to
3.5 × 1.5 cm………………………………………………………………40. S. fallax
92. Ovary and stylopodium densely pubescent but for the short style apex……………….
……………………………………………………………………..101. S. pubistyla
Ovary and stylopodium puberulent, glabrescent to glabrous or if puberulent, then
style equal in length…………………………………………………………………93
93. Connectival appendage exceeding the style at anthesis, very slender, crisped; leaf
blade bullate between intercostal veins; venation below, petiole and twig scabrid
tomentose…………………………………………………………..…19. S. bullata
Connectival appendage not exceeding style at anthesis, awnlike; leaf blade flat,
evenly tomentose or glabrous…………………………………………………..…94
94. Ovary and stylopodium glabrescent, c. 2x the length of style; anther locules large,
strongly tapering to and fused at apex…………………………….…42. S. flaviflora
Ovary and stylopodium puberulent, equal to style length; anther locules not large,
hardly tapering, free………………………………………………………………..95
95. Lateral veins 13–19 pairs; blade golden lepidote (mature trees) below, without
domatia..……………………………………………………………78. S. monticola
Lateral veins 6–8 pairs; blade glabrous, with porelike domatia….…..61. S. kunstleri
96. Stipule scars amplexicaul……………………………………………………………97

Stipule scars prominent but not amplexicaul…………………………………..…104
97. Leaf blade densely goldenbrown tomentose below, with axillary domatia.…………..
………………………………………………………………………….95. S. pilosa
Leaf blade sparsely shorttomentose or glabrous below………………………….…98
98. Stipules with cordate subequal base………………………………...118. S. splendida

Stipules not cordate at base……………………………………………………...…99
99. Leaf base peltate…………………………………………………..…129. S. woodii

Leaf base cordate, obtuse or cuneate………………………………………….…100
203
100. Lateral veins 11–20 pairs; nut to 6 × 4 cm; longer fruit calyx lobes to 11 cm long
…………………………………………………………………………………..…101
Lateral veins 9–12 pairs; nut to 3.7 × 2.5 cm; longer fruit calyx lobes exceeding 15
cm long……………………………………………………………………………102
101. Inflorescences to 35 cm long, axillary on modified sections of the twig with short
internodes and rudimentary or aborted leaves; stipules to 2 cm long, subrevolute……
…………………………………………………………………….119. S. stenoptera
Inflorescences to 20 cm long, in axils of normal leaves on normal twig; stipules to 5
cm long, flat………………………………………………………72. S. macrophylla
102. Leaf blade at least 25 cm long………………………………….…100. S. praestans

Leaf blade at most 21 cm long……………………………………………………103
103. Leaf blade broadly ovate to suborbicular, 10–13 cm wide………107. S. rotundifolia

Leaf blade elliptic, 5–8 cm wide……………………………..…7. S. amplexicaulis
104. Leaf base cordate; undersurface densely persistently golden pubescent………………

………………………………………………………………….76. S. mecistopteryx
Leaf base not cordate; tomentum caducous or not golden, or leaf glabrous……105
105. Stipule scars short, horizontal or somewhat ascending; lateral veins 11–14 pairs…….
……………………………………………………………………..15. S. beccariana
Stipule scars falcate, descending; lateral veins mostly 14–19 pairs…...96. S. pinanga
106. Fruit calyx lobes auriculate at base…………………………………………….…107

Fruit calyx lobes not auriculate at base……………………………………………112
107. Leaf blade narrowly oblong; lateral veins at least 19 pairs…………………….…108

Leaf blade elliptic, oblong to lanceolate; lateral veins at most 15 pairs………...…110
108. Leaf blade concave boatshaped; lateral veins sunken above…….…116. S. slootenii
Leaf blade flat; lateral veins prominent above……………………………………109
109. Leaf blade sparsely tufted pubescent or glabrescent below; lateral veins 24–28 pairs
………………………………………………………………….…81. S. myrionerva
Leaf blade densely shortly persistently scabridpuberulent below; lateral veins 19–25
pairs……………………………………………………………..…..111. S. sagittata
110. Leaf blade 14–26 × 6.5–12 cm, thickly coriaceous…………………….…2. S. acuta

Leaf blade 7–16 × 2.2–6 cm, thinly coriaceous………………………………..…111
111. Leaf blade lustrous below; lateral veins prominent………………73. S. macroptera

Leaf blade dull below; lateral veins hardly raised, very slender……41. S. ferruginea
112. Stipules exceeding 20 mm long, broad, boatshaped, prominently ribbed, coriaceous,

not early caducous……………………………………………102. S. quadrinervis
Stipules at most 19 mm long, chartaceous, flat, not prominently ribbed, early
caducous………………………………………………………………………..113
204
113. Flower buds at least 14 mm long…………………………………………………114

Flower buds at most 10 mm long…………………………………………………115
114. Petiole at most 0.6 cm long; leaf base cordate, unequal………..…70. S. macrantha
Petiole 0.6–1.2 cm long; leaf base obtuse, equal………………………………………
………………………………………………51. S. hemsleyana (subsp. grandiflora)
115. Leaf blade glabrous below………………………………………………………116

Leaf blade sparsely or densely tomentose or pale lepidote below......................…118
116. Apex of leaf blade retuse…………………………………………..…104. S. retusa

Apex of leaf blade acuminate…………………………………………………..…117
117. Lateral veins 8–9 pairs, with prominent axillary porelike domatia…………………...
………………………………………………………………….44. S. foraminifera
Lateral veins 11–12 pairs, with at most a few pairs of small domatia at the midrib
base………………………………………………………..…124. S. teysmanniana
118. Leaf with at least 14 pairs of lateral veins or, if sometimes with as few as 11, greyish
brown scabrid, cream pubescent or lepidote below, or with pale scalelike domatia up
sides of midrib……………………………………………………………….…119
Leaf with at most 13 pairs of lateral veins, or if tomentum as above, then with at
most 10 pairs of lateral veins; domatia and indumentum not as above……………125
119. Leaf blade evenly pinkish brown velutinous below; lateral veins 20–25 pairs, with
prominent intermediates……………………………………….…10. S. argentifolia
Leaf blade not pinkish brown velutinous below; lateral veins less than 21 pairs,
without intermediates…………………………………………………………..…120
120. Leaf venation, petiole and twigs densely scabridtomentose………………………121

Leaf venation, petiole and twigs sparsely or evenly tomentose………………..…124
121. Leaf blade cream lepidote below...................................................…89. S. pallidifolia

Leaf blade not lepidote below...............................................................................…122
122. Leaf blade ovateelliptic, finely scabrid; lateral veins slender below………………….
……………………………………………………………………30. S. dasyphylla
Leaf blade oblongelliptic, oblongobovate, or oblongovate; lateral veins stoutly
prominent below.……………………………………………………………….…123
123. Leaf blade concave, chartaceous; petiole 2.2–3.2 cm long…………125. S. uliginosa

Leaf blade flat, thickly coriaceous; petiole 1.3–2.3 cm long………......110. S. rugosa
124. Leaf blade (mature tree) pinkish buff puberulent below; lateral veins 12–15 pairs…...
……………………………………………………………………….66. S. leprosula
Leaf blade (mature tree) sparsely scabrid below; lateral veins 16–20 pairs…………...
…………………………………………………………………….97. S. platycarpa
125. Leaf blade narrowly ovate to lanceolate, cream to pink lepidote below (mature tree);
lateral veins slender, hardly raised below……………28. S. curtisii (subsp. curtisii)
205
Leaf blade elliptic or broadly ovate, not cream or pink lepidote below; lateral veins
prominent……………………………………………………………………….…126
126. Leaf blade shiny below..………………………………………………………….127

Leaf blade dull below.........................................................................................…128
127. Leaf blade elliptic, 5–9 × 3–5 cm, margin hardly or not revolute, apex shortly
acuminate or frequently retuse....................................................….113. S. scabrida
Leaf blade broadly ovate, 10–15 × 5.5–10 cm, margin prominently revolute, apex
prominently acuminate……………………………………………....105. S. revoluta
128. Leaf blade broadly ovate; lateral veins 8–10 pairs…………………….…86. S. ovata
Leaf blade ovate to elliptic; lateral veins c. 11 pairs…………………………….…129
129. Leaf blade below, petiole and twigs densely deep rufousbrown scabridpuberulent…
…………………………………………………………………………109. S. rubra
Leaf blade below, petiole and twigs densely evenly pale brown puberulent…………..
………………………………………………………………………90. S. parvifolia
Key to Shorea species

(Caution: this key is only reliable for leaves from mature trees; juveniles have smooth bark;
in saplings, leaves often lack pale lepidote blade undersurfaces, blades are larger, narrower,
with more prominent acumens and frequently more veins, petioles are longer, more slender,
frequently geniculate)
1. Wood hard, dense, ray ends glistening on tangential surface (observable with the aid
of hand lens)…………………………………………………………………………2
Wood relatively soft, ray ends not glistening on tangential surface…………………5
2. Inner bark and heartwood orange red to meatred; leaf base sometimes prominently
unequal………………………………………………………………………………3
Inner bark and heartwood creambrown or, if reddish, inner bark evenly textured;
leaf base subequal to equal…………………………………………………………..7
Selangan batu (preferred name); Sabah and Sarawak—selangan (lighter wood
species, Malay), tekam (Iban) (sect. Shorea and sect. Neohopea)
Buttresses prominent, narrow, sharp (except in S. geniculata, S. isoptera, S.
superba). Bark thin, hard, dull greyish brown to fawn, occasionally tawny or
chocolate, indistinctly smalllenticellate, becoming longitudinally cracked and
oblong flaky, the flakes thin, with flat clean surfaces (both sides); inner bark
relatively thin, closetextured, hard, homogeneous yellowish brown paling
towards cambium (rarely pink to red). Sapwood closetextured, glistening on
tangential surface and readily exuding dammar; heartwood chocolatebrown to
dark reddish brown; dammar frequent on bark as cream to cream yellow opaque
incrustations or smears.
206
3. Inner bark dense, homogeneous. Lateral veins typically at least 15 pairs……………..

…………………………………………………………………49. S. guiso (in part)
Inner bark fibrous. Lateral veins at most 13 pairs……………………………….…4
4. Leaf base equal…………………………………………….61. S. kunstleri (in part)

Leaf base prominently unequal..……………………55. S. inaequilateralis (in part)
5. Inner bark distinctly laminated………………………………………………….…35

White meranti; Sabah—melapi (preferred name); Sarawak—raruk (Iban) (sect.
Anthoshorea)
Buttresses more or less prominent, stout. Bark pale or dark, thick irregular
section cracked, eventually rotting off in chunky flakes, the lower and
subsequent surfaces rough, powdery, rarely thinly flaky; inner bark laminated
cream to yellow to orangebrown, thick, fibrous. Sapwood pale yellow, coarse;
heartwood indistinct; dammar incrustations pale to bright yellow. Leaf midrib
obscure, more or less sunken above.
Inner bark not distinctly laminated…………………………………………………6
6. Dammar dark brown to black on fresh surfaces, often as coxcombs on bark, or as

copious incrustations. Bark tawny (except in S. acuminatissima), becoming vertically
cleanly cracked and thinly oblong flaky, the flakes with clean surfaces; inner bark
fibrous, tawny brown with a greenish tinge at the cambium…………………….....45
Yellow meranti; Sabah—seraya kuning, yellow seraya (preferred names);
Sarawak—damar hitam (preferred name), lun (light; Iban), merakunyit (heavy;
Iban) (sect. Richetioides)
Buttresses small or prominent, stout. Sapwood dull yellow to lemon yellow;
heartwood pale tawny brown. Twigs commonly striated or rugulose. Leaf
generally drying yellowish brown to greenish brown; intercostal venation
reticulate or, if subscalariform, with distinct crossconnections, generally drying
darker than blade. Sapling leaves often peltate.
Dammar incrustations opaque, cream to yellow. Bark variously smooth, fissured or
cleanly flaky, greyish to pinkish or chocolatebrown; inner bark meatred or plum to
crimsonbrown, rarely coffeecoloured……………………………………….…76
Red meranti; Sabah—seraya, red seraya (preferred name); Sarawak—lop (Iban),
perawan (Iban) (sect. Brachypterae, sect. Mutica, sect. Ovalis, sect.
Pachycarpae and sect. Rubella)
Buttresses generally prominent, stout. Sapwood cream to pale yellow. Midrib
usually evident, sometimes obscure sunken above; blade drying grey to pink
brown.
7. Bark surface covered with a superficial but distinct pattern of small vsection fissures
and sharp narrow ridges, pale grey, eventually patchily cracked and peeling…………
………………………………………………………………………..16. S. biawak
Bark at first smooth, becoming flaky, yellow to chocolatebrown……………….…8
8. Leaf blade white or cream lepidote below.............................................................…9

Leaf not pale lepidote below……………………………………………………..…24
9. Petiole more than 3.5 cm long…………………………………………………..…10

Petiole less than 3.5 cm long…………………………………………………….…11
207
10. Leaf blade broadly ovate to orbicular; lateral veins drying black; petiole white
lepidote………………………………………………………….…46. S. geniculata
Leaf blade narrowly ovate to elliptic; lateral veins not drying black; petiole cream
pubescent..…………………………………………………………..…27. S. crassa
11. Midrib persistently or caducous pubescent below..............................................…12

Midrib glabrous below………………………………………………………………15
12. Lateral veins less than 9 pairs……………………………………..…83. S. obscura

Lateral veins more than 9 pairs………………………………………………….…13
13. Venation scabridtomentose below..............................................…35. S. exelliptica

Venation evenly tomentose below........................................................................…14
14. Lateral veins at most 15 pairs……………………………….115. S. seminis (in part)

Lateral veins at least 16 pairs………………………………….….121. S. superba
15. Lateral veins at least 15 pairs…………………………………………………….…16

Lateral veins at most 14 pairs………………………………...............................…17
16. Leaf blade cream lepidote below; lateral veins drying cream or yellowish brown
below; petiole 1–1.6 cm long………………………….114. S. scrobiculata (in part)
Leaf blade white lepidote below or drying greyish brown; lateral veins drying black
or greyish brown below; petiole 1.5–2.6 cm long………..…56. S. inappendiculata
17. Petiole 0.7–1 cm long………………………………………..…75. S. maxwelliana

Petiole at least 1.1 cm long……………………………………………………….…18
18. Leaf venation drying black below………………………………………………..…19

Leaf venation drying cream to pale brown below..…………………………………21
19. Leaf blade coriaceous; lateral veins stout, prominent below; petiole stout……………
…………………………………………………………………….12. S. atrinervosa
Leaf blade chartaceous to thinly coriaceous; lateral veins slender, hardly raised;
petiole slender…………………………………………………………………..…20
20. Petiole 1–1.5 cm long; leaf blade falcate; lateral veins without domatia…63. S. laevis
Petiole 1.5–2.5 cm long; leaf blade broadly ovate, subequal; lateral veins with pore
like domatia……………………………………………………..…33. S. domatiosa
21. Leaf lateral veins 11–16 pairs, stout, prominent below…………..…53. S. hypoleuca
Leaf lateral veins at most 12 pairs, elevated but not prominent below……………..22
22. Leaf base unequal; blade subfalcate; petiole and midrib cream lepidote; twig
glabrescent. In kerangas forest……………………………………74. S. materialis
Leaf base subequal; blade not falcate; petiole and twig cream puberulent. In mixed
dipterocarp forest……………………………………………………………………23
23. Leaf blade 6.5–12 × 2.5–5 cm. In forest on sandy soils west of Batang Lupar
(Sarawak)…………………………………………………………..…38. S. falcifera
208
Leaf blade 10–18 × 4.5–8 cm. In forest on clay soils east of Batang Lupar…………...
…………………………………………….39. S. falciferoides (subsp. glaucescens)
24. Midrib puberulent above……………………………………………………….…25

Midrib glabrous above…………………………………………………………..…29
25. Midrib glabrous below........................................................................................…26

Midrib puberulent below……………………………………………………………27
26. Leaf thickly coriaceous; lateral veins c. 9 pairs, lax. In forest on limestone karst hills
……………………………………………………………………..…20. S. calcicola
Leaf thinly coriaceous; lateral veins 9–12 pairs, dense. In kerangas forest or in forest
on organic soils over limestone……………………………………50. S. havilandii
27. Twig compressed……………………………………………….…45. S. foxworthyi

Twig terete………………………………………………………………………..…28
28. Petiole at least 1 cm long…………………………………....115. S. seminis (in part)

Petiole at most 0.8 cm long………………………..…114. S. scrobiculata (in part)
29. Petiole at most 1.1 cm long; lateral veins at most 10 pairs…………………….…30

Petiole at least 1.2 cm long or, if less, then leaf with more than 10 pairs of lateral
veins……………………………………………………………………………….32
30. Leaf blade larger, 10–14 × 4.5–7.5 cm; lateral veins at most 6 pairs…..62. S. ladiana
Leaf blade smaller, 5–10 × 1.8–5 cm; lateral veins at least 6 pairs…………………31
31. Species of ultrabasic soils (Sabah)………………………………..….77. S. micans

Species of shallow humic soils on high ridge crests……………………11. S. asahii
32. Leaf blade base shortly decurrent into petiole; lateral veins somewhat sunken above;
sapling leaf peltate…………………………………………………..…58. S. isoptera
Leaf base not decurrent; lateral veins not sunken; sapling leaf not peltate…………33
33. Inner bark pink. Lateral veins and midrib drying distinctly black below.……………..
…………………………………………………………………49. S. guiso (in part)
Inner bark yellowish brown. Lateral veins and midrib not drying black below.……34
34. Leaf blade at most 6 cm wide; petiole 1–1.5 cm long; intercostal venation
subreticulate…………………………………………………..…18. S. brunnescens
Leaf blade at least 6 cm wide; petiole 2–3.5 cm long; intercostal venation densely
scalariform…………………………………………………….….69. S. lunduensis
35. Leaf venation persistently tomentose below…………………………………….…36

Leaf venation glabrescent or glabrous below..……………………………………...37
36. Leaf undersurface grey tomentose; lateral veins 20–25 pairs……...…64. S. lamellata
Leaf undersurface bright yellow lepidote; lateral veins 25–30 pairs....84. S. ochracea
209
37. Leaf pale pinkish brown lepidote below; lateral veins 20–24 pairs…...31. S. dealbata
Leaf undersurface not pale lepidote below; lateral veins at most 18 pairs……….…38
38. Leaf base cuneate…………………………………………………………………39

Leaf base obtuse or subcordate……………………………………………….…40
39. Midrib slender, acute; lateral veins hardly raised below………..…48. S. gratissima
Midrib stout, terete; lateral veins distinctly raised below.………..…103. S. resinosa
40. Leaf blade chartaceous, drying undulate, chocolatebrown…………………………41

Leaf blade coriaceous, drying not undulate, tawny to pale yellowish brown………42
41. Leaf base usually subcordate; lateral veins 15–18 pairs, prominent below……………
…………………………………………………………………………25. S. cordata
Leaf base obtuse; lateral veins 11–15 pairs, slender and not prominent below………..
……………………………………………………………………17. S. bracteolata
42. Twig compressed apically; lateral veins 22–26 pairs……………..…127. S. virescens

Twig terete apically; lateral veins at most 22 pairs……………………………..…43
43. Leaf base more or less cordate; lateral veins at least 18 pairs……122. S. symingtonii
Leaf base obtuse; lateral veins at most 18 pairs……………………………………44
44. Leaf blade thinly coriaceous; lateral veins (9–)13–18 pairs, slender, not prominent
below, drying paler than the blade……………………………………24. S. confusa
Leaf blade thickly coriaceous; lateral veins 9–13 pairs, stout and prominent below,
drying of the same colour as blade……………………………………3. S. agamii
45. Bark deeply fissured, overall appearing more fissured than flaky………………..46
Bark smooth or appearing flaky rather than fissured………………………………48
46. Leaf blade persistently scabridtomentose or sometimes also creamlepidote below,

drying rich rustbrown. Bark dark chocolatebrown………….1. S. acuminatissima
Leaf blade glabrous or glabrescent on the veins below, drying dull greyish brown to
tawny. Bark tawny brown………………………………………………………..…47
47. Bark surface at first reticulate vsection fissured…………29. S. cuspidata (in part)
Bark surface squaresection fissured…………………..…36. S. faguetiana (in part)
48. Mature tree leaf deeply peltate; smooth barked subcanopy tree…….…94. S. peltata
Juvenile leaf peltate only; main canopy or emergent tree at maturity…………..…49
49. Leaf blade drying dark reddish brown, purplish rufous lepidote below……………….
…………………………………………………………………….99. S. polyandra
Leaf blade drying tawny to greyish green, not lepidote below…………………..…50
50. Midrib and lateral veins more or less persistently tomentose below…………….…51
Leaf blade entirely glabrous…………………………………………………….…58
51. Leaf blade greyish brown scabridtomentose below……………………………..…52

Leaf blade evenly or yellowish brown tomentose below.......................................…53
210
52. Leaf base cordate, margin revolute………………………………57. S. induplicata

Leaf base obtuse, margin not revolute…………………………68. S. longisperma
53. Petiole 2.2–3 cm long; lateral veins 12–14 pairs…………….…54. S. iliasii (in part)
Petiole less than 2 cm long or, if 2.2 cm, leaf with less than 11 pairs of lateral
veins.........................................................................................................................54
54. Lateral veins 14–19 pairs……………………………………………6. S. alutacea

Lateral veins less than 13 pairs……………………………………………………55
55. Midrib hidden and sunken above…………………………………82. S. obovoidea

Midrib clearly evident, at most shallowly furrowed above……………………..….56
56. Leaf base unequal……………………………………………………22. S. chaiana

Leaf base more or less equal…………………………………………………..…57
57. Twig smooth, persistently tomentose; leaf base obtuse……………….65. S. laxa

Twig glabrescent, rugulose; leaf base cuneate………………………47. S. gibbosa
58. Leaf blade dull greyish lepidote below……………………………….23. S. collaris

Leaf tawny brown, glabrous below.…………………………………………….…59
59. Leaf blade more than 3x as long as broad……………………………………….…60

Leaf blade less than 3x as long as broad………………………………………..…62
60. Leaf margin prominently revolute; lateral veins 12–15 pairs. In forest on acid humic
soils……………………………………………………………..…67. S. longiflora
Leaf margin hardly or not revolute; lateral veins at most 13 pairs. In forest on non
humic clay soils………………………………………………………………..….61
61. Leaf blade 12–15 × 4–7 cm; petiole 0.8–1.5 cm long, glabrous, drying black………...
…………………………………………………………………130. S. xanthophylla
Leaf blade 8–14 × 2.5–4 cm; petiole 0.6–0.8 cm long, puberulent, drying brownish
cream........................................................................................….9. S. angustifolia
62. Petiole at least 3 mm diameter…………………………………………………..…63

Petiole at most 2 mm diameter………………………………………………….…64
63. Leaf blade with revolute margin and cordate base…………...…71. S. macrobalanos
Leaf blade with flat margin and cuneate to obtuse base………..54. S. iliasii (in part)
64. Petiole 2–3 cm long; leaf blade frequently subcordate at base, chartaceous; lateral
veins 10–15 pairs…………………………………………….….37. S. faguetioides
Petiole at most 1.8 cm long or, if longer, leaf blade coriaceous and with at most 7
pairs of lateral veins…………………………………………………………….…65
65. Leaf margin revolute………………………………………………………….…66

Leaf margin flat………………………………………………………………..……68
211
66. Leaf base cuneate; lateral veins 5–7 pairs……………………………106. S. richetia

Leaf base obtuse; lateral veins at least 7 pairs………………………………..…67
67. Leaf blade ovate; midrib furrowed above…………………..…79. S. mujongensis

Leaf blade oblonglanceolate; midrib flat above………………..…13. S. bakoensis
68. Leaf blade narrow, at least twice as long as wide…………………………………69

Leaf blade broad, less than twice as long as wide……………………………..…70
69. Midrib and lateral veins evident but shallowly grooved above; canopy tree with flaky
bark……………………………………………………….…120. S. subcylindrica
Midrib and lateral veins not furrowed above; subcanopy tree with smooth bark……...
…………………………………………………………………123. S. tenuiramulosa
70. Bole eventually scrollmarked, smooth overall with patchy flakes……………………

……………………………………………………………29. S. cuspidata (in part)
Bole becoming evenly flaky…………………………………………………….…71
71. Petiole at least 1 cm long; leaf blade 7–15 cm long……………………………..…72

Petiole at most 1 cm long; leaf blade 2.5–8 cm long…………………………..…74
72. Leaf blade thinly coriaceous; lateral veins at most 7 pairs, prominent below…………
………………………………………………………………14. S. balanocarpoides
Leaf blade thickly coriaceous; lateral veins at least 7 pairs, hardly raised below…..73
73. Lateral veins 7–9 pairs; petiole at least 1.7 cm long……………….60. S. kudatensis
Lateral veins 9–12 pairs; petiole at most 1.5 cm long….….36. S. faguetiana (in part)
74. Leaf blade narrowly ovate; lateral veins 7–9 pairs, slender but prominent below…….
……………………………………………………………………92. S. patoiensis
Leaf blade broadly ovate; lateral veins 8–11 pairs, hardly raised below................75
75. Midrib drying dark purplish brown to red below; vast emergent tree…………………
…………………………………………………………………….52. S. hopeifolia
Midrib drying of the same colour as blade; low emergent or main canopy tree……….
…………………………………………………………………….80. S. multiflora
76. Bark remaining smooth, hoopmarked, or eventually becoming cracked or shallowly

vsection fissured, and sometimes patchily flaky…………………………………77
Bark early becoming flaky or deeply fissured……………………………………90
77. Stipule scars amplexicaul……………………………………………………………78

Stipule scars not amplexicaul………………………………………………………85
78. Stipules ovate, cordate, subpersistent; leaf blade glabrous……………………..…79

Stipules oblong to hastate; leaf blade glabrous or pubescent………………………80
79. Twigs with sections with short internodes and no leaves……….…119. S. stenoptera
Twigs without leafless sections with short internodes…………..…118. S. splendida
212
80. Leaf base peltate…………………………………………………..….129. S. woodii

Leaf base cordate, obtuse or cuneate………………………………………………81
81. Leaf blade densely golden brown tomentose below; lateral veins with prominent
tomentose domatia………………………………………………………95. S. pilosa
Leaf blade sparsely puberulent or glabrous below; lateral veins without domatia….82
82. Stipules falcate, subacute; twigs less than 3 mm diameter apically, slender……83
Stipules oblong to hastate, obtuse; twigs at least 3 mm diameter apically, stout…...84
83. Leaf blade broadly ovate to suborbicular, 10–13 cm wide………107. S. rotundifolia

Leaf blade elliptic, 5–8 cm wide…………………………….…7. S. amplexicaulis
84. Petiole 1.5–3 cm long. In forest on clay rich soils…….72. S. macrophylla (in part)
Petiole 3.5–4.5 cm long. In forest on sandy soils………………….100. S. praestans
85. Twigs distinctly compressed apically. Bark remaining smooth or becoming shallowly
flaky……………………………………………………………………………..….86
Twigs more or less terete apically. Bark becoming vsection fissured, eventually
powdery chunkily flaky………………………………………………………….…88
86. Leaf base cordate, blade densely golden tomentose below……………………………

…………………………………………………………76. S. mecistopteryx (in part)
Leaf base cuneate or obtuse, blade not densely golden tomentose below.………….87
87. Stipule scars short, horizontal or ascending. Leaf blade dull below; lateral veins 11–
14 pairs……………………………………………….…15. S. beccariana (in part)
Stipule scars falcate, descending. Leaf blade shiny below; lateral veins 14–19 pairs…
……………………………………………………………………….96. S. pinanga
88. Tomentum on veins and twigs even, short; lateral veins slender, hardly raised below..
.………………………………………………………….41. S. ferruginea (in part)
Tomentum on veins and twig scabrous; lateral veins stout, prominent below….…..89
89. Leaf blade 14–26 × 6.5–12 cm. Inner bark plumred………...…2. S. acuta (in part)
Leaf blade 8–16 × 4–6 cm. Inner bark pale pinkish brown……………………………
…………………………………………………………..73. S. macroptera (in part)
90. Leaf base distinctly unequal, subcordate………………………………………..…91

Leaf base equal or subequal……………………………………………………..…92
91. Leaf blade evenly tomentose below; lateral veins 11–13 pairs………………………..
……………………………………………………….55. S. inaequilateralis (in part)
Leaf blade scabridtomentose below; lateral veins 13–17 pairs……70. S. macrantha
92. Leaf blade cream, pink or golden lepidote below…………………………………...93

Leaf blade not distinctly lepidote below..........................................................….102
93. Twig compressed apically…………………………………………………………94

Twig terete or ribbed apically…………………………………………………..…95
213
94. Lateral veins 11–14 pairs……………………………….…15. S. beccariana (in part)

Lateral veins 16–20 pairs…………………………………………….…4. S. albida
95. Lateral veins 9–11 pairs………………………………………………………..…96

Lateral veins 5–7 or at least 10 pairs………………………………………………97
96. Le af bl ad e pi n k l epi d ot e b el ow; l at eral vein s hardl y rai sed

below……………………………………………….....28. S. curtisii (subsp. curtisii)
Leaf blade more or less cream lepidote below; lateral veins prominent below………..
………………………………………………………………32. S. dispar (in part)
97. Leaf blade golden lepidote below. In montane forest at altitude above 900 m………..
…………………………………………………………………….78. S. monticola
Leaf blade cream lepidote below. In lowland forest at altitude to 800 m………..…98
98. Lateral veins 5–7 pairs…………………………………………..…108. S. rubella

Lateral veins at least 10 pairs…………………………………………………..…99
99. Bark appearing flaky rather than fissured. Leaf blade thinly coriaceous, sparsely
lepidote below, base narrowly cuneate………………………….…..8. S. andulensis
Bark prominently vsection fissured. Leaf blade chartaceous or thickly coriaceous,
evenly or scabridtomentose, base obtuse to cordate (occasionally broadly cuneate)
……………………………………………………………………………….….100
100. Leaf blade chartaceous; lateral veins at least 18 pairs; tomentum if present even…….
……………………………………………………………………….128. S. waltoni
Leaf blade thickly coriaceous; lateral veins at most 19 pairs; tomentum scabrid….101
101. Leaf blade 13–22 × 7–13 cm, ochreous scabrid below...…………..89. S. pallidifolia
Leaf blade 7–16 × 4–9 cm, evenly cream lepidote below except on veins……………
…………………………………………………………………………34. S. elliptica
102. Lateral veins at most 13 pairs………………………………………………..….103

Lateral veins at least 14 pairs…………………………………………………..….130
103. Bark appearing fissured rather than flaky……………………………………….104

Bark appearing flaky rather than fissured………………………………………122
104. Leaf venation below entirely tomentose……………………………………….….105

Leaf blade below glabrous or puberulent on midrib and lateral veins only……..…114
105. Leaf tomentum scabrid or tufted……………………………………………….….106

Leaf tomentum even, short……………………………………………………….109
106. Bark shallowly cracked. Leaf blade somewhat bullate between intercostal veins…….
…………………………………………………………………19. S. bullata (in part)
Bark becoming more or less deeply vsection fissured. Leaf blade flat……….….107
107. Leaf blade sparsely tometose below………………………113. S. scabrida (in part)

Leaf blade densely tomentose below.………………………………………….….108
214
108. Tomentum tawny, prominently tufted……………………..…110. S. rugosa (in part)

Tomentum yellowish brown, scabrous………………………….….30. S. dasyphylla
109. Leaf margin more or less revolute………………………………………………..110

Leaf margin flat……………………………………………………………….…113
110. Leaf margin prominently revolute; blade coriaceous, glabrescent, shiny below. In
kerangas forest………………………………………………….…105. S. revoluta
Leaf margin narrowly slightly revolute; blade thinly coriaceous, not shiny below. In
other forest types………………………………………………………………….111
111. Tomentum rich vinous reddish brown…………………………………109. S. rubra

Tomentum cream to brown…………………………………………………….112
112. Leaf blade 6–11 × 3.5–6 cm; lateral veins 10–13 pairs, the basal pair often with large
pale glabrous domatia……………………………………………..90. S. parvifolia
Leaf blade 4–8 × 2.5–4.5 cm; lateral veins 8–10 pairs, with small hairy domatia in
their axils………………………………………………………………….86. S. ovata
113. Leaf blade 5–9 × 3–5 cm, glabrescent below; lateral veins 8–11 pairs, without
prominent domatia………………………………………….113. S. scabrida (in part)
Leaf blade 8–14 × 3.5–5.5 cm, sparsely pale brown puberulent below; lateral veins
12–15 pairs, sparsely puberulent with pale scalelike domatia in rows up the midrib
(immature), or with densely creambuff puberulent undersurface without domatia…..
………………………………………………………………66. S. leprosula (in part)
114. Lateral veins only 4 pairs; blade concave.............................….102. S. quadrinervis

Lateral veins at least 5 pairs; blade flat…………………………………………...115
115. Petiole 4–6 cm long; blade broadly ovate to orbicular, 10–20 × 9–16 cm, thickly
coriaceous. In peat swamp forest and poorly drained kerangas…..87. S. pachyphylla
Petiole less than 3.5 cm long; blade otherwise. In other types of habitat.…………116
116. Petiole at most 1.7 cm long…………………………………………………….….117

Petiole at least 2 cm long……………………………………………………….…120
117. Leaf apex retuse……………………………………………………….104. S. retusa

Leaf apex acuminate……………………………………………………………….118
118. Leaf base cuneate; lateral veins without domatia. On inland hills……………………..
………………………………………………………………….32. S. dispar (in part)
Leaf base obtuse; lateral veins with domatia. In lowland swamps and valleys……119
119. Lateral veins 8–9 pairs with prominent axillary domatia up the midrib……………….
…………………………………………………………………….44. S. foraminifera
Lateral veins 11–12 pairs with a few pairs of small domatia towards the midrib base
only..………………………………………………………..…124. S. teysmanniana
120. Lateral veins 11–13 pairs; leaf margin revolute……………………......21. S. carapae

Lateral veins 6–8 pairs; leaf margin flat……………………………………….…121
215
121. Leaf blade narrowly ovate, drying rustbrown below; lateral veins slender but
prominent below, ascending. Bark dark chocolatebrown………..…42. S. flaviflora
Leaf blade broadly ovate, drying dull greyish; lateral veins slender and hardly raised
below, spreading. Bark pale greyish brown……………..….61. S. kunstleri (in part)
122. Stipule scars amplexicaul………………………………72. S. macrophylla (in part)

Stipule scars not amplexicaul……………………………………………………..123
123. Leaf blade scabridtomentose below…………………………………………..…124

Leaf blade evenly tomentose, glabrous or glabrescent below............................…..125
124. Leaf blade concave, with revolute margin, more or less bullate between intercostal
veins...……………………………………………………….…19. S. bullata (in part)
Leaf blade flat, margin not revolute……………………91. S. parvistipulata (in part)
125. Lateral veins 8–9 pairs. Inner bark plumred…………………….….93. S. pauciflora

Lateral veins at least 10 pairs. Inner bark rust, plum or yellowbrown……..……126
126. Leaf base with to 3 pairs of pale scalelike domatia in the axils of the basal lateral
veins below..………………………………………………………..59. S. johorensis
Leaf base without pale scalelike domatia……………………………………..…127
127. Leaf blade drying deep reddish brown below, purplish brown above…………………
…………………………………………………………..88. S. palembanica (in part)
Leaf blade drying pale orangerust below, pale pinkish brown above……………128
128. Lateral veins hardly raised below; blade dull lepidote below………………………….
……………………………………………………………..41. S. ferruginea (in part)
Lateral veins prominent below; blade glabrescent, somewhat shiny below..….…..129
129. Leaf blade 14–26 × 6.5–12 cm..…………………………………2. S. acuta (in part)

Leaf blade 8–16 × 4–6 cm……………………………….73. S. macroptera (in part)
130. Leaf blade evenly pubescent or glabrous below………………………………..….131

Leaf blade scabridtomentose below………………………………………………140
131. Lateral veins very slender, hardly raised below………………………………..…132

Lateral veins prominent below………………………………………………….…135
132. Bark cracked, scaly, rather than fissured………………….41. S. ferruginea (in part)
Bark appearing fissured rather than flaky……………………………………..…133
133. Midrib sharply acute, prominent below; blade lanceolate, margin undulate, base
inrolled; petiole 1–1.5 cm long……………………………………98. S. platyclados
Midrib terete below; blade broadly ovate, margin not undulate, base not inrolled;
petiole at least 1.5 cm long………………………………………………………..134
134. Leaf blade 10–15 × 5–8 cm, base subpeltate, margin revolute. Stipule scars cuneate
…………………………………………………………………………26. S. coriacea
216
Leaf blade 6–10 × 3–5 cm, base not subpeltate, margin hardly revolute. Stipule scars
amplexicaul………………………………………………………..…126. S. venulosa
135. Bark flaky…………………………………………………………………….…136

Bark fissured rather than flaky……………………………………………….…138
136. Leaf blade oblong, base cordate, undersurface golden tomentose……………………..

…………………………………………………………76. S. mecistopteryx (in part)
Leaf blade if oblong and with cordate base, then not golden tomentose below…...137
137. Leaf blade densely pale creambuff puberulent below, or sparsely pale brown
puberulent and with pale scalelike domatia running alongside the midrib…………...
……………………………………………………………..66. S. leprosula (in part)
Leaf blade glabrous below but for sparsely buffpuberulent midrib and lateral veins,
without domatia…………………………………………88. S. palembanica (in part)
138. Leaf blade 6–11 × 2.5–4.5 cm, densely pinkish buffvelutinous below……………….
……………………………………………………………………..10. S. argentifolia
Leaf blade at least 8 × 4.5 cm, not densely velutinous below………………….…139
139. Bark deeply fissured. Leaf blade ellipticoblong to broadly ovate; lateral veins 16–20
pairs. In mixed peat swamp forest…………………………97. S. platycarpa (in part)
Bark shallowly fissured. Leaf blade narrowly oblong; lateral veins 24–28 pairs. On
river banks………………………………………………81. S. myrionerva (in part)
140. Bark cracked and flaky, not deeply fissured…………………………………….…141

Bark appearing fissured rather than flaky…………………………………………145
141. Leaf venation densely scabridtomentose below…………………………………..142

Leaf venation sparsely tomentose below.............................................................….144
142. Leaf blade concave……………………………………………...5. S. almon (in part)

Leaf blade flat………………………………………………………………….…143
143. Leaf blade narrowly oblong to lanceolate, coriaceous; tomentum pinkbrown………..

………………………………………………………………………..111. S. sagittata
Leaf blade broadly oblong to elliptic, thinly coriaceous; tomentum pale yellow to
greybrown………………………………………….….91. S. parvistipulata (in part)
144. Leaf blade 14–26 × 6.5–12 cm. Inner bark plumred...……….…2. S. acuta (in part)
Leaf blade 8–16 × 4–6 cm. Inner bark pale pinkish brown……………………………
…………………………………………………………..73. S. macroptera (in part)
145. Bark dark warm brown to blackish, deeply fissured………………………………146

Bark pinkish to greyish brown, shallowly or deeply fissured……………………151
146. Inner bark yellowish or pinkish brown………………………………………..…147

Inner bark rich meat red……………………………………………………….…150
217
147. Leaf blade concave…………………………………………………………………148

Leaf blade flat…………………………………………………………………..…149
148. Petiole 2.2–3.2 cm long. In mixed peat swamp forest………………125. S. uliginosa

Petiole 1–1.8 cm long. In forest on clay soils……………………5. S. almon (in part)
149. Petiole 1.3–2.3 cm long………………………………….……110. S. rugosa (in part)

Petiole 2.5–4 cm long…………………………………………...…101. S. pubistyla
150. Leaf blade ovate, 5–9 × 3–4.5 cm; lateral veins 14–17 pairs. Bark blackish………….
……………………………………………………………………..43. S. flemmichii
Leaf blade broadly oblong, 11–22 × 4–7 cm; lateral veins 25–34 pairs. Bark rich
reddish brown…………………………………………………..……116. S. slootenii
151. Leaf blade at least 3x as long as broad……………………………………………152

Leaf blade at most 2½x as long as broad………………………………………….153
152. Leaf blade 12–17 × 2–4.5 cm, bending back along the midrib, densely tomentose
below; lateral veins 20–25 pairs……………………………………..….85. S. ovalis
Leaf blade 12–25 × 4.5–9 cm, remaining flat, sparsely tomentose below; lateral veins
24–28 pairs…………………………………………….81. S. myrionerva (in part)
153. Bark shallowly patchily flaky…………………………………………………..…154

Bark deeply fissured………………………………………………………….…156
154. Lateral veins 18–22 pairs. Inner bark pink……………91. S. parvistipulata (in part)
Lateral veins 14–17 pairs. Inner bark yellowish brown………………………..…155
155. Lateral veins with short intermediate veins……………………….112. S. scaberrima

Lateral veins without short intermediate veins………………………………………...
………………………………………………51. S. hemsleyana (subsp. grandiflora)
156. Lateral veins without distinct intermediate veins. In mixed peatswamp forest………..
…………………………………………………………….97. S. platycarpa (in part)
Lateral veins with distinct intermediate veins. In mixed dipterocarp forest……157
157. Petiole 0.8–1.5 cm long; leaf not lepidote below; lateral veins 15–20 pairs…………..
……………………………………………………………………………40. S. fallax
Petiole 2.2–2.7 cm long; leaf frequently dull greyish lepidote below (mature trees);
lateral veins 14–17 pairs……………………………………….…..117. S. smithiana
1. Shorea acuminatissima Symington

(Latin, acuminatissimus = prominently acuminate; the leaf apex)
(sect. Richetioides; yellow meranti)
Gard. Bull. S. S. 9 (1938) 340; Browne, FTSB (1955) 162; Ashton op. cit. (1964) 149, op. cit. (1968)
81, op. cit. (1982) 482; Meijer & Wood op. cit. 65; Burgess op. cit. 217; Anderson op. cit. (1980) 121;
PROSEA op. cit. 417; Coode et al. (eds.) op. cit. 74; Newman et al. op. cit. (1996) 109. Type: Ayan
FMS 38723, Borneo, Sabah, Sandakan (holotype KEP).
218
Vast emergent tree, to 70 m tall, to 2 m diameter; buttresses tall, stout. Bark dark
chocolatebrown, deeply squaresection fissured, the ridges only later becoming flaky; inner
bark tawnybrown, fibrous, not distinctly laminated. Twig, inflorescence, petiole, leaf bud,
parts of flower exposed in bud, stipules and venation below more or less persistently rusty
brown scabridpubescent. Twigs c. 1.5 mm diameter apically, terete, muchbranched. Leaf
buds ovoid, to 1.2 mm long. Stipules oblongfalcate, c. 6 × 3.5 mm, base obtuse to
subcordate, somewhat persistent in young trees. Leaves glabrous or sometimes cream
lepidote below, coriaceous, drying rich rustybrown or cream below with the veins usually
darker; blade ovate, 7–10 × 3–4 cm, base obtuse, margin revolute, apex with acumen to 0.7
cm long; midrib evident but sunken above, prominent below; lateral veins 9–12 pairs,
prominent below; intercostal venation densely scalariform, distinctly raised below; petiole
1–1.5 cm long, slender. Inflorescences terminal or axillary; rachis to 8 cm long, singly
branched, branchlets bearing to 6 flowers; bracteoles oblong, to 2 × 2 mm, obtuse. Flowers:
buds ellipsoid, to 5 × 3 mm; sepals broadly ovate; petals yellow; stamens 15, anthers 2
loculed, locules subglobose to broadly elliptic, connectival appendage aristate, sericeous,
exceeding anther apice; ovary and stylopodium glabrous but for band of minute hairs
towards apex, style equal to ovary and stylopodium, stigma not trifid. Fruits subsessile;
calyx lobes unequal, 3 longer lobes spatulate, to 6 × 1.3 cm, tapering to 3 mm above the
tuberculate concave base, 2 shorter ones to 4.5 × 0.8 cm, otherwise similar. Nuts narrowly
ovoid, to 4.5 × 0.8 cm, densely pale rufous pubescent.
Vernacular names. Sabah—seraya kuning runcing (preferred name). Sarawak—lun

runcing (preferred name).
Distribution. Endemic in Borneo. In Sabah known from Beaufort, Kinabatangan,

Sandakan, and Sipitang districts (e.g., SAN 16942, SAN 17767, SAN 17953, SAN 37516, and
SAN 70201) and in Sarawak from Lawas district (e.g., S 26560). Also occurring in Brunei
(e.g., BRUN 3350 and FMS 30404).
Ecology. Locally frequent in mixed dipterocarp forest on friable sandy clay soils, on low
hills at altitudes to 1600 m (near G. Murut). Vulnerable.
2. Shorea acuta P.S.Ashton

(Latin, acutus = sharp; the leaf tip)
(sect. Mutica, subsect. Auriculatae; red meranti)
op. cit. (1980) 124; Coode et al. (eds.) op. cit. 74; Newman et al. op. cit. (1996) 110. Type: Mohsin S
1924, Borneo, Brunei, Belait, Sg. Liang (holotype K; isotype KEP).
Small emergent tree, to 45 m tall, to 1 m diameter; buttresses stout. Bark cracked and flaky
becoming shallowly vsection fissured, pale reddish brown; inner bark plumbrown, densely
fibrous. Young twig, inflorescence, leaf bud, parts of flower exposed in bud, petiole,
venation below and midrib above shortly pale brown puberulent; glabrescent on leaf and
fruit calyx. Twigs terete, to 4 mm diameter apically, compressed at first, with non
amplexicaul stipule scars to 3 mm long. Leaf buds ovoid, 6–10 × 4–6 mm, acute. Stipules
oblong, to 12 × 5 mm, obtuse. Leaves thickly coriaceous, shiny and scabridtomentose or
glabrescent below, drying pale pinkish brown above; blade elliptic, 14–26 × 6.5–12 cm,
base equal or subequal, without domatia, obtuse or occasionally broadly cuneate, apex with
219
tapering acumen to 2 cm long; midrib prominent below, narrowly slightly sunken above;
lateral veins 10–13 pairs, prominent below, arched; intercostal venation densely
scalariform, slender, sinuate; petiole 1.5–2 cm long, stout. Inflorescences terminal or
axillary; rachis compressed, to 8 cm long, singly or doubly branched, branchlets bearing to 9
flowers; bracteoles elliptic, to 3.5 × 2.5 mm, caducous. Flowers: buds narrowly ellipsoid,
obtuse, to 9 × 4 mm; sepals subequal, ovate; petals linear, dark crimson; stamens 15,
filament flat, tapering from base to apex, connectival appendage aristate, not setose, at most
of the same length as anther, becoming reflexed at anthesis, anther 4loculed, locules
broadly oblong to subglobose; ovary and distinct stylopodium pubescent. Fruits: calyx
lobes unequal, 3 longer lobes spatulate, obtuse, to 15 × 2.8 cm, tapering to 1.5 cm towards
the narrowly auriculate base, 2 shorter ones unequal, linearlobed, to 8 × 0.7 cm, similar at
base. Nuts ovoid, to 3 × 2 cm, acute, buff puberulent.
Vernacular names. Sarawak—engkabang tikus (Iban), kawang tikus (Iban), meranti

kawang tikus (Malay).
Distribution. Endemic in Borneo. Recorded in Sarawak from Limbang, Marudi and Miri
(e.g. BRUN 3076, BRUN 3291, S 1906, and SAN 17474). Conserved in the Lambir Hills NP
and Mulu NP, Sarawak.
Ecology. Locally common in mixed dipterocarp forest on deep yellow sands, on hills at
altitudes to 400 m. Endangered outside parks system.
3. Shorea agamii P.S.Ashton

(J. Agama, onetime forest officer in the Sabah Forest Department)
(sect. Anthoshorea, white meranti)
Gard. Bull. Sing. 19 (1962) 270, op. cit. (1964) 161, Gard. Bull. Sing. 22 (1967) 285, op. cit. (1968)
PROSEA op. cit. 409; Coode et al. (eds.) op. cit. 74; Newman et al. op. cit. (1996) 111. Type: Smith
FMS 30525, Brunei, Labi Hills (holotype KEP).
Emergent tree, to 50 m tall, to 2 m diameter, with prominent stout buttresses. Bark

becoming chocolate, ochreous and grey mottled, irregularly cracked and thickly flaky
through surface rotting; inner bark distinctly laminated, orangebrown and creammottled.
Young twig, inflorescence, leaf bud, parts of perianth exposed in bud, ovary, stipule outside,
petiole and midrib below pale brown puberulent, caducous. Twigs terete, 2–3 mm diameter
apically. Leaf buds ovoid to falcate, c. 4 × 2.5 mm. Stipules oblong, obtuse, c. 10 × 3 mm,
caducous. Leaves thickly coriaceous, drying tawny yellow with the venation the same
shade; blade broadly ovate to oblong, 10–15 × 6–10 cm (subsp. agamii) or narrowly ovate,
4.5–10 × 2.5–4 cm (subsp. diminuta), base obtuse or subcordate, apex with broad acumen
of 0.5–1 cm long; midrib prominent below, obscure, furrowed above as also the veins;
lateral veins 9–13 pairs, wellspaced, arched; intercostal venation densely scalariform;
petiole 1–1.5 cm long. Inflorescences terminal or axillary; rachis to 10 cm long, branches
short, bearing to 5 flowers; bracteoles elliptic, to 3 mm long, caducous. Flowers: buds
narrowly ellipsoid, to 5 × 2.5 mm; sepals narrowly ovate, unequal; petals ovateelliptic,
acute, densely pubescent on exposed parts in bud; stamens 15, in 3 unequal whorls,
filaments more or less as long as anthers, tapering gradually, anthers oblong, 4–loculed,
220
connectival appendage c. 3x the length of anther; ovary ovoid, without stylopodium, densely
pubescent, style slightly longer than ovary, sparsely pubescent in the basal half, stoutly
filiform, obscurely trifurcate. Fruits impressed at the c. 1 mm stout pedicel; calyx lobes
unequal, 3 longer lobes to 12 × 2.2 cm, tapering to c. 5 mm above the saccate base, 2
shorter ones linearlobed, unequal, to 6 × 0.4 cm. Nuts ovoid, to 2 × 1.5 cm, glabrescent,
with acicular style remnant to 4 mm long.
Vernacular names. Sabah—melapi agama (preferred name). Sarawak—meranti putih

timbul (preferred name).
Ecology. Scattered in mixed dipterocarp forest on welldrained sandy clay soils, and clay
rich soils on ridges, at altitudes to 700 m.
Notes. Two subspecies, subsp. agamii and subsp. diminuta, are recognised.
Key to subspecies
Leaf blade broadly ovate to oblong, 10–15 × 6–10 cm………………………………………..

subsp. agamii
In Sabah known from Beaufort, Kudat, Sandakan, Sipitang, and Tawau districts (e.g.,
SAN 18476 and SAN 36222) and in Sarawak from Lawas, Marudi and Miri districts
(e.g., S 1537 and S 1936). Also occurring in Brunei (e.g., BRUN 3092 and FMS 35449)
and E Kalimantan (e.g., Ambriansyah Berau 1140 and Meijer 2464). Occurring in
Lambir and Mulu NPs; elsewhere vulnerable following logging.
Leaf blade narrowly ovate, 4.5–10 × 2.5–4 cm…………..……………………………………
subsp. diminuta P.S.Ashton
(Latin, diminutus = made small; the leaf)
Gard. Bull. Sing. 22 (1967) 285, op. cit. (1968) 93, op. cit. (1982) 497; PROSEA op. cit. 409;
Newman et al. op. cit. (1996) 112. Type: Sibat S 25027, Borneo, Sarawak, Bt. Mersing, Anap
(holotype K; isotype KEP).
Known in Sarawak from Kapit, Miri, Serian, Simunjan, and Tatau districts (e.g., S
15227, S 23860, S 27005, S 36655, and S 46520). Also occurring in W Kalimantan
(e.g., bb. 27005, bb. 29076 and Wilkie 94313). Ecology as the species. Vulnerable
following logging. For differences from S. confusa, see there.
4. Shorea albida Symington Fig. 19.

(Latin, albidus = whitish; the leaf undersurface)
(sect. Rubella, red meranti)
Gard. Bull. S. S. 8 (1935) 283; Masamune op. cit. 492; Browne op. cit. 137; Anderson, Gard. Bull.
Sing. 20 (1963) 158, op. cit. (1980) 74; Ashton op. cit. (1964) 175, op. cit. (1968) 104, op. cit. (1982)
499; PROSEA op. cit. 391. Lectotype (designated here): Zainal FMS 30353, Borneo, Brunei, Kuala
Belait (hololectotype KEP; isolectotype L).
221
Fig. 19. Shorea albida. A, flowering leafy twig; B, detail of indumentum on the lower leaf
surface; C, flower bud; D, outer sepal; E, inner sepal; F, adaxial view of petal; G, abaxial
view of petal; H, longitudinal section of open flower; I, adaxial view of stamens; J, abaxial
view of stamens; K, fruit; L, longitudinal section of fruit. (A–J from S 651, K–L from S
2852.)
222
Main canopy tree c. 30 m tall to giant emergent to 75 m tall, to 2.5 m diameter; crown vast,
diffuse, seagreen; buttresses to 4 m tall and wide, stout. Bark becoming deeply vsection
fissured, the surface eventually rotting off in irregular flakes; inner bark meatred, fibrous,
thick. Heartwood crimson red. Twig, inflorescence, leaf bud, stipule and bracteole outside,
parts of perianth exposed in bud, petiole and leaf venation below shortly densely
persistently pinkish brown puberulent; leaf creamlepidote below. Twigs compressed, to 5
× 2 mm apically. Leaf buds ovoid to hastate, compressed, acute, to 10 × 6 mm. Stipules
oblong, obtuse, to 20 × 8 mm, caducous. Leaves coriaceous, drying greyish brown above,
cream below with darker venation; blade oblongelliptic, 7.5–15 × 4.5–6.5 cm, base obtuse,
apex with tapering acumen to 6 m long; midrib obscure, sunken above, slender but
prominent below; lateral veins 16–20 pairs, with shorter intermediates, slender, hardly
elevated below; intercostal venation slender, densely scalariform; petiole 2–3.5 cm long,
slender. Inflorescences terminal or axillary; rachis compressed, lax, to 18 cm long, doubly
branched, branchlets bearing to 3 flowers; bracteoles lanceolate, acute, to 8 × 3 mm.
Flowers: buds broadly ellipsoid, to 9 × 5 mm, acute; sepals deltoid, unequal; petals ovate,
cream; stamens 20–25, filament broad at base, tapering gradually and filiform distally,
anthers 4loculed, locules narrowly oblong, longer than filament, connectival appendage
glabrous, short, slender, erect, shorter than anther; ovary without stylopodium, together
with basal half of style pubescent, style 2x the length of ovary. Fruits: calyx lobes unequal,
3 longer lobes narrowly spatulate, to 8 × 1.4 cm, tapering to 4 mm above the saccate base, 2
shorter ones linearlobed, to 3.5 × 0.4 cm. Nuts ovoid, to 1.2 × 0.9 cm, greybuff
puberulent, with to 2 mm filiform style remnant.
Vernacular name. Sarawak—alan (preferred name).
Distribution. Endemic in Borneo; known in Sarawak from Bintulu, Limbang Marudi, Miri,
Serian, Sibu, and Sri Aman districts (e.g., S 1253, S 2805, S 12250, S 13504, and S 13506),
Brunei (e.g., BRUN 5009 and FMS 37149), and Lower Kapuas in W Kalimantan (e.g., bb.
35243).
Ecology. Formerly gregarious and dominant in three of the ‘phasic communities’ (PC) of
the peat swamp forests (Anderson op. cit. 1963); from within the outermost mixed peat
swamp forest, where this species reaches its greatest size in PC 2, the ‘alan forest’, through
PC 3, the ‘alan bunga forest’, to PC 4, where it is shortest, the ‘padang alan forest’. The
species seldom flowers and juveniles are rare, few surviving, except in the ‘alan bunga
forest’ where there is patchy regeneration; in ‘padang alan’ forest the trees frequently
topple, then shoot from epicormic branches forming new orthotrophic leaders. Alan forest
individuals are generally hollow. Also occasional in poorly drained lower montane kerangas
on Merurong Plateau, Bintulu district, at 800 m altitude. Occurring in Loagan Bunut and
Mulu NPs. Critically endangered owing to failure to regenerate and conversion of the peat
swamp forests.
5. Shorea almon Foxw.

(from a Philippine name—almon)
(sect. Brachypterae, red meranti)
Philip. J. Sci. 67 (1938) 313; Browne op. cit. 150; Meijer & Wood op. cit. 93; Burgess op. cit. 181;
Ashton op. cit. (1968) 104, op. cit. (1982) 507; Anderson op. cit. (1980) 124; PROSEA op. cit. 291;
223
Newman et al. op. cit. (1996) 114. Lectotype (designated here): Whitford FB 11647, the Philippines,
Cadiz, Negros Occidental (hololectotype K; isolectotype L).
Large emergent tree, to 70 m tall, to 1.2 m diameter; crown dense; bole tall, straight;
buttresses to 2.5 m tall, 2 m wide, stout. Bark dark greyish brown, becoming vertically
cracked and thinly flaked; inner bark pinkish brown, fibrous, hard. Twig, inflorescence, bud,
stipule and bracteole outside, parts of perianth exposed in bud, and leaf venation below
more or less densely persistently pinkish brown scabridtomentose; stipule inside and
venation above sparsely so; midrib above, nut and ovary evenly so. Twigs c. 2 mm diameter
apically. Leaf buds ovate, compressed, acute, to 4 × 3 mm. Stipules lanceolate, acute, to 10
× 5 mm, caducous. Leaves chartaceous, somewhat concave, drying pinkish brown with the
midrib and lateral veins typically distinctly paler; blade obovateoblong, 9–16 × 3.5–7 cm,
base equal, broadly cuneate to obtuse, apex with short, broad acumen, to 0.8 cm long;
midrib evident, flat or slightly furrowed above, slender but prominent below as also the
veins; lateral veins 17–20 pairs; intercostal venation densely scalariform, elevated below;
petiole slender, 1–1.8 cm long. Inflorescences terminal or axillary; rachis ascending, lax, to
20 cm long, singly branched, branchlets bearing to 10 flowers; bracteoles elliptic, to 5 × 3
mm, obtuse, not at first caducous. Flowers: buds lanceolate, to 7 × 3 mm; sepals unequal,
deltoid to ovate; petals lanceolate, hirsute on exposed part in bud; stamens 15, in 3 unequal
whorls, filaments compressed at base, abruptly tapering medially and filiform below the
subglobose, 4loculed anthers, connectival appendage slender, glabrous, c. 3x the length of
anther; ovary without distinct stylopodium, ovoid, style c. 1½x the length of ovary, filiform.
Fruits: pedicels to 3 mm long, prominent; calyx lobes unequal, 3 longer lobes to 14 × 2.5
cm, tapering to c. 8 mm above the saccate base, 2 shorter ones loratelobed, to 8 × 1 cm.
Nuts ovoid, to 2 × 1.5 cm, shortly mucronate.
Vernacular name. Sabah—seraya kerukup (preferred name).
Distribution. Borneo and the Philippines. In Borneo, known in Sabah from Keningau,
Kinabatangan, Lahad Datu, and Sandakan districts (e.g., SAN 4472, SAN 23966, SAN
35632, and SAN 83424) and in Sarawak from Bintulu, Lawas and Miri districts (e.g., S
22576 and S 24960). Also occurring in Brunei (e.g., FMS 35679 and S 5792) and NE
Kalimantan (e.g., Arifin et al. Berau 906).
Ecology. Usually rare in mixed dipterocarp forest, on yellow clay soil over shale, on
undulating land, at altitudes to 400 m. Rare in Lambir NP; endangered.
6. Shorea alutacea P.S.Ashton

(Latin, alutaceus = leather colour; the tomentum)
(sect. Richetioides, subsect. Richetioides, yellow meranti)
Newman et al. op. cit. (1996) 115. Type: Bojeng S 10170, Borneo, Sarawak, Lundu district, G.
Gading NP (holotype K).
Mediumsized tree, to 1 m diameter; buttresses stout. Bark smooth; inner bark fibrous.
Twig, inflorescence, bracteole, parts of flower exposed in bud, ovary and nut, stipules, leaf
bud, petiole and midrib shortly densely persistently tawny buff pubescent; leaf blade below
sparsely so; fruit calyx sparsely caducously so. Twigs c. 2 mm diameter apically, terete,
224
sparsely branched, with prominent stipule scars. Leaf buds ovoid, acute, c. 2 × 1.5 mm,
small. Stipules narrowly deltoid, acute, to 8 × 3 mm, relatively large. Leaves chartaceous,
drying pale greenish grey, wrinkling; blade ovate to lanceolate, 15–22 × 6–8 cm, base
cordate, apex with slender acumen to 1 cm long; midrib and veins furrowed above, slender
but prominent below; lateral veins 14–19 pairs; intercostal venation slender,
subscalariform, indistinct; petiole 0.6–1 cm long, 0.2 cm diameter, rugose on drying.
Inflorescences terminal or axillary; rachis to 15 cm long, singly or doubly branched;
bracteoles to 4 × 3 mm, ovate, acute, fugaceous. Flowers: buds to 6 × 3 mm; stamens 15,
anthers 2loculed, locules subglobose, tapering abruptly medially, connectival appendage
setose, 3–4x the length of anther; ovary and stylopodium pyriform. Fruits: pedicel c. 1 mm
long; calyx lobes unequal, 3 longer lobes to 8 × 1.8 cm, tapering to 4 mm above the
tuberculate narrowly saccate base, 2 shorter ones to 6 × 1.2 cm. Nuts narrowly ovoid, to
2.2 × 1 cm, apiculate.
Distribution. Locally endemic in the granodiorite mountains of W Sarawak; so far only

known from G. Gading NP, Lundu district (e.g., S 7975, S 7976 and the type), and the Bau
limestone hills at Tiang Bekap, Kuching district (e.g., S 32684).
Ecology. In mixed dipterocarp forest on deep yellow clay loams, at altitudes below 300 m,
apparently not common. Vulnerable.
7. Shorea amplexicaulis P.S.Ashton Fig. 20.

(Latin, amplexicaulis = stemclasping; the petiole)
(sect. Pachycarpae, red meranti)
al. (eds.) op. cit. 74; Newman et al. op. cit. (1996) 116. Type: Ashton BRUN 3261, Borneo, Brunei,
Andulau FR (holotype K; isotype KEP).
Mediumsized emergent tree, to 50 m tall, to 1.2 m diameter, with stout low buttresses.
Bark pale greyish brown, remaining smooth, hoopmarked; inner bark pale pinkish brown.
Heartwood pink. All exposed fleshy parts typically glabrous, but frequently sparsely
caducous or persistently pale buff to golden pubescent. Twigs slightly compressed at first,
to 3 × 2 mm apically, with prominent pale horizontal amplexicaul stipule scars. Leaf buds
hastate, obtuse, 10–15 × 3–5 mm. Stipules hastate, subacute, to 25 × 8 mm, not early
caducous. Leaves coriaceous, drying dark crimson red often shiny below, paler pinkish
brown above; blade elliptic, 11–21 × 5–8 cm, base cuneate, apex with tapering acumen to 1
cm long; midrib evident, flat above, terete and prominent below; lateral veins 9–12 pairs,
arched, prominent below; intercostal venation remotely scalariform, distinct and slightly
elevated below; petiole distinctly geniculate, 1.5–2.5 cm long. Inflorescences terminal or
axillary; rachis lax, slender, compressed, to 24 cm long, singly branched, branchlets bearing
to 11 distichous flowers; bracteoles lanceolate, to 13 × 8 mm, fugaceous. Flowers: buds to
10 × 3.5 mm; petals pale yellow; anthers narrowly oblong, connectival appendage more
than 2x the length of anther; ovary ovoid, glabrous, without distinct stylopodium, style
columnar. Fruits impressed at base; calyx generally glabrescent, lobes unequal, 3 longer
lobes to 18 × 3 cm, tapering to 2 cm above the saccate base, 2 shorter ones to 13 × 0.8 cm,
similar at base. Nuts broadly ovoid, acute, to 3.7 × 2.5 cm, persistently golden buff
pubescent.
225
Vernacular names. Sabah—kawang bukit (preferred name). Sarawak—engkabang pinang

licin (preferred name), langgai (Iban).
Distribution. Endemic in Borneo, occurring throughout the island except for the
southwestern parts. In Sabah known from Beaufort, Sandakan and Tawau districts (e.g.,
SAN 15106, SAN 22690, SAN 27397, and SAN 62919) and in Sarawak from Belaga, Bintulu,
Kapit, Kuching, Lawas, Limbang, Miri, and Simunjan districts (e.g., S 164, S 10708, S
15840, S 23611, S 32240, and S 68171). Also occurring in Brunei (e.g., BRUN 3006 and S
1633) and Kalimantan (e.g., bb. 29663, Church & Mahyar 1753 and Jarvie and Ruskandi
5756).
Ecology. In mixed dipterocarp forest, especially on clayrich soils but also on sandy clay, at
altitudes to 700 m. Not vulnerable.
Notes. Closely related to S. beccariana which differs in its golden tomentum, generally dull
leaf undersurface and short ascending stipule scars. The two often occur together though S.
beccariana ranges onto more sandy soils.
8. Shorea andulensis P.S.Ashton

(of Andulau FR in Brunei)
Wood op. cit. 97; Burgess op. cit. 184; Anderson op. cit. (1980) 124; Coode et al. (eds.) op. cit. 74;
Newman et al. op. cit. (1996) 117. Type: Ashton BRUN 3263, Borneo, Brunei, Andulau FR (holotype
Mediumsized emergent tree, to 45 m tall, to 1.2 m diameter; crown pale from below,
diffuse, hemispherical; buttresses stout, to 1 m tall. Bark becoming cracked and patchily
flaky, pale pinkish brown mottled. Twig, inflorescence, bracteole, leaf bud, parts of flower
exposed in bud, stipule and petiole densely persistently yellowish buffpuberulent, more
sparsely so on venation below and midrib above. Twigs slender, at first compressed, 1.5–2
mm diameter apically. Leaf buds ovoid, compressed, 3–6 × 1.5–3 mm, subacute. Stipules
hastate, subacute, to 14 × 5 mm. Leaves thinly coriaceous, cream lepidote in mature tree,
otherwise drying rich reddish brown below, pinkish brown above; blade ellipticovate, 6–9
× 3–5 cm, base obtuse or broadly cuneate, margin undulate, apex with acumen to 0.8 cm
long; midrib narrowly furrowed above, slender but prominent below as also the veins;
lateral veins 10–13 pairs, straight and abruptly arched within margin; intercostal venation
slender, scalariform, sinuate; petiole 1–1.2 cm long. Inflorescences terminal or axillary;
rachis terete or slightly compressed, to 12 cm long, singly or doubly branched, branchlets
bearing to 10 flowers; bracteoles ovate, to 3 × 1.2 mm, subacute. Flowers: buds to 8 × 2.5
mm; calyx spreading; petals lilac; stamens 15, connectival appendage c. 3x the length of
anther; ovary small, ovoid, glabrous, stylopodium obscure, style c. 2x the length of ovary,
columnar, pubescent in basal half. Fruits: pedicels to 2 mm long; calyx tube glabrescent,
lobes unequal, 3 longer lobes lorate, to 5 × 0.7 cm, base saccate, 2 shorter ones to 3 cm
long, otherwise similar. Nuts ovoid, to 3.5 × 2.5 cm, apiculate.
Vernacular names. Sabah—seraya daun merah (preferred name). Sarawak—meranti daun

putih (preferred name).
226
Fig. 20. Shorea amplexicaulis. A, flowering leafy twig; B, adaxial view of bract; C, abaxial
view of bract; D, flower bud; E, flower bud with a few sepals and petals removed; F, adaxial
view of inner sepal; G, adaxial view of outer sepal; H, abaxial view of petal; I, adaxial view
of petal with 3 stamens; J, adaxial view of stamens; K, abaxial view of stamens; L, fruit. (A
from S 65194, B–K from S 29202, L from S 23611.)
227
Distribution. Endemic in Borneo. In Sabah known from Kinabatangan and Labuk Sugut
districts (e.g., SAN 23264, SAN 99663 and SAN 131981) and in Sarawak from Bintulu,
Kapit, Kuching, Lawas, and Limbang districts (e.g., S 18408, S 22338, S 29580, S 32368,
and S 44085). Also occurring in Brunei (e.g., BRUN 3030 and BRUN 5435) and W and E
Kalimantan.
Ecology. Local and rare in mixed dipterocarp forest on yellow sandy clay soils, at altitudes
to 600 m. In Sabah the species occurs in forest on ultrabasic substrates. Vulnerable.
9. Shorea angustifolia P.S.Ashton

(Latin, angustus = narrow, folius = leaf; with narrow leaves)
(sect. Richetioides subsect. Richetioides, yellow meranti)
Newman et al. op. cit. (1996) 118. Type: Ashton BRUN 778, Borneo, Brunei, Kuala Temburong
Macang (holotype K; isotypes KEP, L).
Main canopy tree, to 40 m tall, to 90 cm diameter, with frequently misshapen bole and low
rounded buttresses. Bark becoming greyish tawny, thinly irregular flaky. Twig endings and
stipules sparsely caducous pale buffbrown puberulent; leaf bud, parts of flower exposed in
bud, ovary and nut, inflorescence, bracteoles and petiole persistently so. Twigs to 2 mm
diameter apically, straight, terete, superficially cracked, rugose. Leaf buds small, ovoid,
obtuse, to 0.5 × 1 mm. Stipules linear, to 3 mm long, fugaceous. Leaves thinly coriaceous,
drying pale greenish brown; blade deep violet when opening, ovate to oblonglanceolate, 8–
14 × 2.5–4 cm, base cuneate, apex with tapering acumen to 1.5 cm long; midrib flat above,
hardly elevated below as also the veins; lateral veins 8–10 pairs, wellspaced, arched;
intercostal venation subscalariform; petiole 0.6–0.8 cm long, finely cracked and drying
distinct pale brown to creamgrey. Inflorescences terminal or axillary; rachis slender, lax,
terete, to 10 cm long, singly or doubly branched, branchlets bearing to 8 flowers; bracteoles
small, fugaceous. Flowers: buds to 2.5 × 1.5 mm, small; petals cream; stamens 15,
connectival appendage 1–2x the length of anther; ovary and stylopodium conical, style
short, glabrous. Fruits: calyx lobes subequal, broadly ovate, obtuse, to 0.5 × 0.5 cm,
incrassate and clasping the nut. Nuts obovoid, to 0.8 × 0.9 cm, acute.
Vernacular names. Sabah—seraya kuning bukit (preferred name). Sarawak—lun bukit

(preferred name).
Distribution. Endemic in Borneo. In Sabah known from Kota Kinabalu (Gaya Is.), Lahad
Datu and Ranau (G. Kinabalu) districts (e.g., KEP 80302, SAN 20704, SAN 24259, SAN
99630, and SAN 116969) and in Sarawak from Belaga, Kapit and Lawas districts (e.g., S
5682 and BRUN 5736) and W and E Kalimantan (e.g., bb. 29071, Kessler et al. Berau 888,
Kostermans 5412, and Newman et al. 588).
Ecology. Locally frequent on skeletal soils along shale ridges towards the upper limits of
mixed dipterocarp forest, at 500–1200 m altitude. Occurring in Kinabalu and Mulu NPs;
elsewhere becoming vulnerable.
228
10. Shorea argentifolia Symington Fig. 21.

(Latin, argenteus = silvery, folius = leaf; the silvery leaf undersurface)
(sect. Mutica subsect. Mutica, red meranti)
Gard. Bull. Sing. 17 (1960) 489; Ashton op. cit. (1964) 179, op. cit. (1968) 105, op. cit. (1982) 539;
Meijer & Wood op. cit. 98; Burgess op. cit. 166; Anderson op. cit. (1980) 124; PROSEA op. cit. 392;
Coode et al. (eds.) op. cit. 75; Newman et al. op. cit. (1996) 119. Type: Maidin FMS 38838, Borneo,
Sabah, Tawau (holotype KEP).
Emergent tree, to 45 m tall, to 1.5 m diameter, with pale goldensuffused crown with
pendent twig endings; buttresses to 1.5 m tall, stout. Bark becoming vsection fissured,
pinkish brown; inner bark and heartwood pinkish brown. Twig endings, inflorescence, leaf
bud, parts of flower exposed in bud, nut, stipule, bracteole, petiole, leaf blade below, and
midrib above persistently densely pinkish golden velutinous; fruit calyx shortly sparsely
pubescent. Twigs at first compressed, c. 2 × 1 mm apically, sparingly branched; stipule
scars obscure. Leaf buds ovoid, slightly compressed, subacute, c. 4 × 2.5 mm. Stipules
oblonghastate, to 20 × 7 mm, acute, cordate, not early caducous. Leaves thinly
coriaceous, drying pinkish brown below, pale mauvebrown above; blade oblong
lanceolate, 6–11 × 2.5–4.5 cm, base obtuse, apex with acumen c. 0.5 cm long; midrib
narrowly sunken above, terete, slender but prominent below as also the veins; lateral veins
20–25 pairs, dense with short intermediates; intercostal venation slender, densely
scalariform; petiole 0.8–1 cm long, relatively short, rugose. Inflorescences terminal or
axillary; rachis compressed, to 12 cm long, singly branched, branchlets bearing to 6 flowers;
bracteoles oblongelliptic, obtuse, to 8 × 3.5 mm, caducous. Flowers: buds to 6 × 3 mm;
petals pink; stamen 15, connectival appendage short, hardly or not reflexed; ovary and
stylopodium pyriform, shortpubescent, style as long as ovary, glabrous. Fruits: calyx lobes
unequal, 3 longer lobes to 8 × 1.2 cm, tapering to 4 mm above the saccate base, 2 shorter
ones unequal, linearlobed, to 3 × 0.15 cm, similar at base. Nuts ovoid, to 1.4 × 0.9 cm, with
to 2.5 mm long acicular style remnant.
Vernacular names. Sabah—seraya daun perak (preferred name). Sarawak—meranti

binatoh (preferred name).
Distribution. Endemic in Borneo. Known in Sabah from Beaufort, Keningau,

Kinabatangan, Lahad Datu, Labuk Sugut, Ranau, Sandakan, and Tawau districts (e.g, SAN
16976, SAN 23972, SAN 55561, SAN 77288, and SAN 99224) and in Sarawak from Belaga,
Bintulu, Kapit, Lawas, Limbang and Miri districts (e.g., S 15564, S 29660, S 32224, and S
69132). Also occurring in Brunei (e.g., BRUN 126, FMS 28661 and FMS 30534) and NE
Kalimantan (pers. obs.).
Ecology. Locally frequent in mixed dipterocarp forest on clay rich alluvium soils, on hill
slopes and sometimes ridges, at altitudes to 900 m. Occurring in Lambir and Mulu NPs;
elsewhere vulnerable owing to forest conversion.
229
11. Shorea asahii P.S.Ashton

(Asah Unyong, c. 1938–, Sarawakian tree climber and dendrologist extraordinaire)
(sect. Shorea subsect. Barbata, selangan batu)
Wood op. cit. 163; Burgess op. cit. 202; Anderson op. cit. (1980) 118; Coode et al. (eds.) 75; Newman
et al. op. cit. (1998) 169. Type: Flemmich FMS 37183, Borneo, Brunei, Tutong district, Bt. Bedawan
(holotype KEP).
Low emergent tree, to 40 m tall, to 90 cm diameter, with frequently misshapen bole and low
thin buttresses to 80 cm tall. Bark surface becoming dark greyish brown mottled, cracked
and thinly flaky. Young parts at first shortly puberulent, otherwise glabrous but for densely
persistently creambuff puberulent inflorescence; parts of flower exposed in bud, ovary, nut,
and fruit calyx sparsely so. Twigs c. 0.7 mm diameter apically, slender, muchbranched,
terete. Leaf buds falcate, to 1.5 × 1 mm, small. Stipules narrowly deltoid, to 5 mm long,
fugaceous. Leaves coriaceous, lustrous; blade ovate, 6–10 × 3–5 cm, base subequal,
broadly cuneate, apex with slender acumen c. 1 cm long; midrib hardly raised; lateral veins
6–7 pairs, arched, slender, unraised on either surface; intercostal venation densely
scalariform, unraised, obscure; petiole c. 1 cm long, slender. Inflorescences terminal or
axillary; rachis slender, terete, to 2 cm long, singly branched, branchlets zigzag, bearing to
4 close flowers; bracteoles elliptic, to 2 mm, fugaceous. Flowers: buds globose, to 2 mm
diameter; stamens c. 30, filaments, anthers and connectival appendage sparsely barbate
distally, appendage to 2x the length of anther; ovary and stylopodium ovoid, densely
pubescent, style short, slender, glabrous. Fruits: calyx lobes subequal, ovate, to 0.4 × 0.4
cm, subacute, appressed to nut. Nuts oblongovoid, to 1.1 × 0.6 cm, acute.
Vernacular names. Sabah—selangan batu asah (preferred name). Sarawak—kumus bukit

(Iban).
Distribution. Endemic in Borneo. In Sabah known from the Crocker Range (e.g., SAN
24279) and in Sarawak from Kapit, Limbang, Lubok Antu, and Miri districts (e.g., SA 690,
S 29561, S 41410, and S 57238). Also occurring in Brunei (e.g., BRUN 772, BRUN 5632
and S 1183) and W Kalimantan (e.g., bb. 35219 and bb. 35220).
Ecology. Very rare, in the lower limits of upper dipterocarp forest, on thinly organic skeletal
soils over shale and sandstone on minor peaks and spurs, usually at 650–850 m altitude.
Recorded from Lambir NP, elsewhere endangered.
12. Shorea atrinervosa Symington Fig. 22.

(Latin, ater = dull black, nervosus = nerved; the leaf venation drying black)
(sect. Shorea, subsect. Shorea, selangan batu)
Gard. Bull. S. S. 10 (1939) 363, op. cit. (1943) 9; Masamune op. cit. 492; Browne op. cit. 167; Meijer
& Wood op. cit. 163; Burgess op. cit. 202, 210; Ashton op. cit. (1968) 68, op. cit. (1982) 457;
cit. (1998) 170. Type: Alwi FMS 31832, Peninsular Malaysia, Pahang, Bt. Goh FR (holotype KEP).
230
Fig. 21. Shorea argentifolia. A, fruiting leafy twig; B, detail of indumentum on the lower
leaf surface; C, young shoot with stipules, bracts and young inflorescence; D, flower bud; E,
flower bud with exposed gynoecium; F, gynoecium and stamens; G, adaxial view of outer
sepal; H, adaxial view of inner sepal; I, abaxial side of petal; J, adaxial side of petal; K,
adaxial view of stamens; L, abaxial view of stamens; M, fruit. (A–B from S 15564, C–L
from SAN 55561, M from SAN 99224.)
231
Fig. 22. Shorea atrinervosa. A, flowering leafy twig; B, detail of venation on lower leaf
surface; C, detail of indumentum on midrib; D, open flower; E, adaxial view of outer sepal;
F, adaxial view of inner sepal; G, abaxial side of petal; H, adaxial side of petal with
stamens; I, adaxial view of stamens; J, abaxial view of stamens; K, longitudinal section of
open flower; L, fruit. (A–C from SAN 15091, D–K from SAN 27274, L from SAN 16957.)
232
Large emergent tree, to 50 m tall, to 1.2 m diameter, with pale diffuse somewhat irregular
crown; bole tall and straight; buttresses to 3 m tall, thin, spreading. Bark tawnybrown,
irregularly oblongflaky. Bud, twig and petiole fugaceous puberulent; stipules,
inflorescences, parts of perianth exposed in bud, ovary, and nut, persistently so; parts
otherwise glabrous. Twigs c. 2 mm diameter apically, terete, drying black, becoming
ribbed. Leaf buds narrowly ovoid, acute, to 4 × 2 mm. Stipules lanceolate, to 8 × 3 mm
acute, caducous. Leaves thinly coriaceous, undulate, more or less white lepidote below (in
mature trees) with the veins drying distinct black; blade elliptic to ovate, frequently of
irregular shape, 8–16 × 3.5–9 cm, base broadly cuneate to subcordate, apex with tapering
acumen to 1 cm long; midrib elevated above, prominently so below; lateral veins 10–12
pairs, stout and elevated below, often somewhat sinuous, sometimes with small glabrous
domatia; intercostal venation scalariform, slender, sinuous; petiole at first whitish lepidote,
drying black, 1.2–2.2 cm long, geniculate. Inflorescences terminal or axillary; rachis
straight, terete, to 11 cm long, singly branched, branchlets bearing to 6 secund flowers;
bracteoles elliptic, to 3 × 2 mm, acute. Flowers: buds to 8 × 3 mm; petals cream with
crimson median stripe towards base; stamens 25–33, filaments sparsely setose, outer anther
locules glabrous, connectival appendage as long as anther, setose; ovary and stylopodium
ovoid, pubescent, surmounted by a short, glabrous style. Fruits: pedicels to 4 mm long,
broadening into fruit base; calyx lobes unequal, longer lobes to 11 × 2.5 cm, tapering to 1
cm above the saccate base, 2 shorter ones loratelobed, to 8.5 × 0.9 cm. Nuts ellipsoid, to
2.5 × 2 cm, tapering into to 8 mm stout style remnant.
Vernacular name. Sabah and Sarawak—selangan batu hitam (preferred name).
Distribution. Peninsular Malaysia, Sumatra and Borneo. In Sabah known from Beaufort,
Kinabatangan, Labuk Sugut, Lahad Datu, Ranau, Sandakan, Sipitang, Tawau, and Tenom
Sarawak from Bintulu, Kapit, Kuching, Lawas, Lundu, Marudi, Mukah, Simunjan, and
Tatau districts (e.g., S 15215, S 19570, S 22830, S 24698, and S 57601). Also occurring in
Brunei (e.g., BRUN 3337 and BRUN 3342) and W and E Kalimantan (e.g., bb. 23517,
Kessler et al. Berau 793 and Kostermans 13890).
Ecology. Scattered, locally frequent in mixed dipterocarp forest on clayrich soils on low
hills and moist valleys, at altitudes to 600 m. Occurring in Lambir and Mulu NPs; elsewhere
vulnerable.
13. Shorea bakoensis P.S.Ashton

(of Bako NP, Sarawak)
Newman et al. op. cit. (1996) 121. Type: Brunig S 17502, Borneo, Sarawak, Bako NP (holotype K).
Subcanopy or low canopy tree, recorded to 7 m tall, to 5 cm diameter. Bark smooth. All
observed parts glabrous but for the sericeous fruit calyx outside, and densely buffpubescent
nut. Twigs c. 2 mm diameter apically, terete, drying black. Buds and stipules unknown.
Leaves thickly coriaceous, drying dark tawny brown, shiny below; blade narrowly oblong
to lanceolate, 13–18 × 5–6 cm, base obtuse, apex with slender acumen to 1 cm long; midrib
evident but flat above, prominent below; lateral veins 9–10 pairs, prominent below;
233
intercostal venation subscalariform, hardly raised below; petiole 1–1.2 cm long, stout,
drying black. Inflorescences and flowers unknown. Fruits: pedicels short; calyx lobes
subequal, oblong, obtuse, incrassate, clasping the nut. Nuts ellipsoidcylindric, to 2.5 × 1.4
cm, apiculate.
Distribution. Endemic in Borneo; rare and localised, known only by a single collection (the
type) from Bako NP but also observed at G. Santubong, Kuching district, Sarawak, at
altitudes below 200 m.
Ecology. In kerangas forest on shallow podsols on sandstone ridges and plateaux. Rare and
possibly endangered.
14. Shorea balanocarpoides Symington

(Greek, resembling Balanocarpus; the fruit resembles that of Neobalanocarpus, a genus
endemic in Peninsular Malaysia)
Gard. Bull. S. S. 9 (1938) 330, op. cit. (1943) 47; Slooten, Reinwardtia 3 (1956) 340; Ashton op. cit.
(1982) 475; PROSEA op. cit. 418; Coode et al. (eds.) op. cit. 75; Newman et al. op. cit. (1996) 123.
Type: Lambak FMS 15768, Peninsular Malaysia, Pahang, Rompin (holotype KEP). Synonyms:
Balanocarpus pahangensis Foxw., Malay. For. Rec. 10 (1932) 145; Shorea dolichocarpa Slooten op.
cit. (1956) 342, Ashton op. cit. (1964) 151, op. cit. (1968) 83, Anderson op. cit. (1980) 121.
Main canopy tree, to 40 m tall, to 80 cm diameter; bole frequently misshapen; buttresses to

1 m tall, stout. Bark tawny brown, cracked and irregularly flaky. Young twig, leaf bud,
stipule outside, inflorescence, and flower calyx outside sparsely greyish puberulent; parts of
petals exposed in bud, ovary and stylopodium persistently so; nut densely mauvegrey
pubescent. Twigs c. 1.5 mm diameter apically, terete, muchbranched. Leaf buds small,
globose to ovoid, obtuse, to 2 × 1.5 mm. Stipules linear, fugaceous, to 4 mm long. Leaves
coriaceous, drying tawnybrown; blade ovate, 6–12 × 2–7 cm, base obtuse to cuneate
(narrowly peltate in juveniles), decurrent to 1.5 mm down petiole, usually unequal, apex
with slender acumen, to 1 cm long; midrib stout on both surfaces, somewhat elevated above,
raised below; lateral veins 5–7 pairs, arched, widely spaced, raised below; intercostal
venation more or less reticulate to scalariform towards margin, distant, elevated below;
petiole 1.2–2 cm long. Inflorescences terminal or axillary; rachis slender, to 8 cm long,
somewhat irregularly doubly branched, branchlets often zigzag, bearing to 8 distichous
flowers; bracteoles minute, fugaceous. Flowers: buds to 4.5 × 1.2 mm; petals yellowish
brown with bright yellow margin; stamens 15, connectival appendage ciliate towards apex;
ovary and stylopodium subglabrous. Fruits subsessile; calyx lobes subequal, deltoid, c. 0.5
× 0.5 cm, subrevolute at margins, appressed to the nut. Nuts ellipsoid to obovoid, to 3 × 1.3
cm, acute, mauvegrey pubescent.
Vernacular names. Sarawak—barek (Iban), lun gondol (preferred name).
234
Distribution. Sumatra, Peninsular Malaysia and Borneo. Not yet reported from Sabah. In
Sarawak known from Belaga, Bintulu, Kapit, Limbang, and Miri districts (e.g., S 15139, S
BRUN 3365).
Ecology. Local, but there often common, in mixed dipterocarp forest, on clay and sandy
clay soils, particularly on slopes, at altitudes to 700 m. Vulnerable.
15. Shorea beccariana Burck

Ann. Jard. Bot. Buitenz. 6 (1887) 213; Merrill, EB (1921) 404; Browne op. cit. 138; Ashton, Gard.
Bull. Sing. 20 (1963) 280, op. cit. (1964) 180, op. cit. (1968) 105, op. cit. (1982) 525; Meijer & Wood
op. cit. 100; Burgess op. cit. 154, 182; Anderson op. cit. (1980) 125; PROSEA op. cit. 393; Coode et
al. (eds.) op. cit. 75; Newman et al. op. cit. (1996) 124. Lectotype (designated here): Beccari PB
1127, Borneo, ‘Sarawak’ (hololectotype BO). Synonym: Shorea franchetiana F.Heim, op. cit. (1891)
956.
Emergent tree, to 60 m tall, to 1.1 m diameter; crown often goldensuffused from below;
bole tall, often somewhat sinuate; buttresses to 1.5 m tall, stout. Bark smooth, hoop
marked, occasionally somewhat flaky in large trees; inner bark pinkish brown; heartwood
pink. Young twig, inflorescence, parts of perianth exposed in bud, leaf bud, stipule, and
petiole evenly densely rufousbrown puberulent, turning mauvegrey, sparsely so on
venation below; nut buff pubescent. Twigs compressed, 3–6 × 1–2.5 mm apically, dull
greyish brown; stipule scars shortly cuneate to falcate, more or less ascending. Leaf buds
hastate, compressed, acute, 7–11 × 3.5–4.5 mm. Stipules oblong, obtuse, small, to 14 × 5
mm, caducous. Leaves coriaceous, golden lepidote turning to dull mauvegrey below (in
mature trees); blade elliptic to ovate, 11–20 × 5.5–7 cm, base obtuse or broadly cuneate,
apex with broad acumen to 0.8 cm long; midrib prominent below, evident albeit shallowly
sunken above; lateral veins 11–14 pairs, arched, slender but elevated below (more
prominent in immature trees); intercostal venation remotely scalariform; petiole 2–4 cm
long. Inflorescences terminal or axillary; rachis lax, more or less compressed, to 20 cm
long, singly branched, branchlets bearing to 8 distichous flowers; bracteoles oblong, obtuse,
to 15 × 9 mm, caducous. Flowers: buds to 9 × 3 mm; petals pink; stamens 15, connectival
appendage 2–3x the length of anther; stylopodium and style cylindrical. Fruits: base
frequently impressed; calyx glabrescent, lobes unequal, 3 longer lobes to 19 × 2.7 cm,
tapering to 1.2 cm above the saccate base, 2 shorter ones linearlobed, to 10.5 × 0.9 cm,
similar at base. Nuts broadly ovoid, to 4 × 2.8 cm; style remnant conical, to 4 mm long.
Vernacular names. Sabah—seraya langgai (preferred name). Sarawak—langgai (Iban),

meranti langgai (preferred name).
Distribution. Endemic in Borneo. Common and known in Sabah from Labuk Sugut,
Sandakan, Sipitang, Tawau, and Tenom districts (e.g., SAN 16023, SAN 16468, SAN 27397,
SAN 35898, and SAN 97628) and in Sarawak from Bintulu, Kapit, Kuching, Limbang,
Lundu, and Miri districts (e.g., S 10125, S 12612, S 27129, S 32308, and S 46476). Also
occurring in Brunei (e.g., BRUN 418, BRUN 3187, Niga NN 225, S 1933, and S 2172) and
W and NE Kalimantan (e.g., bb. 27726, bb. 30208, Newman et al. 585, and Suzuki K 9494).
235
Ecology. One of the commonest trees in mixed dipterocarp forest on welldrained sites, on
yellow sandy and sandy clay soils, and on welldrained shale ridges, at altitudes to 700 m.
Occurring in Bako, Lambir and Mulu NPs; not vulnerable.
Notes. For comparison with S. amplexicaulis, see there.
16. Shorea biawak P.S.Ashton

(biawak = the monitor lizard, whose skin the bark resembles)
(sect. Shorea subsect. Barbata, selangan batu)
Gard. Bull. Sing. 19 (1962) 281, op. cit. (1964), op. cit. (1968) 68, op. cit. (1982) 464; Meijer & Wood
op. cit. 165; Burgess op. cit. 202; Anderson op. cit. (1980) 118; Coode et al. (eds.) op. cit. 75;
Newman et al. op. cit. (1998) 171. Type: Ashton BRUN 3369, Borneo, Brunei, Bangar (holotype K;
isotype KEP).
Main canopy tree, to 35 m tall, to 70 cm diameter, with straight bole and low thin buttresses.
Bark overall remaining smooth, but patterned by dense minute zigzag vsection surface
fissures. Twigs c. 1 mm diameter apically, slender, terete, muchbranched, smooth and
striated. Young twig, leaf bud, parts of perianth exposed in bud, ovary, nut, inflorescence,
and petiole more or less densely pale buffpuberulent, persistent only on inflorescence, bud
and ovary. Leaf buds to 1 mm long, small. Stipules unknown. Leaves thinly coriaceous,
drying pale greyish brown; blade ovate, 6–10 × 2.5–4.5 cm, base obtuse or broadly
cuneate, apex subcaudate, acumen to 1.5 cm long; midrib slender, somewhat elevated
below, flat or slightly elevated above; lateral veins slender, 5–6 pairs, arched and
ascending, elevated below, slightly furrowed above, with small puberulent domatia;
intercostal venation densely scalariform, not raised; petiole 0.7–1 cm long, slender.
Inflorescences terminal or axillary; rachis terete, to 14 cm long, singly branched, branchlets
densely bearing to 6 flowers; bracteoles ovate, acute, to 2 mm long, fugaceous. Flowers:
buds globose, to 1.5 mm diameter; petals creamwhite; stamens c. 35, filaments glabrous,
anthers sparsely barbate apically, connectival appendage slightly shorter than anther,
densely barbate; ovary and stylopodium ovoid to pyriform, densely pubescent, style short,
glabrous. Fruits: calyx lobes subequal, suborbicular, obtuse, appressed to nut, to 0.7 × 0.9
cm. Nuts subglobose, to 1.3 × 1 cm, obtuse with apiculate style remnant to 1.5 mm long.
Vernacular names. Sabah—selangan batu biawak (preferred name). Sarawak—resak

biawak (Iban), selangan batu biawak (preferred name).
Distribution. Endemic in Borneo. Known in Sabah from Beaufort district (e.g., SAN 15068,
SAN 24825 and SAN 26567) and in Sarawak from Bintulu, Kapit and Miri districts (e.g., S
14453, S 28778 and S 31711). Also occurring in Brunei (e.g., BRUN 3005 and BRUN 3351).
Ecology. Widespread but apparently rare, easily overlooked, in mixed dipterocarp forest on
clay soils, at altitudes to 600 m. Present in Lambir NP; elsewhere vulnerable.
17. Shorea bracteolata Dyer

(Latin, bracteolatus = with bracteoles; the persistent bracteoles of the inflorescence)
236
Fl. Brit. Ind. 1 (1874) 305; Masamune op. cit. 493; Slooten, Bull. Jard. Bot. Buitenz. 3, 18 (1949) 259;
Browne op. cit. 158; Ashton op. cit. (1964) 163, op. cit. (1968) 93 & 496, op. cit. (1982) 496; Meijer
& Wood op. cit. 53; Burgess op. cit. 159; Anderson op. cit. (1980) 123; PROSEA op. cit. 410; Coode
et al. (eds.) op. cit. 75; Newman et al. op. cit. (1996) 125. Type: Maingay 204, Peninsular Malaysia,
Malacca (holotype K; isotype L).
Emergent tree, to 50 m tall, to 1.2 m diameter; bole straight; crown large, hemispherical,
dense; buttresses to 1.2 m tall, stout. Bark surface pale chocolatebrown, becoming cracked
and rather thinly irregularly flaky; inner bark laminated pale and dark brown. All parts at
first sparsely pale brown puberulent, surfaces waxy scurfy; glabrescent except on
inflorescence, buds and ovary. Twigs c. 2 mm diameter apically, somewhat compressed.
Leaf buds shortly falcate, compressed, subacute, 3–5 × 2 mm. Stipules linear, c. 10 mm
long, fugaceous. Leaves chartaceous or thinly coriaceous, drying dull greyish brown; blade
oblongovate to elliptic, 9–14 × 4–6 cm, base obtuse, apex with tapering acumen to 1.5 cm
long; midrib obscurely sunken above, slender but prominent below; lateral veins slender,
12–15 pairs, arched; intercostal venation distantly scalariform to subreticulate; petiole 1–2
cm long, slender. Inflorescence terminal or axillary; rachis slender, straight, terete, to 10 cm
long, singly branched, branchlets bearing to 5 flowers; bracteoles lanceolate, to 12 × 5 mm,
not early caducous. Flowers: buds to 10 × 5 mm; petals pale yellow tinged pink at base
within; stamens 15, anthers narrowly oblong, connectival appendage 3–4x the length of
anther, scabrous; ovary ovoid, scabrous apically without stylopodium, style c. 2x the length
ovary. Fruits: calyx glabrescent; lobes unequal, 3 longer lobes to 10 × 1.7 cm, tapering to 4
mm above the saccate base, 2 shorter ones unequal, to 8 × 0.7 cm. Nuts narrowly ovoid, to
2 × 1 cm, with filiform style remnant to 4 mm long.
Vernacular names. Sabah—melapi pa’ang (preferred name). Sarawak—meranti pa’ang

(preferred name).
Distribution. Sumatra, Peninsular Malaysia, Singapore, and throughout Borneo. In Sabah

known from Lahad Datu and Tawau districts (e.g., SAN 17832, SAN 22651, SAN 31227,
SAN 39093, and SAN 54885) and in Sarawak from Kuching, Lawas and Lundu districts
(e.g., S 10705, S 15199, S 32443, and S 40580). Also occurring in Brunei (e.g., BRUN 907
and Dransfield JD 6662) and Kalimantan (e.g., bb. 14612, bb. 27691, Kostermans 6354, and
Kostermans 13329).
Ecology. In mixed dipterocarp forest on welldrained clay and sandy soils. Mostly scattered
and often uncommon, but locally frequent on coastal hills, at altitudes to 700 m. Vulnerable.
18. Shorea brunnescens P.S.Ashton

(Latin, brunnescens = brownish; the dry leaf)
Newman et al. op. cit. (1998) 172. Type: Smythies S 15218, Borneo, Sarawak, Simunjan districts, Bt.
Gaharu (holotype K; isotypes KEP, L).
Emergent tree, to 45 m tall, to 1.2 m diameter; bole straight, buttresses to 1.2 m tall, thin.
Bark becoming vertically cracked and thinly oblongflaky, pale tawnybrown. Leaf bud,
parts of perianth exposed in bud, ovary, and nut densely more or less persistently buff
pubescent; inflorescence, young twig and petiole sparsely caducously so. Twigs c. 1 mm
237
diameter apically, terete. Leaf buds small, ovoid, acute, to 2.5 × 2 mm. Stipules not seen.
Leaves coriaceous, drying pale ochreousbrown; blade broadly ovate to lanceolate, 6–12 ×
2.5–6 cm, base broadly cuneate, apex with narrow acumen to 1 cm long; midrib obscure,
furrowed above, slender but prominent below; lateral veins 9–11 pairs, slender, arched,
barely elevated below; intercostal venation dense, subreticulate, hardly raised but evident
below; petiole 1–1.5 cm long, subgeniculate. Inflorescences terminal or axillary; rachis
angular, to 9 cm long, singly branched; bracteoles not seen. Flowers: buds to 4 × 2 mm;
stamens 46–62, filaments and anthers glabrous, connectival appendage shorter than anther,
glabrous but for 1–2 apical setae; ovary and stylopodium pyriform. Fruits: calyx lobes
glabrescent, unequal, 3 longer lobes to 6.5 × 1 cm, tapering to 3 mm above the saccate
base, 2 shorter ones to 3 × 0.3 cm, similar at base. Nuts ovoidellipsoid, to 1.2 × 0.7 cm,
with to 3 mm tapering filiform style remnant.
Vernacular name. Sarawak—selangan batu tinteng (preferred name).
Distribution. Endemic in Borneo. In Sabah known from Kota Belud, Kuala Penyu, Ranau
and Sipitang districts (e.g., SAN 16360, SAN 22088, SAN 22406, SAN 50947, and SAN
72276) and in Sarawak from Julau, Kapit, Kuching, Lubok Antu, Lundu, Miri, Samarahan,
Sri Aman, and Tatau districts (e.g., S 15220, S 29623, S 36924, S 37708, and S 41374). Also
occurring in Brunei (e.g., S 1183) and W and SE Kalimantan (e.g., Suzuki K 3565).
Ecology. Locally frequent in mixed dipterocarp forest on leached yellow sandy clay soils
and in upper dipterocarp forest, at altitudes to 1400 m. Occurring in Kinabalu and Mulu
NPs; probably not yet vulnerable.
19. Shorea bullata P.S.Ashton

(Latin, bullatus = blistered; the leaf blade)
Gard. Bull. Sing. 19 (1962) 283, op. cit. (1964) 181, op. cit. (1968) 105; op. cit. (1982) 517; Anderson
op. cit. (1980) 125; Coode et al. (eds.) op. cit. 75; Newman et al. op. cit. (1996) 126. Type: Ladi &
Angga BRUN 2003, Borneo, Brunei, Bangar (holotype K; isotypes KEP, L).
Low emergent tree, to 45 m tall, to 1.2 m diameter; bole straight; crown hemispherical with
golden tinge from below; buttresses stout, to 1 m tall. Bark smooth, hoopmarked at first,
mauvegrey and chocolatebrown mottled, becoming vertically cracked and patchily oblong
flaky; inner bark and heartwood crimson. Twig, outside of stipules, inflorescence, petiole
and venation below persistently scabrid fulvouspubescent; leaf bud, parts of perianth
exposed in bud and leaf midrib above densely evenly so; blade above sparsely caducous
hispid; ovary and nuts densely creambrown pubescent. Twigs c. 2 mm diameter apically,
terete, muchbranched, becoming papery flaky. Leaf buds broadly ovoid, 3–4 × 2–3 mm.
Stipules narrowly hastate, acute, to 7 × 2 mm. Leaves chartaceous, becoming concave and
prominently bullate between the intercostal veins, drying dark reddish brown; blade elliptic
to oblongovate, 6.5–10 × 3–4.5 cm, base narrowly obtuse or broadly acuminate, margin
usually revolute, apex obtuse or shortly broadly acuminate; midrib narrowly furrowed
above, prominent below as also lateral veins; lateral veins 10–12 pairs; intercostal venation
distantly scalariform; petiole 0.8–1 cm long. Inflorescences terminal or axillary; rachis
terete, lax, to 11 cm long, singly branched, branchlets bearing to 8 flowers; bracteoles
broadly elliptic, obtuse, to 6 × 3.5 mm. Flowers: buds to 15 × 3 mm; petals cream with pink
238
at base; stamens 15, connectival appendage c. 4x the length of anther, slender, twisted
towards apex; ovary and stylopodium ovoid, densely shortpubescent, style almost 2x the
length of ovary, columnar, pubescent in basal half. Fruits: calyx lobes unequal, 3 longer
lobes to 9 × 1.3 cm, tapering to 5 mm above the saccate base, 2 shorter ones linearlobed,
to 3 × 0.3 cm, with a similar base. Nuts ovoid, to 1 × 0.7 cm, acute.
Vernacular name. Sarawak—meranti melecur (preferred name).
Distribution. Endemic in Borneo. Known in Sarawak from Bintulu, Kapit, Kuching (sterile
collection), Miri, and Tatau districts (e.g., S 18378, S 27138 and S 63844). Also recorded
from Brunei (e.g., BRUN 924, BRUN 5773 and BRUN 5798).
Ecology. Rare, in scattered populations in mixed dipterocarp forest on yellow sandy clay
soils, on low hills, at altitudes to 400 m. Occurring in Lambir and Mulu NPs; elsewhere
endangered.
20. Shorea calcicola P.S.Ashton

(Latin, calx = limestone, colere = to grow; the natural habitat)
(sect. Shorea, subsect. Shorea)
TFSS 5 (2004) 479. Type: Murthy & Chai S 24690, Borneo, Sarawak, Kuching district, Kuching
Serian Road (holotype KEP; isotype SAR).
Mediumsized tree, occasionally to 50 m tall, to 1.2 m diameter, with low thin buttresses.
Bark chocolatebrown, becoming flaky. Inflorescence, flower calyx, twig and petiole at first
buff puberulent, glabrescent; midrib above, parts of petal exposed in but, ovary and nut
densely persistently buff pubescent. Twig c. 2 mm diameter apically, terete, smooth, dark
brown. Stipules unknown. Leaves thickly coriaceous, somewhat lustrous and drying
orange brown below, dark redbrown above; blade broadly ovate 10–14 × 5–7 cm; lateral
veins c. 9 pairs, slender but raised below, evident but hardly raised above as also the
midrib; intercostal venation densely scalariform, evident but hardly raised; petiole 2–3 cm
long, distinctly geniculate. Complete inflorescences unknown. Flowers: buds to 7 × 3 mm,
fusiform; stamens 25–28, filaments and anther locules glabrous, connectival appendages
surmounted by a single apical seta; ovary and stylopodium broadly pyriform. Fruits:
pedicels c. 2 mm long, slender; 3 longer calyx lobes to 6 × 1.5 cm, broadly spathulate,
obtuse, tapering to c. 4 mm broad above the saccate base, 2 shorter lobes to 2 cm long,
linear, similar at base. Nuts ovoid, to 1.8 × 1.3 cm, tapering to a distinct apiculus.
Distribution. Endemic in Borneo. Not yet known in Sabah. In Sarawak recorded from
Kuching and Serian districts (e.g., S 12582, S 22777 and S 28041). One tentative collection
from G. Subis, Miri district (i.e., S 27282) and unconfirmed records (sterile) from the Mulu
limestone hills.
Ecology. Locally quite common on organic soils over limestone, at altitudes below 400 m.
Not vulnerable.
239
21. Shorea carapae P.S.Ashton

(from Iban word—karapa = swampy pole forest; the habitat)
(sect. Brachypterae, subsect. Brachypterae, red meranti)
Newman et al. op. cit. (1996) 127. Type: Kostermans 13095, Borneo, Kalimantan, W Kutei, Mt.
Palimasan near Tabang, Ulu Belayan (holotype K; isotypes KEP, L).
Mediumsized canopy tree, to 35 m tall, to 65 cm diameter; bole frequently crooked;

buttresses to 1 m tall but usually lower, stout. Bark becoming cracked and patchily flaky,
chocolate and greyish brown mottled; inner bark orangebrown; heartwood yellowbrown.
Twig, leaf bud, stipule, petiole, and parts of corolla exposed in bud persistently densely
evenly buffsericeous; venation below and midrib above sparsely so; fruit calyx and pedicel
sparsely puberulent; nuts glabrous. Twigs compressed, c. 3 × 2 mm apically, stipule scars
prominent, almost amplexicaul. Leaf buds lanceolate, compressed, acute, to 7 × 2 mm.
Stipules oblonglanceolate, subacute, to 25 × 20 mm, caducous. Leaves thickly coriaceous,
drying warm yellowish brown; blade broadly ovate to elliptic, 14–18 × 7–10 cm, base
obtuse to cordate, apex with short, broad acumen; midrib evident, flat to somewhat sunken
above, prominent below; lateral veins 11–13 pairs, stoutly prominent below; intercostal
venation slender, densely scalariform, unraised; petiole 2.3–3.5 cm long. Inflorescences
terminal or axillary; rachis terete, to 15 cm long, singly branched; bracteoles lanceolate,
acute, to 16 × 5 mm, puberulent. Flowers: buds spindleshaped, to 8 × 3 mm; stamens 15,
irregularly whorled, filaments flat at base, shouldered medially, anthers ellipsoid,
connectival appendage c. 3x the length of anther, acicular, erect; ovary small, ovoid, without
stylopodium, finely sericeous, style 3x ovary, slender, stigma obscure. Fruits: pedicel short,
stout; calyx lobes unequal, 3 longer lobes to 7 × 1.5 cm, tapering to c. 5 mm broad above
the saccate base, 2 shorter ones linearlobed, to 2.5 × 0.3 cm, similar at base. Nuts ovoid, to
1 × 0.7 cm, acute.
Distribution. Endemic in Borneo. Known in Sarawak from Belaga, Dalat, Kapit, Lubok
Antu, and Marudi districts (e.g., S 25895, S 40709, S 60087, S 62412, and S 69724). Also
occurring in E Kalimantan (e.g., the type); not yet recorded from Sabah.
Ecology. Locally frequent in swampy pole forest on organic soils on poorly drained
plateaux over acid and also basic volcanic rocks and sandstone, at 800–1200 m altitude.
Becoming vulnerable owing to logging.
22. Shorea chaiana P.S.Ashton

(P.P.K. Chai, former Senior Forest Botanist in the Sarawak Forest Department)
Gard. Bull. Sing. 31 (1978) 42, op. cit. (1982) 477; Newman et al. op. cit. (1996) 128. Type: Suib S
29722, Borneo, Sarawak, Kapit district, Sg. Sepanggil (holotype K; isotype SAR).
240
Emergent tree, to 45 m tall, to 80 cm diameter; buttresses to 1 m tall. Petiole, inflorescence,

parts of perianth exposed in bud, ovary and nut persistently more or less densely creambuff
puberulent; fruit calyx lobes, twig and leaf venation below sparsely more or less caducously
so; other parts glabrous. Twigs c. 1 mm diameter apically, terete, muchbranched. Leaf buds
minute. Stipules not seen. Leaves thinly coriaceous, drying greenish grey; blade elliptic
lanceolate, more or less distinctly falcate, 6–11 × 2–4 cm, base cuneate or obtuse, margin
narrowly subrevolute, apex with caudate acumen to 1.5 cm long; midrib prominent below,
evident but flat to shallowly sunken above; lateral veins 8–11 pairs, slender but distinctly
raised below, evident above, arched; intercostal venation reticulate, distinctly elevated
below; petiole 0.5–0.8 cm long, slender. Inflorescences terminal or axillary; rachis slender,
to 6.5 cm long, singly branched. Flowers: buds to 5 × 2 mm; stamens 15, connectival
appendage c. 2½x the length of anther; ovary ovoidconical, stylopodium columnar,
puberulent, style columnar, glabrous, as long as ovary. Fruits (young): calyx lobes
subequal, short, ovate.
Distribution. Endemic in Borneo. Known in C and NE Sarawak from the Rajang and
Kemena hinterlands in Kapit district (e.g., S 29626, S 29632 and S 57838). Also occurring
in Brunei (e.g., FMS 35478 and FMS 35479).
Ecology. Apparently not common, in mixed dipterocarp forest, at altitudes below 1000 m.
Endangered owing to forest conversion.
23. Shorea collaris Slooten

(Latin, collum = neck; the rimmed fruit calyx)
Reinwardtia 3 (1956) 329; Ashton op. cit. (1968) 82, op. cit. (1982) 477; Anderson op. cit. (1980) 121;
Newman et al. op. cit. (1996) 129. Type: bb. 20650, Borneo, Kalimantan, Long Pahangai, Mahakam
(holotype BO; isotype L).
Emergent tree to 50 m tall, to 1.5 m diameter; bole tall, straight; buttresses to 2 m tall, stout
almost straight. Bark pale yellowish brown, evenly vertically cracked and flaked leaving a
scrollmarked surface below; coxcombs of black dammar frequent. Petiole and leaf bud
persistently greyish buffsericeous; young twig caducously so; inflorescence, parts of
perianth exposed in bud, ovary, stylopodium, and nut greyish buffpuberulent or glabrous.
Twigs c. 2 mm diameter apically, terete. Leaf buds ovoid, subacute, to 2 × 2 mm. Stipules
unknown. Leaves somewhat chartaceous, drying dull greyish green below, greyish brown
above; blade oblonglanceolate, 13–23 × 4.3–8.5 cm, base obtuse, margin usually narrowly
revolute, apex with broad acumen to 1 cm long; midrib slender but prominent below,
evident and flat above; lateral veins c. 13 pairs, slender but elevated below; intercostal
venation slender, hardly raised, scalariform; petiole 1.9–3 cm long, slender. Inflorescences
terminal or axillary; rachis terete, to 9 cm long, singly branched, branchlets bearing to 8
flowers; bracteoles fugaceous. Flowers: buds to 5 × 3 mm; stamens 10, connectival
appendage c. 2½x the length of anther, sparsely sericeous towards apex; ovary and
stylopodium pyriform, style short. Fruits: pedicels to 3 mm long; calyx glabrescent, lobes
equal, ovate, acute, clasping the nut at base, more or less reflexed distally giving a collared
appearance, to 1 × 0.9 cm. Nuts broadly obovoidellipsoid, to 3 × 2 cm, apiculate.
Vernacular names. Sarawak—lun kunyit (preferred name), merakunyit (Iban).
241
Distribution. Endemic in Borneo. Known in Sabah from G. Ampuria, Beaufort district

(e.g., SAN 43580) and in Sarawak from Kapit, Samarahan, Simunjan, and Tatau districts
(e.g., S 19307, S 23246, S 29612, S 60142, and S 67579). Also occurring in SE Kalimantan
(e.g., bb. 20479, bb. 20658 and the type).
Ecology. Frequent in mixed dipterocarp forest on clay rich soils at low altitudes, often near
streams, including on the Iju rhyolite and Mersing basalt, at altitudes to 500 m. Endangered
owing to forest conversion.
24. Shorea confusa P.S.Ashton

(Latin, confusus = confused; long confused with S. virescens)
Gard. Bull. Sing. 31 (1978) 44, op. cit. (1982) 497; Coode et al. (eds.) op. cit. 76; Newman et al. op.
cit. (1996) 129. Synonym: Shorea virescens auct. non Parijs: Ashton op. cit. (1964) 167, op. cit.
(1968) 95, Meijer & Wood op. cit. 60, Burgess op. cit. 159. Type: Meijer SAN 19395, Borneo, Sabah,
Tawau, Mile 24, Cocoa Estate (holotype K; isotypes KEP, SAN).
Large emergent tree, to 50 m tall, to 1.7 m diameter, with large hemispherical crown; bole
straight; buttresses to 3 m tall, stout. Bark pale fawnbrown, closely shallowly cracking and
thinly flaking; inner bark laminated pale orangebrown and cream. Leaf bud, parts of
perianth exposed in bud, ovary, inflorescence, stipule outside, petiole, and very young twig
caducous puberulent. Twigs 2–3.5 mm diameter apically, frequently rugulose. Leaf buds
conical, acute, 3–4 × 2 mm. Stipules linear to deltoid, subacute, c. 8 × 3 mm. Leaves thinly
coriaceous, drying yellowish brown with the venation paler than the blade below; blade
elliptic to slightly obovate, 6–12 × 3.5–5 cm, base obtuse, apex with broad acumen to 1 cm
long; midrib prominent below, obscurely furrowed above; lateral veins (10–)13–18 pairs,
arched, slender but raised below; intercostal venation evident, scalariform; petiole 1–1.5
cm long, c. 1 mm diameter, slender. Inflorescences terminal or axillary; rachis terete, lax, to
22 cm long, singly or doubly branched, branches bearing to 6 flowers; bracteoles not
observed. Flowers: buds to 9 × 5 mm; stamens 15, anther oblong, connectival appendage c.
3x the length of anther; ovary ovoid, without stylopodium small, puberulent, style c. 3x the
length of ovary, sericeous in basal twothirds. Fruits: pedicels stout, base of fruit tapering;
3 longer calyx lobes to 12 × 1.5 cm, hardly tapering above the saccate base, 2 shorter ones
unequal, linearlobed, to 6 × 0.5 cm, with similar base. Nuts ovoid, to 2 × 1.5 cm, with c. 6
mm slender style remnant.
Vernacular name. Sabah—melapi sulang saling (preferred name).
Distribution. Endemic in Borneo. In Sabah known from Kinabatangan, Kota Belud, Kota
Kinabalu, Kota Marudi, Lahad Datu, Sandakan, Tawau, and Tenom districts (e.g., SAN
15027, SAN 16349, SAN 16592, SAN 40334, and SAN 97247) and in Sarawak from Kapit,
Lawas, Marudi, and Miri districts (e.g., S 1533, S 17771, S 23023, and S 41224). Also
occurring in Brunei (e.g., BRUN 2004 and BRUN 5281) and E Kalimantan.
Ecology. Frequent in mixed dipterocarp forest on clayrich soils, on undulating land and
hills, at altitudes to 600 m. Recorded in Lambir NP; elsewhere vulnerable.
242
Notes. Shorea virescens, with which this species was formerly confused, differs in its
velutinate young parts, compressed twigs and larger number of lateral veins. Shorea confusa
differs from S. agami notably in its more slender and paler lateral veins below, and the
tapered base of its fruit. One collection with young fruit (SAN 33016 from Kelumpang,
Tawau district) may represent S. assamica Dyer which occurs in E Kalimantan. Flowering
material is required for confirmation.
25. Shorea cordata P.S.Ashton

(Latin, cordatus = heartshaped; the leaf base)
Newman et al. op. cit. (1996) 130. Type: Smythies SA 678, Borneo, Sarawak, Kuching district, near
Semengoh FR (holotype K).
Large emergent tree, to 50 m tall, to 1 m diameter, with tall straight bole; buttresses to 3 m
tall, stout. Bark pale chocolatebrown, irregularly cracked; inner bark pale brown, obscurely
laminated. Young twig and petiole densely caducous greyish brown puberulent; stipule and
leaf venation sparsely so; leaf bud and parts of perianth exposed in bud persistently so.
Twigs at first somewhat compressed c. 3 × 2 mm apically; stipule scars horizontal, pale,
prominent. Leaf buds ellipsoid, compressed, acute, to 7 × 4 mm. Stipules elliptic, obtuse, to
12 × 6 mm. Leaves coriaceous, drying chocolatebrown below; blade oblong to obovate, 8–
15 × 5.5–10.5 cm, base cordate to obtuse, apex obtuse, retuse or abruptly shortly acuminate;
midrib prominent below, furrowed above; lateral veins 15–18 pairs, prominent below;
intercostal venation slender, densely scalariform; petiole 1.2–2.5 cm long, frequently
becoming rugulose. Inflorescences terminal or axillary; rachis lax, compressed or ribbed, to
12 cm long, singly branched, branchlets bearing to 5 flowers; bracteoles elliptic, obtuse, to
10 × 5 mm, not at first caducous. Flowers: buds to 10 × 5 mm; petals pale yellow tinged
pink at base within; stamens 15, connectival appendage 3–4x the length of anther, scabrous;
ovary puberulent distally, style c. 2x as long as ovary. Fruits: calyx glabrous; 3 longer lobes
to 13 × 2.5 cm, tapering to 8 mm above the saccate base, 2 shorter ones unequal, to 6.5 × 0.8
cm, similar at base. Nuts narrowly ovoid, to 1.4 × 1 cm, with to 4 mm slender style
remnant.
Distribution. Endemic in Borneo. Known in Sarawak from Kuching, Lundu and Tatau
districts (e.g., S 10177, S 13183, S 19306, and S 68788). Also occurring in NW Kalimantan.
Ecology. Rare and probably endangered, in mixed dipterocarp forest, apparently on clay
rich soils especially on igneous rocks, at altitudes below 500 m.
26. Shorea coriacea Burck Plate 3F.

(Latin, coriaceus = leathery; the leaf blade)
Ann. Jard. Bot. Buitenz. 6 (1887) 214; Merrill op. cit. (1921) 404; Masamune op. cit. 493; Browne op.
cit. 146; Ashton op. cit. (1964) 182, op. cit. (1968) 106, op. cit. (1982) 509; Meijer & Wood op. cit.
102; Burgess op. cit. 154; Anderson op. cit. (1980) 125; PROSEA op. cit. 393; Kessler & Sidiyasa,
243
TBSAEK (1994) 102; Coode et al. (eds.) op. cit. 76; Newman et al. op. cit. (1996) 131. Type:
Beccari PB 2948, Borneo, Sarawak (holotype BO; isotypes A, K).
Large emergent tree, to 50 m tall, to 1.4 m diameter, with straight bole; buttresses to 1.5 m
tall, stout. Bark dark chocolatebrown, narrowly fissured and flaky; inner bark pinkish
brown; heartwood deep crimson. Young parts densely pale grey sericeous, caducous except
on stipule outside, leaf bud, parts of perianth exposed in bud and inflorescence; ovary and
nut puberulent except at base. Twigs terete or slightly compressed, 2–3 mm diameter
apically; stipule scars cuneate, pale brown, horizontal or slightly descending. Leaf buds
ovoid, to 6 × 3 mm. Stipules narrowly oblong, acute, to 20 × 5 mm. Leaves coriaceous,
lustrous, drying dark reddish brown below; blade ovate, 10–15 × 5–8 cm, base obtuse,
obscurely subpeltate, margin narrowly revolute, apex with acumen to 1 cm long; midrib
prominent below, obscurely sunken above; lateral veins 16–20 pairs, slender, hardly raised
below, curved within margin; intercostal venation densely scalariform, unraised; petiole
geniculate, drying black, 2.5–4 cm long. Inflorescences terminal or axillary; rachis terete, to
14 cm long, singly or doubly (if terminal) branched, branchlets bearing to 10 flowers;
bracteoles ovate, subacute, to 4 × 2.5 mm. Flowers: buds to 6 × 1.2 mm; petals pink;
stamens 15, connectival appendage 2–3x the length of anther; ovary ovoid, densely short
pubescent except at the base, stylopodium indistinct, style as long as ovary. Fruits: calyx
lobes unequal, 2 longer lobes to 9 × 1.5 cm, tapering to 4–5 mm at the saccate base, 2
shorter ones subequal, linearlobed, to 3.5 cm long, similar at base. Nuts ovoid, to 1.8 × 0.9
cm, with c. 3 mm slender style remnant.
Vernacular names. Sabah—seraya tangkai panjang (preferred name). Sarawak—meranti

tangkai panjang (preferred name).
Distribution. Endemic in Borneo. Known in Sabah from Sipitang district (e.g., SAN 16682
and SAN 69878) and in Sarawak from Bau, Belaga, Kuching, Lawas, Lundu, and Tatau
(e.g., BRUN 3335) and W, C and E Kalimantan (e.g., Ambriansyah and Arifin AA 41, bb.
29692 and Laman et al. TL 1348).
Ecology. Locally abundant in kerangas forest especially on deep podsols, in the lowlands
and on sandstone and acid volcanic plateaux in the lower montane zone, at 100–1200 m
altitude; on ultrabasic substrate in eastern Sabah; occasional in mixed dipterocarp forest in
the kerangas ecotone. Occurring in Mulu NP; elsewhere becoming endangered.
27. Shorea crassa P.S.Ashton

(Latin, crassus = thick; the leaf blade)
op. cit. (1980) 119; Coode et al. (eds.) op. cit. 76; Newman et al. op. cit. (1998) 172. Type: Ismail S
11701, Borneo, Sarawak, Semengoh FR (holotype K; isotype KEP).
Emergent tree, to 45 m tall, to 1.1 m diameter, with straight bole; buttresses to 1.5 m tall,
thin, sharp. Bark tawny to purplish brown, vertically cracked and irregular flaky. Twig,
inflorescence, leaf bud, parts of perianth exposed in bud, ovary, nut, stipule outside, and
petiole densely greyishpuberulent; venation below and fruit calyx sparsely so. Twigs at
244
first somewhat compressed, c. 5 × 2.5 mm apically; stipule scars c. 2 mm long, pale,

ascending. Leaf buds ovoid, compressed, subacute, to 6 × 6 mm. Stipules ovate, acute,
concave, to 8 × 4 mm, fugaceous. Leaves thickly coriaceous, pale cream to golden lepidote
below; blade elliptic to ovate, 10–18 × 5–10 cm, base equal or subequal, cuneate or
narrowly obtuse, occasionally subcordate, apex with tapering acumen to 1 cm long; midrib
prominent below, applanate above to sunken towards apex, evident; lateral veins 7–11
pairs, prominent below, wellspaced, ascending; intercostal venation densely scalariform,
sinuate, unraised; petiole 3.5–5 cm long, stout, somewhat geniculate. Inflorescences
terminal or axillary; rachis terete or somewhat compressed, ribbed on drying, stout, to 13 cm
long, singly branched, branchlets bearing to 12 flowers; bracteoles suborbicular, to 3 mm
across, caducous. Flowers: buds to 15 × 3.5 mm; petals cream, pink at base; stamens 38–46,
filaments hispid, connectival appendage prominent but shorter than the anther, setose;
ovary ovoid, stylopodium cylindrical longer than ovary, style glabrous, short. Fruits: calyx
lobes unequal, 3 longer lobes to 9 × 2.5 cm, tapering to 8 mm above the saccate base, 2
shorter ones narrowly oblong, to 7 × 0.7 cm, similar at base. Nuts ellipsoid, to 2.5 × 2 cm,
tapering into stout to 8 mm long style remnant.
Vernacular name. Sarawak—selangan batu daun tebal (preferred name).
Distribution. Endemic in Borneo. Known in Sarawak from Bintulu, Kapit, Kuching,

Lundu, and Miri districts (e.g., S 6210, S 9468, S 32051, S 46484, and S 59866). Also
occurring in Brunei (e.g., BRUN 570 and BRUN 642) and C and W Kalimantan (e.g., bb.
17036 and bb. 26409).
Ecology. Locally common in mixed dipterocarp forest on deep yellow sandy soils in the
lowlands, and occasionally at altitudes to 1000 m on sandstone. Occurring in Mulu NP;
28. Shorea curtisii Dyer ex Brandis

(C. Curtis, 1853–1928, first Curator of Waterfall gardens, Penang, Peninsular Malaysia)
(sect. Mutica, subsect. Mutica, red meranti)
J. Linn. Soc. Bot. 31 (1895) 101; Symington op. cit. (1943) 67; Browne op. cit. 147; Ashton op. cit.
(1964) 185, op. cit. (1968) 106, op. cit. (1982) 541; Meijer & Wood op. cit. 103; Burgess op. cit. 166;
Anderson op. cit. (1980) 125; PROSEA op. cit. 393; Coode et al. (eds.) op. cit. 76; Newman op. cit.
(1996) 541. Lectotype (designated here): Curtis 1394, Peninsular Malaysia, Penang Hill
(hololectotype K).
Notes. Two subspecies, subsp. curtisii and subsp. grandis P.S.Ashton, are recognised. Of
the two, only subsp. curtisii occurs in Sabah and Sarawak.
subsp. curtisii
Vast emergent tree, to 70 m tall, to 2 m diameter, with huge hemispherical cauliflower

shaped crown, seagreen from a distance; buttresses to 1.8 m tall, stout. Bark becoming
pinkish brown, deeply vsection fissured and eventually crumbly flaky; inner bark thick,
pinkish brown; heartwood pale crimson. Young twig, inflorescence, leaf bud, parts of
perianth exposed in bud, stipule outside, ovary, nut, petiole, and midrib and veins below
245
densely evenly more or less persistently pinkish brown to grey puberulent. Twigs c. 1.5 mm
diameter apically, slender, terete, much branched. Leaf buds ovate to slightly falcate,
subacute, 4–9 × 2.5–3.5 mm. Stipules oblong, obtuse, to 10 × 4 mm. Leaves coriaceous,
pale pink to grey lepidote and dull below; blade ovatelanceolate, 6–9 × 2.5–3.5 cm, base
broadly cuneate, apex with narrow acumen to 0.8 cm long; midrib slender, terete,
prominent below, obscure, slightly sunken above; lateral veins 9–11 pairs, slender, hardly
raised below, arched; intercostal venation obscure, slender, densely scalariform; petiole 1–
1.3 cm long, slender. Inflorescences terminal or axillary; rachis terete, to 6 cm long, singly
or doubly branched, branchlets bearing to 12 flowers; bracteoles elliptic, subacute, to 2.5 ×
2 mm, fugaceous. Flowers: buds to 5 × 3 mm; petals deep crimson; stamens 15, connectival
appendage short or rudimentary; ovary and stylopodium ovoid, style short. Fruits: pedicels
c. 1 mm long; calyx lobes unequal, 3 longer lobes to 7 × 1 cm, tapering to c. 2.5 mm above
the saccate base, 2 shorter ones unequal, linearlobed, to 4 × 0.3 cm, similar at base. Nuts
ovoid, to 1.2 × 0.9 cm, tapering to 1 mm apiculus.
Vernacular name. Sabah and Sarawak—seraya (preferred name).
Distribution. SE Peninsular Thailand, Sumatra (Singkep and Lingga), Peninsular Malaysia,

and Borneo. In Borneo, known in Sabah from Sipitang district (e.g., SAN 15185) and in
Sarawak from Bintulu, Miri and Mukah districts (e.g., S 15146, S 23606, S 23653, and S
46414). Also occurring in Brunei (e.g., BRUN 246, BRUN 2538 and BRUN 3279).
Ecology. Locally common in mixed dipterocarp forest on deep yellow sands along the
Neogene coastal hills. Recorded in Brunei and Ulu Limbang from shallow organic soils
along shale ridges to c. 1200 m altitude in upper dipterocarp forest. Locally frequent in
Lambir and Mulu NPs. Vulnerable in Sarawak outside the parks; critically endangered in
Sabah.
29. Shorea cuspidata P.S.Ashton

(Latin, cuspis = a cusp; the slightly swollen acumen apex)
Newman et al. op. cit. (1996) 134. Type: Galau S 15258, Borneo, Sarawak, Kuching, Semengoh FR
Low emergent tree, to 45 m tall, to 1 m diameter, with straight bole; buttresses to 3 m tall,
rather thin. Bark surface at first vsection zigzag fissured, later flaking in small oblong
pieces leaving a scrollmarked surface. Leaf bud, parts of perianth exposed in bud, ovary,
nut, stipule, bracteole, and inflorescence shortly densely buffpuberulent; young twig and
fruit calyx caducously so; parts otherwise glabrous. Twigs c. 1 mm diameter apically,
slender, becoming minutely striated. Leaf buds ovoid, c. 1 mm long, small. Stipules not
seen. Leaves thinly coriaceous, drying greenish to yellowish brown; blade broadly ovate, 5–
9 × 2–6 cm, base broadly cuneate, apex caudatecuspidate, acumen to 1.5 cm long; midrib
slender, evident and flat or elevated above, prominent below; lateral veins 5–7 pairs,
slender, hardly raised below, arched; intercostal venation obscure, reticulate; petiole 0.7–
1.1 cm long, slender. Inflorescences terminal or axillary; rachis slender, terete, to 9 cm
long, singly branched. Flowers: buds to 3 × 2 mm; petals pale limeyellow; stamens 15,
connectival appendage c. 1½x the length of anther, sericeous distally; ovary and
246
stylopodium pyriform, style glabrous, short. Fruits: pedicels c. 1 mm long; calyx lobes
unequal, 3 longer lobes to 5 × 1.5 cm, tapering to 5 mm above the narrowly saccate
tuberculate base, 2 shorter ones to 4 × 1 cm, otherwise similar. Nuts obovoid, mucronate,
to 2.5 × 1.5 cm.
Vernacular name. Sarawak—lun runching padi (preferred name).
Distribution. Endemic in Borneo; known in Sarawak from Bau, Kuching, Lundu, Miri, and
Simunjan districts (e.g., S 6513, S 9607, S 9843, S 15211, and S 49991). Also occurring in
W Kalimantan (e.g., Suzuki K 9542).
Ecology. Frequent to locally abundant in mixed dipterocarp forest on yellow sandy soils
over the Plateau Sandstone, also on leached clayrich soils on other substrates, on
undulating ground and low hills, at altitudes to 500 m. Common in Bako NP and recorded
from Lambir NP; elsewhere endangered by land conversion.
30. Shorea dasyphylla Foxw.

(Greek, dasy = markedly hairy, phullon = leaf; the pubescent leaf)
Malay. For. Rec. 10 (1932) 224; Browne op. cit. 138; Ashton op. cit. (1968) 106, op. cit. (1982) 546;
Anderson op. cit. (1980) 125; PROSEA op. cit. 394; Newman et al. op. cit. (1996) 135. Type:
Symington KEP 24462, Peninsular Malaysia, Selangor, Sg. Buloh FR (holotype KEP).
Mediumsized emergent tree, to 45 m tall, to 1.1 m diameter, with dense hemispherical

crown; bole straight; buttresses to 1.5 m tall, stout. Bark dark greyish brown, deeply v
section fissured; inner bark and heartwood rich crimson. Twig, leaf bud, inflorescence, parts
of perianth exposed in bud, ovary, nut, stipule, bracteole, petiole, leaf below, and midrib
above densely persistently golden brown scabridpubescent; blade above puberulent. Twigs
c. 2 mm diameter apically, terete. Leaf buds ovoid, obtuse, to 3 × 2 mm. Stipules broadly
ovate, obtuse, to 6 × 4 mm, caducous. Leaves coriaceous, drying pale yellowish brown
below, pale mauvebrown above; blade ovate to elliptic, 7–14 × 3–6 cm, obtuse or broadly
cuneate, margin somewhat revolute, base obtuse or broadly cuneate, apex with slender
acumen to 0.6 cm long; midrib prominent below, terete, striated, furrowed above; lateral
veins 11–15 pairs, prominent below; intercostal venation scalariform, elevated below;
petiole 1.2–1.5 cm long. Inflorescences terminal or axillary; rachis terete or ribbed, to 8 cm
long, singly branched, branchlets bearing to 4 flowers; bracteoles elliptic, obtuse, to 3 × 2
mm, caducous. Flowers: buds to 4 × 3 mm; petals creamyellow; stamens 15, connectival
appendage short, slender, becoming reflexed; ovary and stylopodium ovoid to conical, style
shorter than ovary, glabrous. Fruits: pedicel to 2 mm long; calyx lobes unequal, 3 longer
lobes to 9 × 1.3 cm, tapering to c. 4 mm above the saccate base, 2 shorter ones linear
lobed, to 4 × 0.4 cm, similar at base. Nuts ovoid, shortly apiculate, to 1.8 × 0.9 cm.
Vernacular name. Sarawak—meranti batu (preferred name).
247
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo known in Sabah from
Tambunan district (e.g., KEP 80416) and in Sarawak from Bau, Kuching, Limbang, and
SE Kalimantan (e.g., bb. 34435).
Ecology. Frequent, sometimes common, in mixed dipterocarp forest on yellow sandy clay
soils, at altitudes below 400 m. Common in Bako NP; elsewhere vulnerable owing to land
conversion.
31. Shorea dealbata Foxw.

(Latin, dealbatus = whitewashed; the leaf undersurface)
Malay. For. Rec. 10 (1932) 192; Symington op. cit. (1943) 35; Ashton op. cit. (1968) 93, op. cit.
(1982) 485; Anderson op. cit. (1980) 123; PROSEA op. cit. 410; Newman et al. op. cit. (1996) 136.
Type: Mat Sohor KEP 4188, Peninsular Malaysia, Pahang, Kuantan district, Jambu, Lepar (holotype
KEP).
Low emergent or main canopy tree, to 30 m tall, to 60 m diameter, with straight bole and
low stout buttresses. Bark at first mottled brown and grey, hoopmarked, becoming deeply
irregularly cracked, rotting off in flakes in between; inner bark finely pale rust and cream
laminated. Young twigs, leaf buds, parts of perianth exposed in bud, stipule and bracteole
outside, petiole, and nut densely darkfulvous pubescent; inflorescence similarly hirsute,
venation below fugaceously so; blade below pale pinkbrown lepidote. Twigs compressed,
c. 4 × 2 mm apically. Leaf buds ovoid, conical, subacute, to 6 × 4 mm. Stipules elliptic
oblong, subacute, to 20 × 5 mm. Leaves thickly coriaceous, turned up along the midrib,
drying dull olivebrown above, pale pinkish grey below with petiole; blade ovate to elliptic,
8–16 × 4–7.5 cm, base obtuse or occasionally cuneate, apex with narrow acumen to 1 cm
long; midrib prominent, subacute, below, obscurely sunken above; lateral veins (11–)20–24
pairs, obscure and slightly depressed above, slender and hardly elevated below, ascending;
petiole stout, becoming rugose, 1.5–2.5 cm long. Inflorescences axillary or terminal, rachis
terete or angular, unbranched or singly branched; bracteole unknown. Flowers: buds to 12 ×
5 mm; petals white tinged with pink at base; stamens 17, anthers narrowly oblong,
connectival appendage stout at base, tapering and slender apically, c. 3x the length of
anther; ovary ovoid, sparsely sericeous, without stylopodium, style filiform, trifurcate
apically, sericeous in the basal half. Fruits: pedicels to 5 mm long, slender; calyx lobes
ones loratelobed, to 6.5 × 0.5 cm, similar but narrower at base. Nuts narrowly ovoid, to
1.8 × 0.8 cm, with to 3 mm slender style remnant.
Vernacular name. Sarawak—meranti bumbung (preferred name).
Distribution. Sumatra (Lingga), Peninsular Malaysia and Borneo. In Borneo known only in
Sarawak from Kuching district (e.g., S 9409, S 10139, S 12476, and S 32730).
Ecology. Very local, but there often common, in kerangas forest on white sand terraces and
on sandstone plateau, at altitudes to 600 m. Occurring in Bako NP; elsewhere vulnerable.
248
32. Shorea dispar P.S.Ashton

(Latin, dispar = unlike; the flowers, compared with those of S. parvifolia with similar
leaves)
Gard. Bull. Sing. 31 (1978) 45, op. cit. (1982) 502; Newman et al. op. cit. (1996) 157. Type: Othman
S 29208, Borneo, Sarawak, Kapit, Ulu Baleh above Nanga Mengiong (holotype K; isotypes KEP, L).
Large buttressed emergent tree. Twig, petiole, bud, inflorescence, and parts of perianth
exposed in bud densely persistently paletawny puberulent; leaf venation below sparsely so;
parts otherwise glabrous. Twigs c. 2 mm diameter apically, terete, much branched. Leaf
buds ellipsoid, obtuse, to 3 × 2 mm. Stipules unknown. Leaves coriaceous, somewhat
cream lepidote below; blade elliptic, 4–7 × 2–3.5 cm; midrib prominent below, obscurely
sunken above as also the veins; lateral veins 9–11 pairs, ascending; intercostal venation
scalariform, obscure, unraised; petiole 1.2–1.6 cm long. Inflorescences terminal or axillary;
rachis rigid, ascending, singly branched, to 8 cm long. Flowers: buds ovoid, plump, to 6 × 4
mm; sepals subequal, broadly ovate, acute; petals lanceolate, contorted; stamens c. 25,
filaments lorate, somewhat tapering, anthers oblong, connectival appendage c. ¾x the
length of anther, stout, becoming somewhat reflexed; ovary ovoid, without stylopodium,
small, glabrous, style filiform, c. 2x the length of ovary. Fruits: pedicels to 1 mm; calyx
lobes unequal, 3 longer lobes to 9 × 2 cm, more or less tapering to a more or less
subauriculate base, 2 shorter ones loratelobed, to 2.5 × 0.5 cm, not auriculate. Nuts ovoid,
to 3 × 1.5 cm, apiculate, glabrous.
Distribution. Endemic in Sarawak; known only from the Ulu Baleh, Kapit district (e.g., S
29489 and S 29639).
Ecology. In mixed dipterocarp forest on shale ridges, at 500–600 m altitudes. Rare and
critically endangered.
33. Shorea domatiosa P.S.Ashton

(Latin, domus = a dwelling; the leaf domatia, dwellings for acarines)
Newman et al. op. cit. (1998) 173. Type: G.H.S. Wood SAN 16601, Borneo, Sabah, Sipitang (holotype
KEP).
Large emergent tree, to 50 m tall, to 1.2 m diameter, with diffuse pale hemispherical crown;
bole straight; buttresses to 2 m tall, thin, prominent. Bark becoming vertically cracked and
thinly oblong flaky, pale fawnbrown. Inflorescence, parts of perianth exposed in bud,
ovary except at base and apex, nut, and stipules cream puberulent; domatia fimbriate; parts
otherwise glabrous. Twigs c. 1.5 mm diameter apically, slender, muchbranched. Leaf buds
ovoid, acute, c. 4 × 2 mm. Stipules ovate, narrowly acute, to 6 × 2.5 mm, caducous. Leaves
chartaceous, cream lepidote below (mature tree), drying dark greyish brown; blade broadly
ovate to obovate, 6.5–10 × 3–7 cm, base obtuse to subcordate, subequal, apex with acumen
to 0.8 cm long; midrib raised but not prominent below, more or less flat, evident, above;
249
Fig. 23. Shorea elliptica. A, fruiting leafy twig; B, detail of indumentum on lower leaf
surface; C, flower bud; D, open flower; E, longitudinal section of open flower with petals
removed; F, adaxial view of outer sepal; G, adaxial view of inner sepal; H, abaxial view of
petal; I, adaxial view of petal with stamens; J, adaxial view of stamens; K, abaxial view of
stamens; L, fruit; M, longitudinal section of fruit. (A–B and L from S 15794, C–K from S
37698, M from S 29468.)
250
lateral veins 8–12 pairs, slender, arched, elevated below, with small porelike domatia;
intercostal venation slender, densely scalariform, sinuate; petiole 1.5–2.5 cm long, slender,
geniculate and swollen in the distal half. Inflorescences terminal or axillary; rachis terete,
to 7 cm long, singly branched, branches bearing to 5 flowers. Flowers: buds to 13 × 4 mm;
stamens 25–30, anthers sparsely setose towards apices, connectival appendage c. ¼x the
length of anther, stout, setose; ovary small, ovoid, stylopodium 1½x the length of ovary,
slender, style glabrous, short. Fruits: calyx glabrous, lobes unequal, 3 longer lobes to 13 ×
3 cm, tapering to 8 mm above the saccate base, 2 shorter ones to 7 × 1.2 cm, otherwise
similar. Nuts ovoid, to 3.5 × 2.5 cm, with to 1 cm prominent style remnant.
Vernacular names. Sabah—selangan batu matamata (preferred name). Sarawak—

selangan batu lubang hidung (preferred name).
Distribution. Endemic in Borneo. Known in Sabah from Beaufort, Sipitang and Tawau
districts (e.g., SAN 15164, SAN 21492 and SAN 37110) and in Sarawak from Bintulu, Lawas
and Miri districts (e.g., S 1785, S 22069 and S 24984). Also occurring in Brunei (e.g., BRUN
3395 and BRUN 5666), and NE Kalimantan.
Ecology. Scattered, usually as single mature trees, but widespread, in mixed dipterocarp
forest on shale ridges, at altitudes to 700 m. Vulnerable.
34. Shorea elliptica Burck Fig. 23.

(Latin, ellipticus; the elliptic leaf blade)
cit. 138; Ashton op. cit. (1968) 151, op. cit. (1982) 500; Anderson op. cit. (1980) 125; PROSEA op.
cit. 428; Newman et al. op. cit. (1996) 137. Type: Beccari PB 2574, Borneo, Sarawak, Matang
(holotype BO).
Mediumsized emergent tree, to 40 m tall, to 1.2 m diameter, with hemispherical dense

crown pale from below; bole straight; buttresses to 1.5 m tall, stout. Bark becoming dark
mauvebrown dappled, pale grey, deeply vsection fissured; inner bark orangebrown;
heartwood pink. Twig, leaf bud, parts of perianth exposed in bud, stipule and bracteole
outside, inflorescence, nut, and veins and midrib below densely ochreous to cream scabrid
puberulent; lateral veins above and stipule within sparsely evenly so; blade densely cream
lepidote below. Twigs c. 2 mm diameter apically, terete to somewhat compressed, much
branched; stipule scars short. Leaf buds ovoid, obtuse, compressed, to 8 × 6 mm. Stipules
narrowly ovate, acute, to 10 × 6 mm, caducous. Leaves thickly coriaceous, drying dull
greyish brown above, cream below; blade ellipticoblong, 7–10 × 4–9 cm, base broadly
cuneate to subcordate, apex obtuse to shortly broadly acuminate; midrib evident but
shallowly furrowed above, prominent, terete below; lateral veins 11–16 pairs, prominent
below; intercostal venation slender, densely scalariform, more or less obscure; petiole 1.5–
3.3 cm long. Inflorescences terminal or axillary; rachis more or less compressed, to 12 cm
long, singly or doubly branched, branchlets bearing to 3 flowers; bracteoles elliptic, obtuse,
to 2 × 1 mm, caducous. Flowers: buds to 4 × 3 mm; petals pale yellow; stamens 19–20,
anthers oblong, almost as long as filaments, connectival appendage stout, somewhat shorter
than anther; ovary without stylopodium, glabrous, style c. 2x as long as ovary. Fruits:
pedicels to 3 mm, expanding into receptacle; calyx lobes unequal, 3 longer lobes narrowly
251
spatulate to sublorate, to 8 × 1.5 cm, tapering to 4 mm above the saccate base, 2 shorter
ones to 6 × 0.8 cm, otherwise similar. Nuts ovoid, acute, to 1.5 × 1.2 cm.
Vernacular name. Sarawak—meranti lang (preferred name).
Distribution. Endemic in Borneo. Known in Sarawak from Kuching and Lundu districts
(e.g., S 10059, S 27769, S 29468, S 37765, and S 49960) and in Kalimantan from Lower
Kapuas valley.
Ecology. Locally common, in mixed dipterocarp forest on leached sandy clay soils, at
altitudes to 500 m. Occurring in Bako NP; elsewhere endangered.
35. Shorea exelliptica Meijer Plate 4A.

(Latin, ex = excluded from, elliptica = S. elliptica; species formerly confused with S.
elliptica)
Act. Bot. Neerl. 12 (1963) 323; Meijer & Wood op. cit.167; Ashton op. cit. (1964) 132, op. cit. (1968)
Coode et al. (eds.) op. cit. 76; Newman et al. op. cit. (1998)174. Lectotype (designated here):
Rahman S 1641, Borneo, Brunei, Belait (hololectotype K; isolectotypes KEP, L). Synonyms: Shorea
?elliptica auct. non Burck: Symington op. cit. (1943) 13; S. sp. Browne op. cit. 170.
Emergent tree, to 60 m tall, to 1 m diameter, with dense hemispherical crown pale from
below; bole straight; buttresses to 1.5 m tall, spreading, rather thin. Bark becoming closely
cracked, then thinly oblong flaky, the flakes becoming persistent in big trees. Twig,
inflorescence, leaf bud, parts of perianth exposed in bud, stipule, bracteole, petiole, midrib
on both surfaces, and veins below densely persistently purplish brown to golden brown
scabridpubescent, sometimes glabrescent on midrib above; ovary and nut evenly buff
pubescent. Twigs ridged, at first somewhat compressed or terete, c. 2 mm diameter apically.
Leaf buds ovoid, more or less compressed, to 3.5 × 3 mm. Stipules broadly ovate, acute, to
8 × 4 mm, not at first caducous. Leaves coriaceous, golden to silver lepidote below (mature
trees), drying purplish brown with cream undersurface, the veins and midrib dark purplish;
blade oblong to broadly ovate, 9–15 × 3.5–7 cm, base broadly cuneate, apex with tapering
acumen to 1 cm long; midrib prominent below, evident but furrowed above; lateral veins
12–18 pairs, prominent below; intercostal venation scalariform, slender, sinuate; petiole
1.2–1.7 cm long, rugose. Inflorescences terminal or axillary; rachis ribbed and slightly
compressed, to 12 cm long, singly branched, branchlets bearing to 8 flowers; bracteoles
elliptic, acute, to 4 mm long, caducous. Flowers: buds to 10 × 3 mm; petals cream, pink at
base; stamens 30–40, filaments slightly hispid, connectival appendage as long as anther,
setose; ovary and stylopodium shortly ovoid, densely tomentose except at the base, style as
long as both, slender, glabrous. Fruits: calyx glabrous, lobes unequal, 3 longer lobes to 8 ×
2.4 cm, tapering to 4 mm above the saccate base, 2 shorter ones to 4.5 × 1.7 cm, otherwise
similar. Nuts ovoid, to 1.5 × 1 cm, with to 3 mm tapering style remnant.
Vernacular name. Sabah and Sarawak—selangan batu tembaga (preferred name).
252
Distribution. Peninsular Malaysia and Borneo. In Borneo known in Sabah from

Kinabatangan, Lahad Datu, Sandakan, Sipitang, and Tawau districts (e.g., SAN 15165, SAN
16986, SAN 19559, and SAN A 4242) and in Sarawak from Belaga, Kapit, Kuching, Lawas,
Limbang, Marudi, and Miri districts (e.g., S 1432, S 32326, S 38750, S 41230, and S 68994).
Also occurring in Brunei (e.g., BRUN 880 and BRUN 5661) and Kalimantan (e.g.,
Ambriansyah & Arifin Berau 1075 and bb. 29311).
Ecology. Frequent in mixed dipterocarp forest on yellow clay and sandy clay soils on the
sedimentary rocks of the inland ridges, at altitudes to 600 m; and less so on undulating land
in the lowlands. Occurring in Lambir and Mulu NPs; elsewhere vulnerable owing to
logging.
36. Shorea faguetiana F.Heim

(Faguet, 19th century French botanical assistant at Paris)
Bull. Mens. Soc. Linn. Paris 2 (1891) 975; Merrill op. cit. (1921) 414; Symington, Gard. Bull. S. S. 7
(1933) 148, op. cit. (1939) 381, op. cit. (1943) 50; Masamune op. cit. 493; Browne op. cit. 162;
op. cit. 217; Anderson op. cit. (1980) 121; PROSEA op. cit. 418; Coode et al. (eds.) op. cit. 76;
Newman et al. op. cit. (1996) 138. Lectotype (designated here): Beccari PB 2491, Borneo, Sarawak,
Matang (hololectotype P).
Large emergent tree, to 50 m tall, to 1.5 m diameter, with straight bole and dense
hemispherical crown; buttresses to 1.5 m tall, stout. Bark fawnbrown, vertically cracked
and thinly oblong flaky; dammar coxcombs present, dark greyish brown. Leaf bud, parts of
perianth exposed in bud, young twig, inflorescence, stipule, and bracteole pale greyish
brown puberulent; young leaves fugaceous puberulent below; ovary, stylopodium, and nut
densely cream buff pubescent. Twigs c. 1 mm diameter apically, slender, rugulose,
lenticellate. Leaf buds ovoid to conical, c. 2 × 1.5 mm, small. Stipules hastate, to 4 × 1.5
mm. Leaves coriaceous, drying greyish green; blade elliptic to oblonglanceolate or ovate,
7–12 × 3–5 cm, base cuneate to obtuse, equal or subequal, apex with tapering acumen to 1
cm long; midrib raised and terete below, evident and shallowly furrowed to flat above;
lateral veins 9–12 pairs; intercostal venation rather dense, subscalariform, slightly elevated;
petiole 1–1.5 cm long, fairly stout. Inflorescences terminal or axillary, rachis lax, terete, to
20 cm long, doubly or trebly branched, branchlets bearing to 8 flowers; bracteoles small,
fugaceous. Flowers: buds small, to 3 mm long; petals cream; stamens 15, connectival
appendage as long as anther, sericeous towards apex; ovary subglobose, stylopodium as
long as ovary, style short, glabrous. Fruits: calyx sparsely puberulent, lobes unequal, 3
longer lobes to 6 × 1.2 cm, tapering to 2.5 mm above the tuberculate saccate base, 2
shorter ones to 4.5 cm long, narrower, otherwise similar. Nuts narrowly ellipsoid to
obovoid, acute, to 1.5 × 0.5 cm.
Vernacular names. Sabah—seraya kuning siput (preferred name). Sarawak—lun siput

(preferred name).
Distribution. SE Peninsular Thailand, Peninsular Malaysia and Borneo. In Borneo, occur

throughout Sabah and recorded from Beaufort, Keningau, Kinabatangan, Sandakan,
Sipitang, Tawau, and Tenom districts (e.g., SAN 15143, SAN 22789, SAN 36401, SAN
61239, SAN 63063, and SAN 68597) and in Sarawak from Bau, Belaga, Bintulu, Kapit,
253
Kuching, Lawas, Limbang, Lundu, Simunjan, and Tatau districts (e.g., S 15221, S 22485, S
27122, S 29635, and S 49931). Also occurring in Brunei (e.g., BRUN 3019 and BRUN 3178)
and E Kalimantan (e.g., Arifin et al. Berau 710 and Arifin and Ambriansyah Berau 1040).
Ecology. Widespread and frequent, in mixed dipterocarp forest on welldrained clay and
sandy clay soils, at altitudes to 1000 m. Well represented in Kubah, Lambir and Mulu NPs;
probably not vulnerable.
37. Shorea faguetioides P.S.Ashton

(Greek, oides = resembling; similar to S. faguetiana)
Wood op. cit. 80; Anderson op. cit. (1980) 122; Coode et al. (eds.) op. cit. 77; Newman et al. op. cit.
(1996) 139. Type: Drahman S 1771, Borneo, Sarawak, Lawas, Bt. Pengalih (holotype K; isotype
KEP).
Large emergent tree, to 50 m tall, to 1.4 m diameter, with straight cylindrical bole and dense
hemispherical crown. Bark pale greyish tawny, shallowly cracked and evenly flaky,
appearing smooth overall; dammar in greyish coxcombs, black within. Inflorescence and
parts of perianth exposed in bud persistently greyish brown puberulent; ovary, stylopodium
and nut densely buff puberulent; young twig sometimes caducously so; parts otherwise
glabrous. Twigs c. 3 mm diameter apically, somewhat compressed at first, drying blackish,
rugulose. Leaf buds ovoid, obtuse, compressed, 3–4 × 2.5 mm. Stipule narrowly deltoid,
saccate, obtuse, to 18 × 6 mm, not at first caducous. Leaves brilliant magenta when
opening, chartaceous, drying tawnybrown lustrous below with darker midrib; blade
narrowly ovate, 12–18 × 5–7 cm, base subequal, broadly cuneate to subcordate, apex with
prominent acumen to 2 cm long; midrib evident, flat or shallowly furrowed above,
prominent below; lateral veins 10–15 pairs, slender but distinctly raised below, arched;
intercostal venation scalariform, slender, evident; petiole 2.2–3 cm long, c. 2 mm diameter,
slender, drying black, weakly geniculate. Inflorescences terminal or axillary; rachis terete,
lax, to 15 cm long, doubly or sometimes trebly branched, branchlets bearing to 7 flowers;
bracteoles small, fugaceous. Flowers: buds to 2.5 × 1.5 mm; petals cream; stamens 15,
connectival appendage c. 3x the length of anther, sericeous towards apex; ovary ovoid,
stylopodium shorter than ovary, cylindrical, style short, glabrous. Fruits: calyx puberulent
towards base, otherwise glabrous, lobes unequal, chartaceous, 3 longer lobes to 7 × 1.5 cm,
tapering to 4 mm above narrow saccate tuberculate base, 2 shorter ones to 5 × 0.7 cm,
otherwise similar. Nuts ellipsoid to obovoid, to 2 × 0.6 cm, with 3 mm slender style
remnant.
Vernacular names. Sabah—seraya kuning (preferred name). Sarawak—barek (Iban).
Distribution. Endemic in Borneo. In Sabah known from Beaufort, Keningau and Kota
Kinabalu districts (e.g., SAN 16242, SAN 38713 and SAN 66242) and in Sarawak from
Belaga, Bintulu, Kapit, Lawas, Limbang, Lundu, Miri, Simunjan, and Tatau districts (e.g., S
23743, S 25025 and S 27004). Also occurring in Brunei (e.g., BRUN 3340 and BRUN 5645)
and W Kalimantan (e.g., bb. 29609 and bb. 29610).
254
Ecology. Locally frequent in mixed dipterocarp forest on clayrich soils, on shale and
volcanic rocks, at altitudes to 700 m. Occurring in Lambir NP; elsewhere vulnerable owing
to land conversion.
38. Shorea falcifera Dyer ex Brandis

(Latin, falx = a sickle, ferre = to bear; carrying a sickle, the sickleshaped leaf)
J. Linn. Soc. Bot. 31 (1895) 64; Merrill op. cit. (1921) 571; Ashton op. cit. (1978) 36, op. cit. (1982)
456; PROSEA op. cit. 428; Newman et al. op. cit. (1998) 177. Type: Beccari PB 3046, Borneo,
Sarawak, Santubong (holotype K). Synonyms: Hopea linggensis Boerl., Cat. Hort. Bog. 2 (1901) 105;
Shorea flava Meijer op. cit. 325, Ashton op. cit. (1968) 69, Anderson op. cit. (1980) 119.
Emergent tree, to 45 m tall, to 1.4 m diameter, with seagreen diffuse hemispherical crown;
bole straight; buttresses to 1.5 m tall, fairly thin. Bark yellowish brown, patchily becoming
rich reddish brown, cracked and patchily oblong flaky. Twig, bud, stipules (sparsely so
within) venation, and blade below densely persistently pale yellow lepidote; inflorescence,
parts of perianth exposed in bud and nut densely persistently buffpuberulent; fruit calyx
sparsely caducously so; ovary and stylopodium sericeous. Twigs c. 1 mm diameter apically,
slender, terete. Leaf buds small, elliptic, subacute, to 2 × 1 mm. Stipules elliptic, obtuse, to
6 × 4 mm, fugaceous. Leaves coriaceous, drying mauvegrey above, cream below; blade
narrowly ovate to lanceolatefalcate, 6.5–12 × 2.5–5 cm; midrib shallowly sunken, slender
but evident above, prominent below; lateral veins c. 10 pairs, slender but slightly raised
below; intercostal venation obscure, scalariform; petiole slender, somewhat geniculate, 1.1–
1.8 cm long. Inflorescences terminal or axillary; rachis terete or ribbed, to 11 cm long,
singly branched, branchlets bearing to 5 flowers. Flowers: buds to 8 × 2 mm; petals cream;
stamens 33–34, filaments setose, anthers sparsely so on distal margin, connectival
appendage shorter than anther, shortly setose; ovary and stylopodium pyriform, style short,
glabrous. Fruits: calyx lobes unequal, 3 longer lobes to 9.5 × 1.8 cm, tapering to 7 mm
above the saccate base, 2 shorter ones to 7 × 0.8 cm, otherwise similar. Nuts ovoid, to 2.5
× 1.5 cm, with to 7 mm prominent filiform style remnant.
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo known in Sarawak from
Kuching, Lundu, Simunjan, and Sri Aman districts (e.g., S 10288, S 15433, S 25291, S
58976, and S 68888).
Ecology. Common in mixed dipterocarp forest on sandy soils on hills, often rocky, near the
sea, at altitudes to 200 m. Occurring in Bako NP; vulnerable outside parks system.
39. Shorea falciferoides Foxw.

(Greek, oides = resembling; similar to S. falcifera)
Philip. J. Sci. 13 (1918) Bot. 189. Lectotype (designated here): Mayor FB 25664, the Philippines,
Luzon, Zambales Province, Masinloc (hololectotype K). Synonym: Shorea gisok Foxw., op. cit.
(1938) 294.
255
Notes. Two subspecies, subsp. falciferoides and subsp. glaucescens are recognised, of
which the former is endemic in the Philippines.
subsp. glaucescens (Meijer) P.S.Ashton
Gard. Bull. Sing. 31 (1978) 37, op. cit. (1982) 458; PROSEA op. cit. 428; Coode et al. (eds.) op. cit.
77; Newman et al. op. cit. (1998) 178. Basionym: Shorea glaucescens Meijer op. cit. 327, Meijer &
Wood op. cit. (1964) 170, Ashton op. cit. (1964) 134, op. cit. (1968) 71, Burgess op. cit. 202,
Anderson op. cit. (1980) 119. Type: G.H.S. Wood & Agama SAN 15484, Borneo, Sabah, Sepilok FR
Emergent tree, to 50 m tall, to 1.1 m diameter; crown sparse, irregularly hemispherical; bole
straight or misshapen; buttresses to 3 m tall, spreading, thin. Bark reddish brown, becoming
cracked and irregularly oblong flaky. Young twig, inflorescence, petiole, leaf bud, stipules,
bracteole, parts of perianth exposed in bud, ovary, stylopodium, and nut densely evenly
persistently cream puberulent; fruit calyx sparsely so towards base. Twigs 2–3.5 mm
diameter apically, terete or somewhat compressed and ribbed; stipule scars c. 1.5 mm long,
pale, horizontal. Leaf buds globose to ovoid, subacute, slightly compressed, to 5 × 3.5 mm.
Stipules obtuse, to 10 × 4 mm, fugaceous. Leaves chartaceous, drying pale tawny above,
cream below; blade broadly ovatefalcate, 10–18 × 4.5–8 cm, base subequal, obtuse to
cuneate, apex with tapering acumen c. 0.8 cm long; midrib evident, flat to shallowly
furrowed above, prominent below; lateral veins 8–12 pairs, slender, wellspaced,
ascending, raised but not prominent below; intercostal venation slender, densely
scalariform, not raised; petiole 1.5–2 cm long, stout. Inflorescences terminal or axillary;
rachis ribbed and somewhat compressed, to 4 cm long, singly branched, branchlets bearing
to 6 close flowers; bracteoles elliptic, to 3 mm long, fugacious. Flowers: buds to 5 × 2.5
mm; petals cream; stamens c. 45, filaments and anthers glabrous, connectival appendage
very short, stout, shortly setose with one long apical bristle; ovary and stylopodium ovoid,
style short, glabrous. Fruits: calyx lobes unequal, 3 longer lobes to 9.5 × 2.5 cm, tapering
to 5 mm broad above the saccate base, 2 shorter ones to 7 × 1 cm, otherwise similar. Nuts
broadly ovoid, to 1.5 × 1.5 cm, with to 4 mm tapering style remnant.
Vernacular names. Sabah—selangan batu laut (preferred name). Sarawak—selangan batu

daun nipis (preferred name).
Distribution. Endemic in Borneo; widespread but absent from the W and SW parts. In
Sabah known from Beaufort, Beluran, Kudat, and Sandakan districts (e.g., SAN 16423, SAN
16523, SAN 16962, SAN 25439, and SAN 61155) and in Sarawak from Belaga, Bintulu,
Kapit, Lawas, Limbang, and Miri districts (e.g., S 14741, S 27130, S 29677, S 32312, and S
43355). Also occurring in Brunei (e.g., BRUN 795, BRUN 5718, KEP 80140, and S 2116)
and in E Kalimantan (e.g., Ambriansyah Berau 981 and bb. 29684).
Ecology. Locally frequent in mixed dipterocarp forest on clayrich soils, at altitudes to 600
m, especially on moist slopes. Occurring in Lambir NP and Sepilok FR; elsewhere
vulnerable owing to conversion of its habitat.
40. Shorea fallax Meijer

(Latin, fallax = deceptive; not at first clearly distinguished from S. scaberrima)
256
Act. Bot. Neerl. 12 (1963) 335; Meijer & Wood op. cit. 105; Ashton op. cit. (1964) 186, op. cit. (1968)
Coode et al. (eds.) op. cit. 77; Newman et al. op. cit. (1996) 140. Lectotype (designated here): G.H.S.
Wood SAN A 1734, Borneo, Sabah, Beaufort Hill (hololectotype K; isolectotypes KEP, L).
Synonyms: Shorea squamata auct. non Benth. & Hook.f.: Browne op. cit. 144; S. oleosa Meijer op.
cit. 338, Meijer & Wood op. cit. 124, Burgess op. cit. 183.
Emergent tree, to 50 m tall, to 1.3 m diameter but usually smaller; crown dense,
hemispherical; bole straight, cylindrical; buttresses to 2 m tall, stout. Bark warm brown,
deeply cracked and becoming oblong flaky; inner bark dull reddish brown; heartwood pink.
Young twig, inflorescence, leaf bud, parts of perianth exposed in bud, stipule and bracteole
outside, petiole, leaf below, and midrib above sparsely persistently pale brown scabrid
pubescent; ovary, stylopodium and nut densely evenly buffpubescent. Twigs 2–2.5 mm
diameter apically, terete, ribbed below the c. 2 mm long pale subfalcate stipule scars. Leaf
buds ovoid, subacute, 5–8 × 4–6 mm, large. Stipules hastate, to 15 × 5 mm, caducous.
Leaves coriaceous, drying chocolatebrown below, mauvegrey above; blade broadly
oblong to ovate, 12–24 × 5.5–11 cm, base obtuse, apex with tapering acumen to 1.2 cm
long; midrib evident, flat above, prominently terete below; lateral veins 15–19 pairs,
prominent below, arched, generally with many short intermediate veins; intercostal
venation scalariform, distant, prominent below; petiole 1–1.5 cm long. Inflorescences
terminal or axillary; rachis terete or ribbed, lax, to 22 cm long, singly or doubly branched,
branchlets bearing to 7 flowers; bracteoles ovate, acute, to 7 × 3 mm, not at first caducous.
Flowers: buds to 8 × 4 mm; petals creamyellow; stamens 15, connectival appendage c. 2x
the length of anther; ovary and stylopodium pyriform, style equally long, columnar,
glabrous. Fruits subsessile; calyx glabrescent, lobes unequal, 3 longer lobes to 5 × 0.8 cm,
tapering to 3.5 mm broad above the saccate base, 2 shorter ones linearlobed, to 2.5 × 0.15
cm, similar at base. Nuts ovoid, to 2.7 × 1 cm, apiculate.
Vernacular names. Sabah—seraya daun kasar (preferred name). Sarawak—engkabang

pinang (Iban), engkabang layar (preferred name).
Distribution. Endemic in Borneo. In Sabah known from Beaufort, Keningau,

Kinabatangan, Labuk Sugut, Sandakan, and Tawau districts (e.g., SAN 16466, SAN 28687,
SAN 36910, SAN 61257, and SAN 123774) and in Sarawak from Belaga, Bintulu, Kapit,
Lawas, Lundu, Marudi, Miri, and Tatau districts (e.g., S 15113, S 22386, S 31702, S 46429,
and S 53496). Also occurring in Brunei (e.g., BRUN 769 and BRUN 3065) and Kalimantan
(e.g., Endert 4783).
Ecology. Frequent in mixed dipterocarp forest, on clay soils mostly over shale, on moist
hillsides, low hills and alluvium, at altitudes to 600 m. Occurring in Danum Valley
Conservation Area and Sepilok FR in Sabah and Lambir and Mulu NPs in Sarawak;
elsewhere vulnerable owing to forest conversion.
41. Shorea ferruginea Dyer ex Brandis

(Latin, ferrugineus = rustcoloured; the dry leaf)
(sect. Mutica, subsect. Auriculatae, red meranti)
257
J. Linn. Soc. Bot. 31 (1895) 91; Merrill op. cit. (1921) 405; Masamune op. cit. 493; Browne op. cit.
77; Newman et al. op. cit. (1996) 142. Type: Beccari PB 2604, Borneo, Sarawak, Matang (holotype
K).
Emergent tree, to 45 m tall, to 1.6 m diameter, with diffuse hemispherical, cauliflower

shaped crown; bole straight, cylindrical; buttresses to 3.5 m tall, stout. Bark smooth at first,
becoming yellowish fawn, shallowly vsection fissured and powdery thinly oblong flaky,
occasionally becoming somewhat shaggy; inner bark and heartwood yellowish brown.
Young twig, inflorescence, leaf bud, parts of perianth exposed in bud, stipule and bracteole
outside (sparsely so within), petiole, and blade below persistently pale brown scabrid
puberulent; ovary, stylopodium and nut sparsely cream buff pubescent. Twigs 2–2.5 mm
diameter apically, terete, ribbed below the short pale falcate stipule scars. Leaf buds ovoid,
subacute, 5–8 × 4–6 mm. Stipules hastate, acute, to 15 × 5 mm. Leaves thinly coriaceous,
drying dull orangebrown below, paler above; blade broadly oblong to ovatelanceolate, 6–
15 × 3.3–5.3 cm, base obtuse, apex with slender acumen to 1.2 cm long; midrib obscure,
sunken above, prominent below; lateral veins 11–14 pairs, slender and hardly raised below;
intercostal venation slender, scalariform, obscure; petiole 1–1.5 cm long. Inflorescences
terminal or axillary; rachis terete or ribbed, lax, to 22 cm long, singly or doubly branched,
branchlets bearing to 7 flowers; bracteoles ovate, acute, to 7 × 3 mm, fugaceous. Flowers:
stamens 15, connectival appendage curved but not becoming fully reflexed; ovary and
stylopodium pyriform, style equally long, columnar, glabrous. Fruits subsessile; calyx
glabrescent; lobes unequal, 3 longer lobes to 12 × 3.5 cm, tapering to to 1.3 cm at the
auriculate base, saccate at the incrassate centre, 2 shorter ones linearlobed, to 4 × 0.8 cm,
hardly auriculate at base. Nuts ovoid, to 2.7 × 1 cm, prominently apiculate.
Vernacular names. Sabah—seraya melantai kecil (preferred name). Sarawak— engkabang

kali (Iban), meranti menalit (preferred name).
Distribution. Endemic in Borneo; widespread. In Sabah recorded from Beaufort, Kota

Merudu, Labuk Sugut, Nabawan, Ranau, and Tenom districts (e.g., SAN 15089, SAN 16363,
SAN 94903, SAN 99272, and SAN 99547) and in Sarawak from Belaga, Bintulu, Kapit,
Kuching, Lawas, Lubok Antu, Miri, and Samarahan districts (e.g., S 10703, S 15833, S
29640, S 36929, and S 68123). Also occurring in Brunei (e.g., BRUN 422 and BRUN 3396).
Ecology. Locally common in mixed and upper dipterocarp forests on dry sandy clay and
skeletal clay soils, especially on ridges, at altitudes to 1100 m. Occurring in Kinabalu,
Lambir and Mulu NPs; not vulnerable.
42. Shorea flaviflora Wood ex P.S.Ashton

(Latin, flavus = yellow, flos = flower; yellowflowered)
Wood op. cit. 107; Anderson op. cit. (1980) 126; Coode et al. (eds.) op. cit. 77; Newman et al. op. cit.
(1996) 518. Type: G.H.S. Wood SAN 15109, Borneo, Sabah, Beaufort, Pangi (holotype KEP; isotype
L).
258
Emergent tree, to 45 m tall, to 1.4 m diameter, with irregular crown; bole cylindrical or
frequently misshapen; buttresses to 4 m tall, stout. Bark dark brown to blackish, becoming
deeply vsection fissured and eventually thinly flaky, with occasional cream dammar
smears; inner bark rich crimson; heartwood dark yellowish brown. Young parts shortly pale
grey puberulent with minute adpressed hairtufts, caducous except on inflorescence, leaf
bud, stipule, and parts of perianth exposed in bud; ovary, stylopodium and nut densely buff
pubescent. Twigs 2–3 mm diameter apically, terete, glabrous, smooth; stipule scars narrow,
pale, ascending, half encircling twig. Leaf buds falcate, acute, 6–12 × 1.5–3 mm. Stipules
narrowly hastate, to 24 × 7 mm, caducous. Leaves thinly coriaceous, undulate, drying rich
fawnbrown below with paler veins, pale tawny above; blade narrowly ovate, 12–18 × 5.5–
7 cm, base broadly cuneate to obtuse, apex tapering to c. 1.5 cm slender acumen; midrib
flat, evident above, prominent below; lateral veins 6–7 pairs, ascending, arched, slender but
prominent below, with small porelike axillary domatia; intercostal venation very slender,
hardly raised, densely scalariform; petiole 2–3 cm long, geniculate. Inflorescences terminal
or axillary, pendent; rachis terete, to 15 cm long, doubly branched, branchlets bearing to 12
flowers; bracteoles elliptic, obtuse, to 5 × 2.5 mm, fugaceous. Flowers: buds large, to 14 ×
4.5 mm; petals rich yellow; stamens 15, anthers large, reniform, locules indistinct,
connectival appendage c. 3x the length of anther; ovary and stylopodium pyriform, style
filiform, as long as ovary and stylopodium, glabrous. Fruits: calyx glabrous, lobes unequal,
3 longer lobes to 13 × 3 cm, tapering to 11 mm above the saccate base, 2 shorter ones to 8
× 1.3 cm, otherwise similar. Nuts ovoid, to 2.5 × 1.3 cm, with tapering style remnant to 1
cm long.
Vernacular names. Sabah—seraya daun besar (preferred name). Sarawak—selangan

merah bukit (preferred name).
Distribution. Endemic in Borneo; known in Sabah from Beaufort, Ranau, Sipitang,

Tambunan, and Tenom districts (e.g., SAN 15098, SAN 16389, S 23484, SAN 24817, and
SAN 69870) and in Sarawak from Bintulu, Kapit, Limbang, and Tatau districts (e.g., S
5276 and BRUN 5664).
Ecology. Locally frequent in mixed dipterocarp but especially upper dipterocarp forest on
high shale ridges, at (150–)400–1300 m altitude. Occurring in Kinabalu and Mulu NPs;
elsewhere vulnerable owing to logging.
43. Shorea flemmichii Symington

(C.O. Flemmich, Forestry Officer in Peninsular Malaysia, who served in Brunei 1938–1941
and collected important dipterocarp types)
Gard. Bull. S. S. 10 (1939) 378; Masamune op. cit. 493; Browne op. cit. l.c. 147; Ashton op. cit.
(1964) 190, op. cit. (1968) 108, op. cit. (1982) 507; Anderson op. cit. (1980) 126; PROSEA op. cit.
395; Coode et al. (eds.) op. cit. 77; Newman et al. op. cit. (1996) 144. Type: Flemmich FMS 32585,
Borneo, Brunei, Belait district, Tenajor, Bt. Rotan (holotype KEP; isotype KEP).
Magnificent emergent tree, to 60 m tall, to 2.5 m diameter, with vast cauliflowershaped

crown suffused golden brown from below; bole tall, cylindrical; buttresses to 1.5 m tall,
stout. Bark purplish grey, black and ochremottled, becoming densely deeply vsection
259
fissured, the intervening ridges oblongflaky; dammar in prominent ochreyellow smears;

inner bark deep crimson; heartwood reddish rust. Young twig, inflorescence, leaf bud, parts
of perianth exposed in bud, stipule and bracteole outside, petiole, midrib below and at base
above, and venation below densely persistently golden tawny scabridtomentose; leaf above
scabridpuberulent; ovary and nut buffpubescent. Twigs stout though tapering to 2 mm
diameter apically, much branched, terete, verruculose. Leaf buds ovoid, subacute, slightly
compressed, 3–6 × 2–4 mm. Stipules narrowly hastate, acute, to 8 × 2.5 mm, fugaceous.
Leaves coriaceous, cream lepidote below, otherwise drying chocolatebrown; blade broadly
elliptic or ovate (young trees), 5–9 × 3–4.5 cm, base obtuse, margin revolute, apex with
narrow acumen to 0.5 cm long; lateral veins 14–17 pairs, dense, prominent below;
intercostal venation scalariform, sinuate; petiole 0.7–0.9 cm long, stout. Inflorescences
terminal or axillary; rachis terete or slightly compressed, to 8 cm long, singly or doubly
irregularly branched, branchlets somewhat zigzag, bearing to 8 flowers; bracteoles
suborbicular, obtuse, to 6 × 5 mm, caducous. Flowers: buds subglobose, to 4.5 × 3.5 mm;
corolla dark crimson; stamens 15, connectival appendage c. 3x the length of anther; ovary
and stylopodium ovoidconical, style filiform, glabrous, as long as ovary. Fruits subsessile;
calyx puberulent, calyx lobes unequal, 3 longer lobes to 6.5 × 1.5 cm, tapering to 2.5 mm
above the saccate base, 2 shorter ones linearlobed, to 2 × 0.15 cm, similar at base. Nuts
ovoid, to 1.5 × 1 cm, subacute,
Vernacular name. Sarawak—meranti raya (preferred name).
Distribution. Endemic in Borneo. In Sarawak known from Kuching, Lawas, Miri,

Samarahan, and Simunjan districts (e.g., S 6525, S 16753 and SFN 10246).
Ecology. Very local and rare or scattered in mixed dipterocarp forest on deep yellow sands,
at altitudes below 400 m. Well represented in Lambir NP; elsewhere critically endangered.
44. Shorea foraminifera P.S.Ashton

(Latin, foramen = opening, ferre = to bear; the prominent porelike leaf domatia)
Coode et al. (eds.) op. cit. 77; Newman et al. op. cit. (1996) 143. Type: Sabli S 1916, Borneo, Brunei,
Andulau FR (holotype K; isotype KEP).
Emergent tree, to 50 m tall, to 1.2 m diameter; bole straight, cylindrical; buttresses to 1 m

tall, stout. Bark pinkish brown, prominently vsection fissured; inner bark pinkish brown;
heartwood deep pink. Young parts, leaf and stipules excepted, fugaceous puberulent; calyx
sparsely puberulent towards base; nut persistently sericeous, known parts otherwise
glabrous. Twigs at first slightly compressed, 1–2 mm diameter apically. Leaf buds ovoid,
compressed, subacute, to 8 × 5 mm. Stipules oblong to elliptic, obtuse, to 10 × 5 mm,
caducous. Leaves coriaceous, drying warm brown below, greyish brown above; blade
broadly ovate, 6–9 × 4–7 cm, base cordate to obtuse, apex with somewhat falcate acumen
to 0.5 cm long; midrib obscure and slightly sunken above, prominent, terete, below; lateral
veins 8–9 pairs, prominent below, arched, with prominent pustular porous domatia;
intercostal venation very slender, scalariform, hardly raised; petiole 1–1.5 cm long.
Inflorescences and flowers unknown. Fruits: pedicels c. 1 mm long; calyx lobes
puberulent towards base, unequal, 3 longer lobes to 7 × 1.2 cm, tapering to c. 4 mm above
260
the saccate base, 2 shorter ones loratelobed, to 4.5 × 0.4 cm, similar at base. Nuts ovoid, to
1.5 × 1 cm, shortly apiculate.
Vernacular name. Sarawak—meranti lubang hidung (preferred name).
Distribution. Endemic in Borneo. Rare, known definitely from Brunei (e.g., BRUN 243,
FMS 28659, FMS 28675, S 1916, and SAN 17531). Fallen leaves observed at Similajau FR,
Bintulu district and Ulu Arip, Mukah district in Sarawak.
Ecology. Occasional, in small groups on periodically flooded sandy alluvium, including

shallow peat, at low altitude. Critically endangered.
45. Shorea foxworthyi Symington

(F.W. Foxworthy, 1877–1950, first research officer at the Forest Research Institute,
Kepong, Malaysia)
Gard. Bull. S. S. 8 (1935) 272, op. cit. (1943) 14; Keith op. cit. 44; Meijer & Wood op. cit. 169;
PROSEA op. cit. 428; Coode et al. (eds.) op. cit. 77; Newman et al. op. cit. (1998) 179. Type:
Foxworthy KEP 9507, Peninsular Malaysia, Pahang, Baloh FR (holotype KEP).
Large emergent tree, to 60 m tall, to 1.2 m diameter, with dense irregularly hemispherical
crown; bole straight, cylindrical; buttresses to 3 m tall, slender, spreading. Bark greyish
fawn, becoming irregularly cracked then shallowly flaked, and eventually shaggy leaving
scrollmarked surfaces below. Twig, inflorescence, leaf bud, parts of perianth exposed in
bud, stipule outside, and petiole densely shortly evenly goldentawny velutinous; leaf
venation below, and inside of stipules and bracteoles sparsely so, caducous on venation.
Twigs somewhat compressed, c. 2 × 3 mm apically. Leaf buds ellipsoid, obtuse, to 3 × 2
mm. Stipules narrowly elliptic to falcate, subacute, to 15 × 4 mm, caducous. Leaves
coriaceous, drying tawnybrown below; blade somewhat bullate in between lateral veins,
elliptic, 8–13 × 3–6.5 cm, base broadly cuneate, subequal, apex with broad acumen to 1 cm
long; midrib evident but somewhat sunken above, prominent below; lateral veins 10–14
pairs, ascending, arched, prominent below; intercostal venation slender, densely
scalariform; petiole 1.1–2 cm, drying rugose, velutinous. Inflorescences terminal or
axillary; rachis to 5 cm long, singly branched, branchlets bearing to 3 secund flowers;
bracteoles not seen. Flowers: buds to 10 × 3 mm; petals cream with a pink base; stamens
32–41, filaments densely setose, anther with sparsely setose distal margin, connectival
appendage densely setose, hardly exceeding anther; ovary ovoid, stylopodium conical,
glabrous, filiform, as long as ovary. Fruits: calyx lobes unequal, 3 longer lobes to 10 × 2.5
cm, tapering to 7 mm above the saccate base, 2 shorter ones linearlobed, to 8 × 1 cm,
similar at base. Nuts ovoid, to 2.5 × 1.4 cm, with prominent tapering style remnant to 6 mm
long.
Vernacular names. Sabah—selangan batu bersisik (preferred name). Sarawak—selangan

batu bukit (Malay).
Distribution. Sumatra, extreme SE Peninsular Thailand, Peninsular Malaysia, and Borneo.

In Sabah known from Lahad Datu, Sandakan, Sipitang, and Tawau districts (e.g., SAN
261
15183, SAN 19686 and SAN 36224); and in Sarawak from Bintulu, Kapit, Lawas, Marudi,
Miri, and Simunjan districts (e.g., S 4640, S 15149, S 18380, S 19577, and S 29622). Also
occurring in Brunei (e.g., BRUN 676) and Kalimantan (e.g., Wilkie 94306).
Ecology. Uncommon in mixed dipterocarp forest on yellow clay and sandy clays soils, at
altitudes to 700 m. Occurring in Lambir NP; elsewhere endangered owing to the loss of its
natural habitat.
46. Shorea geniculata Symington ex P.S.Ashton

(Latin, geniculatus = kneed; the petiole)
op. cit. (1980) 119; PROSEA op. cit. 429; Coode et al. (eds.) op. cit. 77; Newman et al. op. cit. (1998)
180. Type: Ashton BRUN 3264, Borneo, Brunei, Andulau FR (holotype K; isotypes KEP, L).
Large emergent tree, to 50 m tall, to 1.8 m diameter; crown irregularly hemispherical, pale
from below; bole straight, cylindrical; buttresses stout. Bark dark yellowish brown,
vertically cracked and irregularly flaked, sometimes becoming shaggy. Inflorescence,
bracteoles, parts of perianth exposed in bud, ovary, stylopodium, and fruit densely
persistently greyishbuff puberulent; vegetative parts glabrous. Twigs stout, terete, much
branched, c. 3 mm diameter apically, with swollen nodes. Leaf buds linearfalcate, 4–7 × 1–
1.5 mm. Stipules narrowly oblong, acute, to 10 × 3 mm, caducous. Leaves coriaceous,
drying cream lepidote below with black venation; blade broadly ovate to suborbicular, 11–
17 × 7–13 cm, base obtuse to subcordate, apex with short acumen to 0.8 cm long; midrib
stout, terete, slightly elevated above, more so below as also lateral veins; lateral veins 9–11
pairs, slender, distant, arched; intercostal venation densely scalariform, sinuate, slender;
petiole 4–6 cm long, geniculate. Inflorescences terminal or axillary; rachis terete, to 12 cm
long, singly branched, branchlets bearing to 4 flowers; bracteoles oblong, to 4 mm long,
fugaceous. Flowers: buds to 20 × 4 mm; petals cream; stamens c. 55, filament and anther
glabrous, connectival appendage somewhat shorter than anther, stout, setose; ovary and
stylopodium broadly ovoid, tapering abruptly, style short, glabrous. Fruits: calyx lobes
subequal, short, incrassate, deltoid, c. 1.5 × 1.5 cm, appressed to nut. Nuts globose, very
large, to 5 cm diameter, shortly mucronate.
Vernacular name. Sarawak—upun penyau (Malay, preferred name).
Distribution. Endemic in Borneo. Recorded in W Sarawak from Bau, Kuching, Miri, and
Samarahan districts (e.g., S 21420). Also occurring in Brunei (e.g., BRUN 3060, BRUN
3264, S 1929, and SAN 17470).
Ecology. Extremely local, but there frequent, in mixed dipterocarp forest on deep yellow
sands on low hills, at altitudes to 300 m. Occurring in Lambir NP; elsewhere critically
endangered owing to land conversion and logging.
47. Shorea gibbosa Brandis

(Latin, gibbosus = pouched; the staminal filament)
262
J. Linn. Soc. Bot. 31 (1895) 99; Symington op. cit. (1933) 143, op. cit. (1943) 51; Masamune op. cit.
493; Keith op. cit. 17, 23; Browne op. cit. 162; Meijer & Wood op. cit. 71; Burgess op. cit. 217;
Ashton op. cit. (1968) 83, op. cit. (1982) 482; Anderson op. cit. (1980) 122; PROSEA op. cit. 418;
Kessler & Sidiyasa op. cit. 103; Coode et al. (eds.) op. cit. 78; Newman et al. op. cit. (1996) 146.
Type: Ridley 6079, Singapore (holotype K). Synonym: Hopea grisea Brandis op. cit. 63.
Vast emergent tree, to 70 m tall, to 2 m diameter; bole tall, cylindrical; crown vast,
cauliflowershaped, somewhat diffuse; buttresses to 5 m tall, spreading, stout. Bark pale
tawny, becoming cracked and peeling in thin small oblong flakes, leaving a scrollmarked
surface. Twig, inflorescence, leaf bud, parts of perianth exposed in bud, stipules, bracteole,
ovary, nut, and petiole persistently shortly evenly pale buff to rufous puberulent; leaf
venation and fruit calyx sparsely so, glabrescent. Twigs c. 1 mm diameter apically, slender,
terete, muchbranched, pale greyish brown, striated. Leaf buds minute. Stipules narrowly
elliptic, to 3 × 1 mm, acute, fugaceous. Leaves chartaceous, undulate, pink when opening,
drying dark tawnybrown and wrinkling; blade ovate, 5–13 × 2–5 cm, base broadly cuneate
to obtuse, apex with tapering acumen to 1.5 cm long; midrib evident but somewhat furrowed
above, prominent below; lateral veins 7–9 pairs, slender but distinctly raised below, arched;
intercostal venation subreticulate, slightly elevated below; petiole 0.8–1.2(–1.6) cm long,
slender. Inflorescences terminal or axillary; rachis terete, to 10 cm long, singly branched,
branchlets bearing to 6 flowers; bracteoles oblong, obtuse, to 2 × 1 mm, fugaceous.
Flowers: buds to 5 × 3 mm; petals cream with pink base; stamens 15, connectival
appendage somewhat longer than anther, glabrous; ovary ovoid, style columnar, pubescent
in basal half. Fruits: pedicels to 2 mm long, slender; sepals unequal, 3 longer lobes to 9 ×
2 cm, tapering to 4 mm above the narrow tuberculate saccate base, 2 shorter ones to 6 ×
0.5 cm, otherwise similar. Nuts narrowly ellipsoid, acute, to 1.8 × 1.2 cm.
Vernacular names. Sabah—seraya kuning gajah (preferred name). Sarawak—lun gajah

(preferred name).
Distribution. SE Sumatra, Peninsular Malaysia, Singapore, and Borneo. In Sabah

widespread and recorded from Beaufort, Keningau, Kinabatangan, Kota Belud, Kota
Kinabalu, Labuk Sugut, Lahad Datu, Pensiangan, Ranau, Sandakan, Tambunan, Tawau, and
Tenom districts (e.g., SAN 9273, SAN 15452, SAN 16906, SAN 17004, and SAN 100174)
and in Sarawak from Bau, Belaga, Kapit, Kuching, Lundu, Marudi, Miri, and Tatau districts
(e.g., S 7971, S 15357, S 22090, S 43602, and S 43368). Also occurring in Brunei (BRUN
3128 and FMS 35461) and C, E and SE Kalimantan (e.g., bb. 13877, bb. 31384, Endert
5256, Kostermans 5261, and Kostermans 13247).
Ecology. Local dense populations occur in mixed dipterocarp forest on deep wellstructured
clay loams, particularly on low hills and undulating land over intermediate and igneous
rocks but rare, scattered, on the prevailing sedimentaries, at altitudes to 1000 m. Recorded
from Kubah and Mulu NPs; elsewhere endangered.
Notes. This variable species which can be confused with S. mujongensis which differs in its
glabrous leaf and twigs, and S. polyandra in which the leaf dries purplish brown with darker
venation below, and in which the inner bark is distinctly albeit faintly laminated.
48. Shorea gratissima (Wall. ex Kurz) Dyer
263
(Latin, gratus = pleasing; very pleasing)
Fl. Brit. Ind. 1 1874) 307; Symington op. cit. (1943) 36; Meijer & Wood op. cit. 54; Burgess op. cit.
159; Ashton op. cit. (1982) 487; PROSEA op. cit. 411; Newman et al. op. cit. (1996) 147. Basionym:
Hopea gratissima Wall., Cat. (1828) 960, nom. nud.; H. gratissima Wall. ex Kurz, J. As. Soc. Beng.
42, 2 (1873) 61. Type: Wallich s.n., Singapore (holotype K).
Magnificent tree, to 60 m tall, to 2 m diameter; bole straight or frequently twisted; crown

large, hemispherical with a few large twisted branches; buttresses to 2 m tall, stout. Bark
surface pale greyish fawn, shallowly vertically cracked and thinly oblong flaked; inner bark
laminated orangebrown and yellow. Exposed young parts evenly pale fulvous puberulent,
glabrescent except on stipules. Twigs c. 1.5 × 1 mm apically, at first somewhat compressed.
Leaf buds ovoid, subacute, to 2 × 1 mm. Stipules lanceolate, acute, to 10 × 3 mm, caducous.
Leaves thinly coriaceous, drying pale tawny with paler venation; blade ovate to elliptic, 4–
10 × 1.5–4.5 cm, base cuneate, margin undulate, apex acute or acuminate with slender
acumen to 1 cm long; midrib obscure and sunken above, slender and acute below; lateral
veins 12–14 pairs, slender, arched, hardly raised below; intercostal venation
subscalariform, obscure; petiole 0.8–1.5 cm long, slender. Inflorescences terminal or
axillary; rachis to 10 cm long, singly (if axillary) or doubly branched, branchlets bearing to
8 flowers; bracteoles to 2 mm long, linear, fugaceous. Flowers: buds to 5 × 3 mm; stamens
25, anthers oblong, tapering, connectival appendage c. 3x the length of anther, villous in the
distal half; ovary narrowly ovoid, without stylopodium, style c. 1½x as long as ovary.
Fruits: pedicels c. 1.5 mm long; calyx lobes unequal, 3 longer lobes loratespatulate, to 7 ×
1.3 cm, tapering to c. 6 mm above the saccate base, 2 shorter ones to 5.5 × 0.6 cm,
otherwise similar. Nuts ovoid, apiculate, to 1.5 × 0.8 cm.
Vernacular name. Sabah—melapi laut (preferred name).
Distribution. Myanmar, Peninsular Thailand, Sumatra, Peninsular Malaysia, Singapore,

and Borneo. In Borneo known in Sabah from Kudat, Lahad Datu and Tawau districts (e.g.,
SAN 15453, SAN 16408, SAN 20162, SAN 31644, and SAN A 4345). Also occurring in
Kalimantan (pers. obs.).
Ecology. Locally abundant in mixed dipterocarp forest on dry hills near the coast, at
altitudes to 500 m. Endangered.
49. Shorea guiso (Blanco) Blume

(from a Philippino name)
Mus. Bot. Lugd.Bat. 2 (1852) 343; Merrill, Sp. Blancoan. (1918) 270, PEB (1929) 203; Symington
op. cit. (1935) 266; Masamune op. cit. 494; Keith op. cit. 34; Browne op. cit. 153; Meijer & Wood op.
cit. 171; Burgess op. cit. 173; Ashton op. cit. (1968) 71, op. cit. (1982) 447; Anderson op. cit. (1980)
119; PROSEA op. cit. 429; Newman et al. op. cit. (1998) 181. Basionym: Mocanera guiso Blanco, Fl.
Filip. ed. 1 (1837) 449. Neotype (designated here): Merrill Sp. Blancoan. 407 (= US 904091), the
Philippines, Luzon, Bataan province, Limay (K, US). Synonyms: Euphoria malaanonan Blanco op.
cit. (1837) 286; Dipterocarpus guiso (Blanco) Blanco, Fl. Filip. ed. 2 (1845) 313; Anisoptera guiso
(Blanco) A.DC., op. cit. 616; Shorea pierrei Hance, J. Bot. 16 (1878) 302; S. vulgaris Pierre ex
264
Lanessan, Pl. Util. Colon. Fr. (1886) 301; S. vidaliana Brandis op. cit. 83; Isoptera burckii Boerl. op.
cit. 111; S. obtusa Wall. var. kohchangensis F.Heim, Bot. Tidsskr. (1902) 263; S. robusta Gaertn.f.
var. schmidtii F.Heim op. cit. (1902) 263; S. longipetala Foxw. op. cit. (1932) 174.
Large emergent tree, to 60 m tall, to 1.6 m diameter, with dense cauliflowershaped crown;
bole straight, cylindrical; buttresses to 3 m tall, long, thin, prominently concave. Bark
greyish tawny, becoming vertically shallowly cracked and narrowly oblong flaky; inner
bark rich pink, finely textured. Sapwood strawyellow merging to dark reddish brown
heartwood, moderately hard. Young parts at first greyish buff puberulent, early caducous
except on inflorescence, parts of perianth exposed in bud, ovary and nut; becoming sparsely
so as parts expand except on inflorescence and nut. Twigs slender, c. 1 mm diameter
apically. Leaf buds compressed, ovatefalcate, to 5 × 3 mm. Stipules oblong, subacute, to 7
× 5 mm, caducous. Leaves thinly coriaceous, drying undulate, purplish above, pale
chocolatebrown below with blackish veins and midrib; blade oblonglanceolate, 5–14 ×
2.5–6 cm, base obtuse to broadly cuneate, apex with broad, prominent acumen to 1 cm
long; midrib slender but prominent below, evident but somewhat furrowed above; lateral
veins (11–)15–19 pairs, slender but prominent below, dense; intercostal venation densely
scalariform, more or less obscure; petiole slender, somewhat geniculate, 1–1.8 cm long.
Inflorescences slender, lax, pendent; rachis to 10 cm long, branchlets bearing to 5 flowers;
bracteoles minute, linear, fugaceous. Flowers: buds slender, to 1 cm long; petals bright
yellow, red towards base within; stamens 20–28, anthers glabrous, connectival appendage
c. ½x the length of anther, with 1–4(–8) long terminal bristles; ovary ovoidconical, without
stylopodium, style columnar, c. ½x as long as ovary. Fruits: calyx lobes unequal, 3 longer
lobes to 5.5 × 1 cm, tapering to 3 mm broad above the saccate base, 3 shorter ones linear
lobed, to 3 cm long, similar at base. Nuts ovoid, apiculate, small, to 0.8 × 0.5 cm, hidden
within calyx lobes.
Vernacular name. Sabah and Sarawak—selangan batu merah (preferred name).
Distribution. CochinChina, Thailand, Sumatra, Peninsular Malaysia, Borneo, and the

Philippines. In Sabah locally abundant on the coastal hills in Kinabatangan, Kudat, Lahad
Datu, Sandakan, Semporna, and Tawau districts (e.g., SAN 5230, SAN 16409, SAN 29721,
SAN 38959, SAN 39221, and SAN 62396) and in Sarawak rare on limestone slopes in Bau,
Kapit and Miri districts (e.g., S 27265, S 24692, S 24694, and S 41542). Also occurring in E
Kalimantan (e.g., Kostermans 5889, Kostermans 13324 and Kostermans 13470).
Ecology. In mixed dipterocarp forest on dry sites, at altitudes to 300 m; locally common
along the east coast of Sabah, as also in the weakly seasonal tropics throughout its range.
Vulnerable.
50. Shorea havilandii Brandis

(G.D. Haviland, 1857–1901, first Sarawak Medical Officer, plant and insect collector)
J. Linn. Soc. Bot. 31 (1895) 82; Merrill op. cit. (1921) 405; Masamune op. cit. 494; Browne op. cit.
78; Newman et al. op. cit. (1998) 182. Type: Haviland 2395/1899, Borneo, Sarawak, near Kuching
(holotype K; isotype L). Synonym: Hopea ovalifolia Boerl. op. cit. 102.
265
Main canopy tree, to 40 m tall, to 50 cm diameter, with narrow dense crown; bole more or
less fluted and twisted; buttresses to 0.7 m tall, small, slender. Bark yellowish brown,
becoming thinly flaky eventually cracked and oblong flaky. Twig, petiole, midrib above,
leaf bud, stipule, inflorescence, bracteoles, parts of perianth exposed in bud, ovary, and nut
more or less persistently greyish tawny puberulent. Twigs terete, slender, c. 1 mm diameter
apically. Leaf buds ovoid, obtuse, c. 2 × 1.5 mm. Stipules oblong, subacute, c. 6 × 2.5 mm,
fugaceous. Leaves thinly coriaceous, drying pale chocolatebrown below, mauvebrown
above, curling; blade ovateelliptic, 8–16 × 2.5–6 cm, base obtuse to broadly cuneate, apex
with narrow acumen to 0.7 cm long; midrib slender but prominent below, evident but
grooved above; lateral veins 9–12 pairs, dense, slender but prominent below, with small
axillary pubescent domatia; intercostal venation slender, densely scalariform; petiole 0.8–
1.2 cm long. Inflorescences terminal or axillary; rachis straight, terete or slightly
compressed, to 12 cm long, singly branched, branchlets bearing to 7 flowers; bracteoles
elliptic, subacute, to 4 mm long, fugaceous. Flowers: buds to 8 × 2.5 mm; petals cream,
pink at base; stamens 30–50, filaments and anthers glabrous, connectival appendage short,
with single terminal bristle; ovary broadly ovoid, without stylopodium, style c. ½x the length
of ovary. Fruits: calyx lobes subequal, ovate, thin, narrowly acuminate, saccate, to 1 × 0.8
cm. Nuts globose, to 1.5 × 1 cm, obtuse but for 2 mm style remnant.
Vernacular name. Sabah and Sarawak—selangan batu pinang (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from Kinabatangan and Tawau

districts (e.g., SAN 16451, SAN 19490, SAN 23135, and SAN 26589) and in Sarawak from
Bintulu, Kuching, Lawas, Lundu, Marudi, and Miri districts (e.g., S 1517, S 4428, S 9620, S
27555, and S 46595). Also occurring in Brunei (e.g., BRUN 3007 and BRUN 5562)
wherever its habitat exists.
Ecology. Locally frequent, in kerangas on white sand podsols, on poorly drained white and
yellow sandy alluvium, also on organic soil over limestone, at altitudes to 400 m. Occurring
in Bako, Lambir and Mulu NPs; elsewhere vulnerable owing to land conversion.
51. Shorea hemsleyana (King) King ex Foxw.

(W.P. Hemsley, 1843–1924, botanist at Kew)
(sect. Mutica, subsect. Mutica, selangan batu)
Malay. For. Rec. 10 (1932) 167, p.p.; Symington op. cit. (1933) 129; Ashton op. cit. (1967) 293, op.
cit. (1982) 536. Basionym: Balanocarpus hemsleyanus King, J. As. Soc. Beng. 62, 2 (1893) 134.
Lectotype (designated here): King’s Collector (Kunstler) 6670, Peninsular Malaysia, Perak, Larut
(hololectotype K). Synonym: Pachychlamys hemsleyanus (King) Ridl., op. cit. (1922) 234.
Notes. Two subspecies, viz. subsp. hemsleyana and subsp. grandiflora, are recognised. The
former occurs in Peninsular Thailand, Sumatra and Peninsular Malaysia and the latter is
confined to Sarawak and W Kalimantan in Borneo.
subsp. grandiflora (Brandis) P.S.Ashton

(Latin, grandis = large, flora = flower; largeflowered)
Gard. Bull. Sing. 22 (1967) 293, op. cit. (1968) 109, op. cit. (1982) 536; Browne op. cit. 147;
Anderson op. cit. (1980) 126; PROSEA op. cit. 395; Newman et al. op. cit. (1996) 147. Basionym:
266
Shorea grandiflora Brandis op. cit. 93. Type: Haviland 2121, Borneo, Sarawak, near Kuching
(holotype K).
Main canopy or subcanopy tree, to 20 m tall, to 40 cm diameter, with dense hemispherical

crown; bole cylindrical; buttresses low, rounded. Bark pink and greymottled, becoming
shallowly densely vsection fissured; inner bark yellowish brown, faintly laminated;
heartwood pink. Twig, inflorescence, leaf bud, parts of perianth exposed in bud, stipule and
bracteole outside, petiole, and leaf venation below densely persistently more or less
yellowish brown to fulvous scabridtomentose; fruit calyx lobes, ovary and nut, stipule and
bracteole inside, and midrib above evenly so. Twigs terete, stout, 2–4 mm diameter
apically. Leaf buds lanceolate, subacute, to 6 × 3 mm. Stipules lanceolate, subacute, to 16
× 5 mm, not at first caducous. Leaves coriaceous, drying pale mauvebrown above, rufous
scabrid below; blade oblong, 10–23 × 4–11 cm; midrib and venation prominent below,
somewhat furrowed above; lateral veins 14–17 pairs; intercostal venation remotely
scalariform; petiole short, stout, 0.6–1.2 cm long. Inflorescences axillary; rachis terete, to 4
cm long, singly branched, flowers crowded on short branchlets; bracteole elliptic, obtuse, to
3 mm. Flowers: buds large, to 18 × 6 mm; petals dark red within, pink outside; stamens 15,
connectival appendage becoming reflexed; ovary ovoid, stylopodium short, pubescent, style
as long as ovary, glabrous. Fruits subsessile; calyx lobes subequal, 3 longer lobes as short
as ripe nut, lanceolate, acute, chartaceous, to 2.5 × 1.8 cm, with thickened and saccate
base, 2 shorter ones ovate, to 2 × 1.8 cm, otherwise similar. Nuts ovoid, to 7 × 3 cm,
shortly apiculate.
Distribution. Endemic in Borneo; recorded from Kapit, Kuching and Lundu districts in
Sarawak (e.g., S 584, S 18860, S 35964, and S 35966) and Lower Kapuas, in W Kalimantan.
Ecology. Rare, in mixed dipterocarp forest on yellow leached sandy clay soils, and on
shallow peat, at altitudes to 600 m. Recorded from Kubah NP; elsewhere endangered.
52. Shorea hopeifolia (F.Heim) Symington

(Latin, Hopealike, folia = leaf)
Gard. Bull. S. S. 7 (1933) 150, Gard. Bull. S. S. 8 (1934) 36, op. cit. (1943) 52; Masamune op. cit.
494; Browne op. cit. 163; Meijer & Wood op. cit. 73; Burgess op. cit. 217; Ashton op. cit. (1968) 84,
op. cit. (1978) 43, op. cit. (1982) 482; Anderson op. cit. (1980) 122; PROSEA op. cit. 419; Coode et
al. (eds.) op. cit. 78; Newman et al. op. cit. (1996) 148. Basionym: Cotylelobium hopeifolium F.Heim
op. cit. (1891) 971. Type: Beccari PB 2491, Borneo, Sarawak, Matang (holotype P). Synonyms:
Shorea ridleyana King op. cit. 115, p.p.; Hopea heimiana Brandis op. cit. 63, Merrill op. cit. (1921)
402; H. albescens Ridl., J. Str. Br. Roy. As. Soc. 73 (1916)142; S. kalunti Merr., Philip J. Sci. 26
(1925) 475; H. hopeifolia (F.Heim) Slooten, Bull. Jard. Bot. Buitenz. 3, 10 (1929) 396.
Vast and elegant emergent tree, to 65 m tall, to 2 m diameter, with large diffuse cauliflower
shaped crown; bole majestic, columnar; buttresses massive to 4 m tall, spreading, stout.
Bark tawny brown, shallowly vertically cracked and thinly oblong flaky; dammar
coxcombs few, small, blackish. Vegetative parts glabrous; inflorescence, bracteole outside,
parts of perianth exposed in bud, ovary, base of style and nut persistently pale buff woolly
puberulent; fruit calyx glabrescent except at base. Twigs slender, muchbranched, terete,
minutely rugulose, c. 1 mm diameter apically. Leaf buds ovoid, acute, minute, c. 1 × 1 mm.
Stipules lanceolate, acute, to 5 × 2 mm, fugaceous. Leaves somewhat chartaceous and
267
undulate, drying dark tawnybrown with the midrib distinctly dark red below; blade ovate,
3.5–8 × 2–4 cm, base broadly cuneate, apex with prominent acumen to 1 cm long; midrib
flat above, prominent below, usually with prominent porelike domatia either side at base;
lateral veins 9–11 pairs, with distinct shorter intermediates, slender, hardly raised below,
arched; intercostal venation reticulate, evident; petiole slender, geniculate, 0.8–1 cm long.
bearing to 9 flowers; bracteoles elliptic, obtuse, to 2 × 2 mm. Flowers: buds to 5 × 2 mm;
petals pale yellow; stamens 15, connectival appendage slightly shorter than anther; ovary
ovoid, without stylopodium, style columnar. Fruits: pedicels to 3 mm long; calyx lobes
unequal, 3 longer lobes to 7 × 1.5 cm, tapering to 4 mm above the saccate tuberculate base,
2 shorter ones to 4 × 0.7 cm, otherwise similar. Nuts ellipsoid, to 2.7 × 1.2 cm, shortly
apiculate.
Vernacular names. Sabah—seraya kuning jantan (preferred name). Sarawak—lun siput

jantan (preferred name).
Distribution. Sumatra, Peninsular Malaysia, Borneo, and the Philippines (Mindanao). In

Sabah, recorded from Beaufort, Tambunan, Tawau, and Tenom districts (e.g., SAN 15104,
SAN 17161, SAN 19492, SAN 32051, and SAN A 4313) and in Sarawak from Bintulu,
Kuching, Lundu, Miri and Tatau districts (e.g., S 9603, S 20285, S 25041, S 32446, and S
53414). Also occurring in Brunei (BRUN 2529 and FMS 30383) and C Kalimantan (e.g.,
Argent et al. 9461, bb. 24181 and Jarvie & Ruskandi 5771).
Ecology. As S. gibbosa and often occurring with it, but less common; occasionally to 1200
m altitude. Recorded from Lambir NP; elsewhere endangered.
Notes. Easily confused with S. multiflora but the midrib here dries dark red below, while the
tree becomes vastly taller.
53. Shorea hypoleuca Meijer

(Greek, hypo = underneath, leucos = white; the leaf undersurface)
Act. Bot. Neerl. 12 (1963) 329; Meijer & Wood op. cit. 174; Burgess op. cit. 202; Ashton op. cit.
(1968) 72, op. cit. (1982) 459; Anderson op. cit. (1980) 119; Newman et al. op. cit. (1998) 183.
Lectotype (designated here): G.H.S. Wood SAN 16048, Borneo, Sabah, Sandakan district, Sepilok FR
(hololectotype K).
Large emergent tree, to 60 m tall, to 1.8 m diameter; bole tall, cylindrical; crown
hemispherical, pale from below; buttresses low, thin. Bark fawnbrown, irregularly flaky.
Midrib above, leaf bud, inflorescence, parts of perianth exposed in bud, ovary, nut, stipule,
and bracteole shortly evenly persistently ochreous puberulent; fruit calyx glabrescent except
towards base; mature tree twig, midrib above, petiole, and leaf blade below yellowish
cream lepidote. Twigs ribbed at first, becoming terete, 1.5–2 mm diameter apically. Leaf
buds ovoid, obtuse, to 3 × 2 mm. Stipules oblongovate, to 7 × 4 mm, subacute, fugaceous.
Leaves coriaceous, drying tawnybrown above, pale yellow lepidote below; blade ovate to
narrowly elliptic, 8.5–17 × 3.5–8 cm, base cuneate to obtuse, margin more or less undulate
distally, apex with slender acumen to 1.5 cm long; midrib and lateral veins evident but
furrowed above, prominent below; lateral veins 11–16 pairs, ascending, with small
268
glabrous axillary domatia; intercostal venation scalariform, unraised; petiole 1–1.6 cm

long. Inflorescences terminal or axillary; rachis ribbed, to 14 cm long, singly branched,
branchlets bearing to 5 flowers; bracteoles small, fugaceous. Flowers: buds to 10 × 2 mm;
petals pale yellow; stamens c. 33, filaments and anthers glabrous, connectival appendage as
long as anther, setose with the apical bristles longer; ovary and stylopodium pyriform, style
short, glabrous. Fruits: calyx lobes unequal, 3 longer lobes to 8.5 × 1.7 cm, tapering to c. 3
mm above the saccate base, 2 shorter ones to 6 × 0.4 cm, similar at base. Nuts ovoid, to 1.5
× 1 cm, with to 3 mm long tapering stylopodium.
Vernacular name. Sabah and Sarawak—selangan batu kelabu (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from Beaufort, Beluran, Kudat, Lahad
Datu, Sandakan, and Tawau districts (e.g., FMS 35636, KEP 80505, SAN 36654, SAN
37368, and SAN A 214) and in Sarawak from Lawas and Lundu districts (e.g., S 1751, S
7967, S 10158, and S 10175).
Ecology. In mixed dipterocarp forest on wellstructured clay soils over sedimentary and
basic to intermediate igneous rocks, and around the base of limestone hills, at altitudes to
400 m; local but frequent where it occurs. Occurring in Sepilok FR; elsewhere vulnerable,
possibly endangered, by forest conversion.
54. Shorea iliasii P.S.Ashton

(Ilias Paie, research assistant and sometime curator of the Kuching herbarium, avid and
discriminating plant hunter)
Newman et al. op. cit. (1996) 149. Type: Ilias S 15102, Borneo, Sarawak, Bintulu district, Ulu Segan
FR (holotype K).
Emergent tree, to 50 m tall, to 1 m diameter; crown dense, hemispherical; bole cylindrical,

tall; buttresses to 2 m tall, stout. Bark fawnbrown, becoming irregularly flaky. Young parts
greyish sericeous caducous except on inflorescence, parts of perianth exposed in bud, base
of fruit calyx, ovary, and nut. Twigs slightly compressed at first, minutely striated, c. 2 mm
diameter apically. Leaf buds conical, subacute, c. 2 × 2 mm. Stipules not seen. Leaves
coriaceous, drying tawnybrown, darker above than below; blade oblongovate, 19–25 × 9–
12 cm, base subequal, obtuse but shortly decurrent with petiole, margin narrowly revolute,
apex with broad, short acumen to 0.8 cm long; midrib flat or slightly raised above,
prominent below; lateral veins 12–14 pairs, prominent below, obscurely furrowed above,
arched; intercostal venation remotely subscalariform; petiole 2.2–3 cm long, stout, 0.3–0.4
cm diameter. Inflorescences terminal or axillary; rachis terete or compressed; bracteoles not
seen. Flowers: buds to 4 × 2 mm; petals cream; stamens 15, connectival appendage c. 1½x
the length of anther; ovary narrowly ovoid, without stylopodium, style short, glabrous.
Fruits: calyx lobes unequal, 3 longer lobes to 8 × 2.2 cm, tapering to c. 4 mm above the
tuberculate saccate base, 2 shorter ones to 5.5 × 1.2 cm, otherwise similar. Nuts ovoid, to 1
× 0.8 cm, acute.
269
Vernacular name. Sarawak—lun siput daun besar (preferred name).
Distribution. Endemic in Borneo. Known in Sarawak from Belaga, Bintulu, Kapit, and
Lubok Antu districts (e.g., S 14455, S 22041, S 29604, S 33551, and S 43608) and from Ulu
Barito in C Kalimantan.
Ecology. Locally frequent in mixed dipterocarp forest on yellow sandy clay soils on
sedimentary rocks, on undulating land and hillsides, at altitudes to 400 m. Vulnerable owing
to land conversion.
55. Shorea inaequilateralis Symington

(Latin, inaequalateralis = unequalsided; the leaf base)
(sect. Brachypterae, red selangan, red meranti)
Gard. Bull. S. S. 8 (1935) 281; Masamune op. cit. 494; Browne op. cit. 153; Anderson op. cit. (1963)
158, op. cit. (1980) 126; Ashton op. cit. (1964) 191, op. cit. (1968) 109, op. cit. (1982) 505; PROSEA
op. cit. 395; Coode et al. (eds.) op. cit. 78; Newman et al. op. cit. (1996) 505. Lectotype (designated
here): Zainal FMS 30351, Borneo, Brunei, Kuala Belait (hololectotype KEP).
Noble emergent tree, to 45 m tall, to 1.2 m diameter, with large crown and pendent
branches; bole often somewhat sinuate; buttresses to 2 m tall, stout, spreading. Bark
becoming deeply densely vsection fissured and narrowly oblong flaky, reddish brown and
greymottled; inner bark deep reddish brown, fibrous; heartwood hard, deep reddish brown.
Young parts densely tawny pubescent, persistent on young twigs; leaf bud, outside of stipule,
inflorescence, bracteole, parts of perianth exposed in bud, ovary, nut, petiole, and midrib
above sparsely shortly so; appearing glabrescent on inside of stipule, leaf blade on both
surfaces, and venation below. Twigs c. 2.5 × 1.5 mm apically, compressed at first,
becoming terete, ribbed below the petiole insertion; stipule scars falcate, to 1.5 mm long.
Leaf buds ovoid, acute, somewhat compressed, 3–6 × 1.5–1.5 mm. Stipules not at first
caducous, narrowly hastate, acute, to 20 × 5 mm. Leaves chartaceous, drying tawny
brown; blade ovate, 9–14 × 4.5–7.5 cm, base strongly unequal, subcordate, apex with
prominent caudate acumen to 2 cm long; midrib evident, flat above, slender, terete but
prominent below; lateral veins 11–13 pairs, slender, raised below, arched; intercostal
venation scalariform, wellspaced; petiole stout, 0.5–0.8 cm long. Inflorescences terminal
or axillary; rachis terete, pendent, to 7 cm long, branchlets bearing to 4 flowers; bracteoles
narrowly deltoid, to 10 × 3 mm. Flowers: buds to 12 × 3 mm; petals red; stamens 15,
connectival appendage c. 3x as long as anther; ovary ovoid, without distinct stylopodium,
style c. 2x as long as ovary. Fruits: calyx lobes unequal, 3 longer lobes narrow, to 14 × 1.2
cm, tapering to c. 5 mm above the saccate base, 2 shorter ones to 12 × 0.9 cm, similar at
base. Nuts ovoid, to 1.8 × 1.4 cm, with to 6 mm slender filiform style remnant.
Vernacular name. Sarawak—semayur (preferred name).
270
Distribution. Endemic in NW Borneo. Known in Sarawak from Marudi and Sibu districts
(e.g., KEP 79319, S 1263 and S 11244) and in Brunei from Belait district (e.g., BRUN 990,
FMS 30499, FMS 35705, and S 2229).
Ecology. Very local, but there semigregarious, on the banks of peat swamp streams into the
mixed peat swamp forest. Critically endangered.
56. Shorea inappendiculata Burck

(Latin, inappendiculatus = lacking an appendix; the short connectival appendage of the
stamen)
cit. 171; Meijer & Wood op. cit. 175; Burgess op. cit. 211; Ashton op. cit. (1968) 73, op. cit. (1982)
455; Anderson op. cit. (1980) 120; PROSEA op. cit. 430; Coode et al. (eds.) op. cit. 455; Newman et
al. op. cit. (1996) 455. Type: Beccari PB 3009, Borneo, Sarawak, Matang (holotype BO).
Large emergent tree, to 60 m tall, to 1.5 m diameter; crown large, hemispherical; bole tall,
cylindrical; buttresses to 3 m tall, thin, spreading. Bark greyish fawn, thinly irregularly
flaky. Twigs, leaf buds, stipules, inflorescence, parts of perianth exposed in bud, and
bracteole densely persistently rufous scabridpubescent; leaf venation below sparsely so
(more dense on young trees), ovary and nut evenly so; blade below more or less densely
silvery lepidote. Twigs 2–4 mm diameter apically, prominently ribbed and somewhat
compressed at first, blackish with pale horizontal stipule scars. Leaf buds ovoid, conical, to
6 × 4 mm. Stipules broadly lanceolate, to 10 × 4 mm. Leaves coriaceous, often billowed
between lateral veins, drying purplish brown above, more or less grey to silvery lepidote
below with black venation; blade oblongelliptic, (7–)10–16 × (2–)3–8 cm, base more or
less unequal, obtuse to cordate, apex shortly acuminate to obtuse; midrib and lateral veins
prominent below, evident but furrowed above; lateral veins 13–24 pairs; intercostal
venation densely scalariform, slightly elevated below; petiole stout, 1.5–3 cm long.
Inflorescences axillary; rachis ribbed when dry, to 10 cm long, singly branched, branchlet
bearing to 7 flowers; bracteoles elliptic, obtuse, to 3 × 2 mm, fugaceous. Flowers: buds to 7
× 3 mm; stamens 28–34, filaments and anthers setose distally, connectival appendage as
long as or longer than outer anther, with a few terminal bristles; ovary and stylopodium
ovoid, style short. Fruits: pedicels to 4 mm long, broadening into receptacle; calyx lobes
unequal, 3 longer lobes to 13 × 3 cm, tapering to c. 8 mm above the saccate base, 2 shorter
ones loratelobed, to 10 × 0.9 cm, similar at base. Nuts ovoid, to 3.3 × 1.6 cm, prominently
apiculate.
Distribution. N Sumatra, Peninsular Malaysia (NW Johor) and Borneo. In Sabah known
from Kinabatangan and Tawau districts (e.g., SAN 16454, SAN 17833 and SAN 23270) and
in Sarawak from Bintulu, Kapit, Lundu, Miri, Samarahan, and Tatau districts (e.g., S 294, S
10060, S 15527, S 15885, and S 41128). Also occurring in Brunei (e.g., BRUN 3080) and E
Kalimantan.
Ecology. Scattered in mixed dipterocarp forest on wellstructured clay soils, mostly in the
coastal hills, at altitudes to 400 m; apparently generally rare, as single individuals with little
regeneration. Recorded from Kubah and Lambir NPs; elsewhere endangered.
271
Fig. 24. Shorea isoptera. A, flowering leafy twig; B, part of inflorescence; C, flower bud;
D, adaxial view of outer sepal; E, adaxial view of inner sepal; F, abaxial view of petal; G,
adaxial view of petal; H, longitudinal section of flower bud; I, flower bud with two sepals,
petals and stamens removed; J, adaxial view of stamens; K, abaxial view of stamens; L,
fruit; M, longitudinal section of fruit. (A–K from S 7978, L–M from SAN 39018.)
272
57. Shorea induplicata Slooten

(Latin, induplicatus = sharply turned in; the leaf margins)
Newman et al. op. cit. (1996) 151. Type: Knight SA 582, Borneo, Sarawak, Kuching district, Sampadi
FR (holotype KEP).
Canopy or low emergent tree, to 40 m tall, to 60 cm diameter; bole frequently misshapen;

crown dense, small, hemispherical, conspicuously creamcoloured in new leaf; buttresses
low but continuing up bole as ribs to 3 m tall. Bark grey and rustmottled, becoming thinly
irregularly flaky. Young parts pale rufous scabridpubescent, turning dark greyish brown
and persisting on twig, leaf bud, stipules, petiole, and venation below; leaf below at first
creampuberulent, turning pale grey. Twigs terete, becoming verruculose, c. 2 mm diameter
apically. Leaf buds ellipsoid, obtuse, to 3 × 2 mm. Stipules linear, not at first caducous, to
5 × 2 mm. Leaves coriaceous, drying pale greyish green above, pale grey below; blade
lanceolate, 8–17 × 2–5 cm, base subequal, cordate, margin prominently narrowly revolute,
apex with slender acumen to 2 cm long; midrib evident but shallowly furrowed above,
prominent below; lateral veins 12–16 pairs, prominent below; intercostal venation laxly
subscalariform; petiole rather short, 0.6–1.3 cm long. Inflorescences and flowers
unknown. Fruits: pedicels to 2 mm long; calyx glabrescent, lobes subequal, ovate, acute,
incrassate, the apices becoming patent, to 0.8 × 0.6 cm. Nuts broadly ellipsoid to obovoid,
to 2.5 × 2 cm, subacute.
Vernacular name. Sarawak—lun putih (preferred name).
Distribution. Endemic in the extreme NW Borneo. Known in Sarawak from Sampadi FR,
Kuching district (e.g., S 12611, S 18858 and SFN 10399) and in W Kalimantan from
Sanggau district (e.g., bb. 17608).
Ecology. Rare, in the mixed dipterocarp forestkerangas ecotone, on leached pale yellow
and grey sandy soils, at altitude below 200 m. Critically endangered, perhaps extinct.
58. Shorea isoptera P.S.Ashton Fig. 24.

(Greek, iso = same, pteron = wing; the subequal fruit calyx lobes)
(sect. Neohopea, selangan batu)
Newman et al. op. cit. (1998) 185. Type: Ashton BRUN 3018, Borneo, Brunei, Temburong, Bt. Patoi
(holotype K; isotype L).
Emergent tree, to 60 m tall, to 1.5 m diameter, with stout cylindrical bole; crown dense,
hemispherical; buttresses stout, low. Bark greyish tawny, becoming irregularly rather thinly
flaky; inner bark thin, pale pinkish brown, eventextured. Sapwood straw yellow, hard;
heartwood dark brown; dammar exudations greyish white. All vegetative parts glabrous;
inflorescence, bracteole, parts of perianth exposed in bud, apex of ovary, and nut
persistently greyish puberulent. Twigs smooth, sparsely branched, slender, c. 1.5 mm
273
diameter apically. Leaf buds globose, small, c. 1.5 mm diameter. Stipules caducous, not
seen. Leaves thinly coriaceous, satiny, drying tawnygrey; blade ovate, 9–16 × 6–8 cm,
base broadly cuneate to obtuse, decurrent to 2 mm down petiole (peltate in juveniles),
margin undulate, apex with slender acumen to 1.5 cm long; midrib terete, elevated below,
flat above; lateral veins 9–11 pairs, sunken above, raised below, slender, arched, sometimes
with a few minute porelike axillary domatia; intercostal venation sinuate, scalariform, very
slender, unraised; petiole slender, 1.3–2 cm long. Inflorescences terminal or to 3axillary;
rachis slender, terete or somewhat compressed, lax, regularly doubly branched, branchlets
bearing to 6 close flowers; bracteoles linear, to 1 mm long, fugaceous. Flowers: buds
globose, to 1.5 mm long; sepals subequal, imbricate, acute, 3 outer lobes ovate, 2 inner ones
deltoid; petals oblong, obtuse, contorted, falling separately; stamens 15, filaments broad,
short, glabrous, anthers broadly oblong, outer anther locules slightly larger than that of the
inner ones, connectival appendage short, stout, glabrous; ovary and stylopodium ovoid
conical, style short glabrous. Fruits: calyx lobes subequal, lobes to 5.5 × 1.5 cm, tapering
to 5 mm broad above the short saccate base, becoming rotate when ripe, barely twisted.
Nuts ovoid, to 0.9 × 1 cm, with to 4 mm tapering style remnant, on a broad shallow
receptacle.
Vernacular names. Sabah—selangan batu gelombang (preferred name). Sarawak—

selangan batu bulu ayam (preferred name).
Distribution. Endemic in Borneo, widespread but local. In Sabah, known from Beaufort,
Kota Belud, Sandakan, Sipitang, and Tenom districts (e.g., SAN 15111, SAN 16603, SAN
36909, SAN 39018, and SAN A 4319) and in Sarawak from Bau, Kuching, Lundu, and
Marudi districts (e.g., S 7978, S 10169, S 22828, S 22829, and S 32659). Also occurring in
Brunei (BRUN 3343, BRUN 5676 and KEP 80132) and E Kalimantan.
Ecology. Very local in mixed dipterocarp forest on clayrich soils over shale and
intemediate igneous rocks, at altitudes to 500 m; rather rare. Occurring in G. Gading NP;
elsewhere endangered as its habitat is choice for oil palm plantation.
59. Shorea johorensis Foxw.

(of Johor, Peninsular Malaysia)
(1982) 513; PROSEA op. cit. 395; Kessler & Sidiyasa op. cit. 103; Coode et al. (eds.) op. cit. 78;
Newman et al. op. cit. (1996) 152. Type: Bain KEP 5992, Peninsular Malaysia, Johor, G. Panti
(holotype KEP). Synonym: Shorea leptoclados Symington op. cit. (1939) 376, op. cit. (1943) 77,
Keith op. cit. 12, Browne op. cit. 140, Ashton op. cit. (1964) 195, op. cit. (1968) 110, Meijer & Wood
op. cit. 112, Burgess op. cit. 154, 184, Anderson op. cit. (1980) 126.
Large emergent tree, to 50 m tall, to 1.6 m diameter; crown large, hemispherical; bole tall,
straight, symmetrical; buttresses to 3 m tall, stout, prominent. Bark tawnygrey to fawn,
overall appearing smooth, shallowly densely cracked and thinly oblong flaky. Twig,
inflorescence, leaf bud, parts of perianth exposed in bud, stipules, bracteole, ovary, nut,
petiole, venation below, and midrib above shortly evenly persistently greyish buff pubescent;
274
young leaf caducously so. Twigs 2–3 mm diameter apically, somewhat ribbed and
compressed at first, much branched; stipule scars to 2 mm long, pale, falcate, descending.
Leaf buds ovoid, compressed, 6–8 × 2–3 mm. Stipules lanceolate, to 35 × 7 mm, caducous.
Leaves chartaceous, drying tawnygrey below; blade ovate, 9–14 × 4.5–7.5 cm, base obtuse
to subcordate, apex with slender and tapering acumen to 0.7 cm long; midrib evident, more
or less flat, above, slender but prominent below; lateral veins 10–12 pairs, slender but
prominent below, arched towards their ends, the basal 3–6 pairs usually with paired scale
like domatia; intercostal venation densely scalariform, very slender; petiole 1.5–2 cm long.
Inflorescences terminal or axillary; rachis terete or somewhat compressed, slender, to 15
cm long, branchlets long, bearing to 18 flowers; bracteoles narrowly ovate, obtuse, to 7 mm
long, caducous. Flowers: buds to 8 × 3 mm; petals pale yellow; stamens 15, connectival
appendage c. 3x as long as anther; ovary globose, stylopodium indistinct, style 2x as long as
ovary, filiform. Fruits: calyx lobes unequal, 3 longer lobes to 12 × 2.3 cm, tapering to c. 6
mm above the saccate base, 2 shorter ones linearlobed, to 6.5 × 0.6 cm, similar at base.
Nuts broadly ovoid, to 2 × 1.4 cm, shortly apiculate.
Vernacular names. Sabah—seraya majau (Malay, preferred name). Sarawak—meranti

majau (Malay, preferred name).
Distribution. E Sumatra, SE Peninsular Malaysia and Borneo. In Sabah recorded from

Beaufort, Beluran, Kinabatangan, Lahad Datu, Sandakan, Sipitang, and Tawau districts
(e.g., SAN 16620, SAN 17233, SAN 17952, SAN 93829, and SAN A 512) and in Sarawak
from Kapit, Lawas, Limbang, Marudi, and Miri districts (e.g., S 1527, S 1802 and S 29616).
Also occurring in Brunei (e.g., BRUN 355, FMS 30533 and S 1951) and E and C
Kalimantan (e.g., Sidiyasa 1121).
Ecology. Formerly one of the commonest light red meranti species in the eastern part of its
Borneo range, uncommon to the west; in mixed dipterocarp forest on moist clay and silty
soils, especially on low hills and floodplains; occasionally at altitude to 800 m. Occurring in
Lambir and Mulu NPs; elsewhere its natural habitat is greatly reduced by forest conversion;
vulnerable.
Notes. The dull leaden dry leaf with several pairs of small pale scalelike domatia in the
axils of the first few pairs of lateral veins distinguishes this species from S. palembanica and
S. pauciflora.
60. Shorea kudatensis Wood ex Meijer

(of Kudat, Sabah)
(1982) 483; Newman et al. op. cit. (1996) 155. Lectotype (designated here): G.H.S Wood SAN 15363,
Borneo, Sabah, Kudat district, Tamalang FR (hololectotype K).
Low emergent tree, to 45 m tall, to 1.9 m diameter; buttresses low, rounded. Bark deeply
cracked, flaky. Inflorescence, parts of petals exposed in bud, ovary, and nut persistently
cream puberulent; sepals becoming sparsely so in fruit; twig caducously so. Twigs c. 3 mm
diameter apically, terete, more or less rugulose, stout. Leaf buds small, ovoid, obtuse.
Stipules fugaceous. Leaves coriaceous, lustrous above, drying greyish green; blade ovate,
275
8–15 × 5–9 cm, base broadly cuneate to obtuse, subequal, shortly decurrent, apex shortly
broadly acuminate; midrib stout, distinctly elevated on both surfaces; lateral veins 7–9
pairs, slender, arched; intercostal venation subreticulate, elevated below, evident above;
petiole rather stout, 1.7–2.2 cm long. Inflorescences terminal or axillary; rachis to 14 cm
long, doubly or more branched, manyflowered; bracteoles not seen. Flowers: buds c. 3 × 2
mm; petals cream; stamens 15, connectival appendage short, equal in length with anther,
scarious apically; ovary ovoid, without stylopodium, style short. Fruits subsessile; calyx
lobes unequal, 3 longer lobes to 6 × 1 cm, tapering to 3 mm above the saccate base, 2
shorter ones to 4.3 × 0.5 cm, otherwise similar. Nuts narrowly ovoid, to 2 × 0.8 cm.
Vernacular name. Sabah—seraya kuning (preferred name).
Distribution. Endemic in NE Sabah; known from Kota Kinabalu, Kuala Penyu, Kudat,
Labuan, Papar, and Pitas districts (e.g., SAN 5905, SAN 15494, SAN 15552, SAN 30903, and
SAN 37701).
Ecology. Locally common in mixed dipterocarp forests on dry hills near the sea.
Endangered.
61. Shorea kunstleri King

(H.H. Kunstler, 1837–1887, King’s plant collector in Perak, Peninsular Malaysia)
J. As. Soc. Beng. 62, 2 (1893) 116; Browne op. cit. 154; Ashton op. cit. (1964) 192, op. cit. (1968)
126; PROSEA op. cit. 430; Coode et al. (eds.) op. cit. 78; Newman et al. op. cit. (1996) 155.
Lectotype (designated here): King’s Collector (Kunstler) 3674, Peninsular Malaysia, Perak, Larut
(hololectotype K).
Large emergent tree, to 55 m tall, to 1.9 m diameter; crown large, cauliflowershaped,

somewhat diffuse, with a few large ascending branches; bole tall, cylindrical; buttresses to
1.5 m tall, spreading, stout. Bark becoming pale mauve to greybrown, deeply narrowly v
section fissured and flaky, with yellow dammar smears; inner bark orange brown. Sapwood
hard; heartwood dark reddish brown. Young parts shortly pale buff puberulent; more or less
caducous on twig, petiole and leaf blade; more dense and persistent on inflorescence, leaf
bud, ovary, nut, stipules and bracteole outside (sparse within). Twigs c. 2 mm diameter
apically, terete, muchbranched. Leaf buds falcate, 5–7 × 2–2.5 mm. Stipules narrowly
lanceolate, to 12 × 4 mm, fugaceous. Leaves coriaceous, drying pale greyish tawny; blade
broadly ovate and frequently twisted to one side, 8–12 × 4.5–7 cm, base broadly cuneate to
obtuse, apex with tapering acumen to 1 cm long; midrib evident but shallowly furrowed
above, broadly elevated below; lateral veins 6–8 pairs, slender, more or less raised but not
prominent below, arched, with or without minute axillary porelike domatia; intercostal
venation slender, scalariform; petiole 2–3.5 cm long, geniculate. Inflorescences terminal or
rarely axillary; rachis slender, slightly compressed on drying, to 15 cm long, branchlets lax,
bearing to 9 flowers; bracteoles elliptic, obtuse, to 5 × 3.5 mm, fugaceous. Flowers: buds to
9 × 3 mm; corolla pale yellow, pink at base; stamens 15, connectival appendage more than
276
2x the length of anther; ovary and stylopodium pyriform, style somewhat shorter than ovary,
glabrous. Fruits: calyx glabrescent, lobes unequal, 3 longer lobes to 8.5 × 1.8 cm, tapering
to c. 5 mm above the saccate base, 2 shorter ones linearlobed, to 4 × 0.4 cm, similar at
base. Nuts ovoid, to 2 × 1.5 cm, with c. 3 mm long conical style remnant.
Vernacular names. Sabah—seraya sirap (preferred name). Sarawak—selangan merah

(preferred name).
Distribution. N Sumatra, Peninsular Malaysia and Borneo. In Sabah recorded from Lahad
Datu, Sipitang, and Tawau districts (e.g., SAN 16822, SAN 21503, SAN 24282, SAN 25468,
and SAN A 294) and in Sarawak from Bintulu, Kuching, Lawas, Miri, and Samarahan
(e.g., BRUN 346, FMS 48490 and S 1905) and in scattered localities throughout Kalimantan
(e.g., bb. 11080, bb. 19164 and bb. 26855).
Ecology. Locally frequent in mixed dipterocarp forest on yellow sandy and sandy clay,
usually deep, soils over sandstone; on ultrabasic substrates in E Sabah; occasionally on high
ridges at altitudes to 800 m. Occurring in Kubah NP and common in Lambir NP;
endangered outside parks system.
62. Shorea ladiana P.S.Ashton Fig. 25.

(Ladi Bikas, plant collector in Brunei, 1957–1962)
(sect. Shorea, subsect. Barbata, selangan batu)
op. cit. (1980) 120; Coode et al. (eds.) op. cit. 78; Newman et al. op. cit. (1998) 186. Type: Ghazalli S
11076, Borneo, Sarawak, Kuching district, Semengoh FR (holotype K; isotype KEP).
Main canopy tree, to 40 m tall, to 1.5 m diameter; bole cylindrical; crown dense,
hemispherical; buttresses to 1 m tall, thin. Bark yellowish brown, becoming irregularly
oblong flaky, sometimes shaggy. Young twig and petiole cinereous; leaf bud, parts of
perianth exposed in bud, ovary, nut, inflorescence, and bracteole greybuff pubescent; parts
otherwise glabrous. Twigs terete, muchbranched, 1.5–2 mm diameter apically. Leaf buds
small, conical, c. 1 × 0.5 mm. Stipules unknown. Leaves coriaceous, shiny; blade ovate,
10–14 × 4.5–7.5 cm, base obtuse or broadly cuneate, margin narrowly revolute, apex with
tapering acumen to 1.5 cm long; midrib evident and raised within a shallow furrow above,
stout, raised but not prominent below as also the lateral veins; lateral veins 5–6 pairs, well
spaced, arched; intercostal venation indistinct, densely scalariform; petiole 1–2.2 cm long.
Inflorescences terminal or axillary; rachis somewhat compressed, straight, lax, with short
branchlets bearing to 9 flowers; bracteoles ovoiddeltoid, acute, to 4.5 mm long, fugaceous.
Flowers: buds globose, to 1.5 mm diameter; petals cream; stamens 30–35, anther locules
shortly barbate apically, connectival appendage longer than anther, densely barbate; ovary
and stylopodium ovoid, style short, glabrous. Fruits: pedicels to 3 mm long; calyx lobes
subequal, shorter than ripe nut, broadly ovate, acute, thickened, saccate, to 0.8 × 0.8 cm.
Nuts ovoidglobose, to 1.7 × 1.4 cm, sharply acute at the c. 1 mm long style remnant.
Vernacular name. Sarawak—selangan batu kilat (preferred name).
277
Distribution. Endemic in NW Borneo; known in W Sarawak from Bintulu, Kuching and

Miri districts (e.g., S 15255, S 27171, S 29309, and S 32416) and in Brunei from Belait
district (e.g., BRUN 2622 and BRUN 5543).
Ecology. Locally frequent in mixed dipterocarp forest on deep yellow sands, at altitudes to
300 m. Recorded from Kubah NP; elsewhere endangered.
63. Shorea laevis Ridl.

(Latin, laevis = smooth, hairless; the leafy shoots)
FMP 1 (1922) 232; Browne op. cit. 169; Ashton op. cit. (1964) 139, op. cit. (1968) 73, op. cit. (1978)
120; PROSEA op. cit. 431; Kessler & Sidiyasa op. cit. 104; Coode et al. (eds.) op. cit. 78; Newman et
al. op. cit. (1998) 187. Lectotype (designated here): Sahak FMS 1905, Peninsular Malaysia, Negeri
Sembilan, Kuala Pilah, Serting FR (hololectotype K). Synonyms: Hopea laevifolia Parijs in Fedde,
Rep. 33 (1933) 244; Shorea laevifolia (Parijs) Endert, Tectona 28 (1935) 292; S. rogersiana Raizada
& Smitinand, Thai For. Bull. Bot. 1 (1954) 7.
Huge emergent tree, to 70 m tall, to 2.4 m diameter; bole stout, often twisted; crown vast
irregular diffuse cauliflowershaped, pale from below; buttresses to 6 × 4 m, stout. Bark
pale orange to greybrown, becoming vertically cracked and irregularly thinly flaky, with
pale yellow dammar smears. Inflorescence, bracteole, parts of perianth exposed in bud,
ovary, and nut persistently pale cream puberulent; fruit calyx sparsely so; vegetative parts
epilose; leaf undersurface, petiole, and young twig cream lepidote (mature trees). Twigs
slender, terete, muchbranched, c. 1 mm diameter apically. Leaf buds narrowly ovoid, 3.5 ×
1.5 mm. Stipules narrowly lanceolate, to 8 × 2 mm, fugaceous. Leaves thinly coriaceous,
drying greyish brown with cream undersurface and black venation (mature trees); blade
narrowly ovatelanceolate, falcate, 6.5–10 × 2.5–4 cm, base subequal, broadly cuneate,
apex with slender acumen to 2 cm long; midrib and venation hardly raised on either
surface, midrib evident above; lateral veins slender, dense, 11–14 pairs; intercostal
venation densely scalariform, slender; petiole 1–1.5 cm long, slender, geniculate.
Inflorescences terminal or axillary; rachis slender, terete, to 12 cm long, branchlets bearing
to 9 flowers; bracteoles linear, to 2 mm long, fugaceous. Flowers: buds globose, c. 2 mm
diameter; petals cream; stamens c. 50, filaments sparsely barbate distally, anther locules
barbate at base and apex, connectival appendage small, barbate; ovary and stylopodium
small, conical, style short, glabrous. Fruits: calyx lobes unequal, 3 longer lobes
chartaceous, to 6.5 × 1 cm, tapering to c. 4 mm above the saccate base, 2 shorter ones to 4
× 0.5 cm, otherwise similar. Nuts ovoid, to 1.5 × 0.9 cm, with to 4 mm long tapering style
remnant.
Vernacular names. Sabah—selangan batu kumus (preferred name). Sarawak—kumus

(preferred name), mikai (Iban).
Distribution. Peninsular Myanmar, Peninsular Thailand (Pattani), N Sumatra, Peninsular

Malaysia, and Borneo. In Borneo widespread. In Sabah known from Beaufort, Keningau,
Kinabatangan, Kota Merudu, Kudat, Labuk Sugut, Lahad Datu, Ranau, Sipitang, Tawau,
and Tenom districts (e.g., SAN 15207, SAN 17037, SAN 26086, SAN 97661, and SAN
99560) and in Sarawak from Belaga, Bintulu, Kapit, Lawas, Limbang, Miri, and Sri Aman
278
Fig. 25. Shorea ladiana. A, flowering leafy twig; B, longitudinal section of flower bud; C,
open flower; D, open flower with sepals and two petals removed; E, adaxial view of outer
sepal; F, adaxial view of inner sepal; G, adaxial view of petal with stamens; H, abaxial view
of petal; I, adaxial view of stamens; J, abaxial view of stamens; K, fruit; L, longitudinal
section of fruit. (A from S 583, B–J from S 26229, K–L from S 15255.)
279
(e.g., BRUN 409 and BRUN 3190) and E Kalimantan (e.g., bb. 23427, bb. 34270 and bb.
34453).
Ecology. One of the commonest emergent trees in mixed and upper dipterocarp forests
along shale ridges, at 200–1000 m altitude, usually on leached skeletal clay and sandy clay
soils. Occurring in Mulu NP; elsewhere vulnerable owing to logging.
64. Shorea lamellata Foxw.

(Latin, lamellatus = arranged in layers; the inner bark)
Malay. For. Rec. 10 (1932) 278; Symington op. cit. (1943) 39; Slooten op. cit. (1949) 236; Ashton op.
cit. (1968) 94, p.p., op. cit. (1982) 493; Anderson op. cit. (1980) 123, p.p.; PROSEA op. cit. 412;
Kessler & Sidiyasa op. cit. 104; Newman et al. op. cit. (1996) 156. Type: Arnot FMS 28082,
Peninsular Malaysia, Perak, Keledang Saiong FR (holotype KEP).
Large emergent tree, to 50 m tall, to 1.75 m diameter; bole tall, cylindrical; crown dense,
hemispherical; buttresses to 3 m tall, stout. Bark greyish to purplish brown, becoming
irregularsection fissured and coarsely flaky; inner bark laminated pale and dark yellow.
Twig, leaf bud, stipule and bracteole outside, inflorescence, and parts of calyx exposed in
bud persistently tawny brown scabridpubescent; fruit calyx, stipule and bracteole within,
petiole, and leaf venation below sparsely so; petals outside, ovary, and nut evenly pale
brown pubescent. Twigs terete, 2–3 mm diameter apically. Leaf buds ovoidfalcate, acute,
to 7 × 4 mm. Stipules oblongobtuse, to 20 × 6 mm. Leaves thinly coriaceous, drying
greyish, frequently bullate in between lateral veins; blade ellipticoblong to ovate or
occasionally obovate, (6.5–)10–15 × (3.5–)4–8 cm, base obtuse to shallowly cordate, apex
with short, abrupt acumen to 0.7 cm long; midrib slender but prominent below, obscure,
sunken above; lateral veins 19–25 pairs, slender but prominent below; intercostal venation
densely scalariform, evident and somewhat elevated below; petiole slender, 1.6–2.2 cm
long. Inflorescences terminal or axillary; rachis slender, lax, to 14 cm long, singly
branched, branchlets bearing to 3 flowers; bracteoles elliptic, acute, to 5 × 3 mm, tardily
caducous. Flowers: buds ellipsoid, to 10 × 5 mm; petals white; stamens 15, anthers oblong,
twice as long as broad, connectival appendage 2½–3x the length of anther, scarious
towards apex; ovary small, ovoid, tapering into a slender pubescent stylopodium twice its
length and glabrous columnar style 3x its length. Fruits: pedicels c. 2 mm long, fruit base
obtuse; calyx lobes unequal, 3 longer lobes to 18 × 1.5 cm, tapering to c. 7 mm above the
saccate base, 2 shorter ones loratelobed, to 7 × 0.5 cm, similar at base. Nuts ovoid, to 1.4
× 1 cm, prominently apiculate.
Vernacular name. Sarawak—meranti lapis (preferred name).
Distribution. Sumatra (Lingga and Singkep), Peninsular Malaysia and Borneo. Absent from
Sabah. In Sarawak known from Bintulu, Kuching, Mukah, and Tatau districts (e.g., S 902, S
15126, S 15828, and S 18082). Also occurring in E and SE Kalimantan (e.g., bb. 34264,
Kostermans 5813, Kostermans 7695, and Kostermans 8723).
280
Ecology. Rare, scattered in mixed dipterocarp forest on low hills on sandy clay soils, on
sedimentary substrates and also the Arip rhyolite, at altitudes to 500 m. Recorded from
Kubah NP; elsewhere endangered owing to land conversion.
Notes. Confused by Ashton (op. cit. (1964) 164, op. cit. (1968) 94), Meijer and Wood (op. cit.
56), Burgess (op. cit. 159) and Anderson (op. cit. (1980) 123) with S. confusa and S.
virescens. The typically oblongovate leaf blade, terete twig and scabrid indumentum serve
to distinguish this species.
65. Shorea laxa Slooten

(Latin, laxus = loose; possibly referring to the inflorescence)
Reinwardtia 3 (1956) 345; Ashton op. cit. (1964) 155, op. cit. (1968) 86, op. cit. (1982) 474; Meijer &
al. (eds.) op. cit. 78; Newman et al. op. cit. (1996) 157. Type: Flemmich FMS 48191, Borneo, Brunei,
Bt. Puan (holotype KEP).
Low emergent tree, to 50 m tall, to 1.6 m diameter; bole tall, cylindrical; crown dense,
hemispherical; buttresses to 2.5 m tall, stout. Bark pale yellowish brown, becoming
shallowly densely vertically cracked and thinly oblong flaky. Inflorescence, bracteole
outside, parts of perianth exposed in bud, ovary, nut, leaf bud, stipule outside, twig, petiole,
and the basal half of the midrib below persistently sparsely creambuff puberulent; calyx
eventually glabrescent in fruit. Twigs terete, 2 mm diameter apically. Leaf buds globose or
ovoid, subacute, c. 3 × 2 mm. Stipules lanceolate, to 5 × 2 mm, fugaceous. Leaves thinly
coriaceous, drying warm brown, slightly shiny; blade ovate to elliptic, 7–14 × 4–9 cm, base
obtuse to broadly cuneate (peltate in juveniles), margin narrowly revolute, apex with
slender acumen to 1.5 cm long; midrib evident, flat above, prominent below as also the
veins; lateral veins 8–10 pairs, arched and somewhat following the margin; intercostal
venation scalariform, wellspaced; petiole 1.5–2.2 cm long. Inflorescences terminal or
axillary; rachis terete, slender, lax, to 18 cm long, branchlets bearing to 7 distichous flowers;
bracteoles minute, to 1.5 mm long, fugaceous. Flowers: buds to 8 × 2 mm; petals cream;
stamens 15, connectival appendage 2x as long as anther, ciliate towards apex; ovary and
stylopodium ovoidconical, style short, pubescent at base. Fruits: calyx lobes subequal,
broadly ovate, incrassate, saccate, tuberculate, c. 1 × 1 cm, the obtuse thin apices recurved.
Nuts obovoid, acute, to 3.5 × 2.5 cm, on germination splitting to reveal brilliant red
cotyledons.
Vernacular names. Sabah—seraya kuning keladi (preferred name). Sarawak—lun timbul

(preferred name).
Distribution. Endemic in the northern parts of Borneo. Known in Sabah from Labuk Sugut
and Sandakan districts (e.g., SAN 24299 and SAN 39305) and in Sarawak from Bintulu, Miri
and Tatau districts (e.g., S 15141, S 46410 and S 46598). Also occurring in Brunei (e.g.,
BRUN 3010 and BRUN 3061).
Ecology. Very local on account of its local habitat, but there often common in mixed
dipterocarp forest on deep yellow sandy soils of coastal hills comprised of soft neogene
281
sandstone in Sarawak, ultrabasic substrates in Sabah, at altitudes below 400 m. Common in

Lambir NP; elsewhere vulnerable.
Notes. Closely related to S. peltata, see there.
66. Shorea leprosula Miq.

(Latin, leprosulus = leprous; the appearance of the domatia)
Fl. Ned. Ind., Suppl. (1861) 487; Beccari, Nelle For. Born. (1902) 570; Merrill op. cit. (1921) 405, op.
cit. (1929) 203; Keith op. cit. 12; Masamune op. cit. 494; Slooten op. cit. (1949) 262; Browne op. cit.
Burgess op. cit. 154, 182; Anderson op. cit. (1980) 126; PROSEA op. cit. 396; Kessler & Sidiyasa op.
cit. 105; Coode et al. (eds.) op. cit. 79; Newman et al. op. cit. (1996) 158. Lectotype (designated
here): Teijsmann HB 1122, Sumatra (hololectotype L). Synonyms: Hopea ?maranti Miq., op. cit. 489;
Shorea maranti (Miq.) Burck op. cit. 217.
Large late successional tree, to 60 m tall, to 1.5 m diameter; crown pale pinkish from below,
diffuse, cauliflowershaped; bole tall, cylindrical; buttresses to 2 m tall, stout. Bark pale
fawnbrown, deeply widely vsection fissured, the intervening ridges only becoming
crumbly flaky in old trees; inner bark reddish brown. Sapwood light; heartwood pale
reddish brown. Twig, inflorescence, leaf bud, parts of flower exposed in bud, stipule,
bracteole, petiole, and leaf venation below persistently densely pinkish buffpuberulent;
blade undersurface densely pinkish brown sericeous (mature trees only). Twigs terete,
ridged at first, muchbranched, c. 1.5 mm diameter apically. Leaf buds broadly ovoid,
subacute, compressed, 3–5 × 2–3 mm. Stipules oblong to broadly hastate, obtuse, to 10 ×
3.5 mm, fugaceous. Leaves thinly coriaceous, drying pale pinkish brown above, pinkbuff
below (mature trees); blade ovateoblong, 8–14 × 3.5–5.5 cm, base obtuse or broadly
acuminate, apex with short acumen to 0.8 cm long; midrib obscure, sunken above,
prominent below, beset with lines of scalelike domatia in young trees; lateral veins 12–15
pairs, slender but prominent below, arched near margin; intercostal venation slender,
densely scalariform; petiole 1–1.5 cm long. Inflorescences terminal or axillary; rachis
terete, lax, slender, to 14 cm long, generally singly branched, branchlets bearing to 12
flowers; bracteoles elliptic, obtuse, to 3 × 2 mm, fugaceous. Flowers: buds to 6 × 3 mm;
petals pale yellow; stamens 15, connectival appendage short, becoming reflexed; ovary and
stylopodium ovoid, glabrous, style twice their length. Fruits: calyx more or less
glabrescent; calyx lobes unequal, aliform, 3 longer lobes to 10 × 2 cm, tapering to c. 5 mm
above the saccate base, 2 shorter ones to 5.5 × 0.3 cm, otherwise similar. Nuts ovoid, to 1
× 1.3 cm, with c. 2 mm long tapering style remnant.
Vernacular names. Sabah—seraya tembaga (preferred name). Sarawak—meranti tembaga

(preferred name; Malay).
Distribution. Extreme SE Peninsular Thailand (Pattani), Sumatra, Peninsular Malaysia, and

Borneo (throughout the island in suitable habitats). In Sabah known from most districts
(e.g., SAN 15509, SAN 18002, SAN 29715, SAN 69199, and SAN 94912) and in Sarawak
from Bintulu, Kapit, Kuching, Limbang, Lundu, Marudi, Miri, and Tatau districts (e.g., S
802, FMS 30484 and FMS 37463) and Kalimantan (e.g., bb. 34268, Kostermans 12650 and
Meijer 2273).
282
Ecology. Absent from the widespread sandy soils of Sabah and Sarawak, but frequent,
locally quite common in mixed dipterocarp forest on clay soils on shales and clays, also acid
and intermediate igneous rocks and periodically flooded alluvium, at altitudes to 600 m. A
light demander, competitive in early to late succession and flowering more frequently than
other species of sect. Mutica. Occurring in G. Gading, Lambir and Mulu NPs in Sarawak,
and Danum Valley Conservation Area and Sepilok FR in Sabah; not vulnerable.
67. Shorea longiflora (Brandis) Symington

(Latin, longus = long, flos = flower)
Gard. Bull. S. S. 9 (1938) 330; Slooten op. cit. (1956) 318; Browne op. cit. 163; Anderson op. cit.
(1963) 158, op. cit. (1980) 122; Ashton op. cit. (1964) 156, op. cit. (1968) 86, op. cit. (1982) 473;
PROSEA op. cit. 419; Coode et al. (eds.) op. cit. 79; Newman et al. op. cit. (1996) 160. Basionym:
Hopea longiflora Brandis op. cit. 63. Type: Haviland 2120, Borneo, Sarawak, probably near Kuching
(holotype K). Synonym: Balanaocarpus longiflorus (Brandis) Foxw. ex Symington op. cit. (1934) 29.
Understory or low main canopy tree, to 35 m tall, to 50 cm diameter; bole frequently

misshapen; crown loose, oblong, with pendent branches and leaves; buttresses low, stout,
occasionally with a few stilt roots. Bark fawn and dark grey, becoming patchily cracked
and thinly irregularly flaky. Young twig, leaf bud, inflorescence, parts of perianth exposed
in bud, and stipule fulvous powdery puberulent, more or less caducous. Twigs terete, stout,
rugulose, becoming papery scaly, c. 2.5 mm diameter apically. Leaf buds globose or
ellipsoid, slightly compressed, c. 3 × 2 mm. Stipules oblongelliptic, to 10 × 6 mm, obtuse,
concave, caducous. Leaves thickly coriaceous; blade narrowly ovatelanceolate, 10–24 ×
4–6 cm, base obtuse, margin prominently revolute, apex with slender acumen to 2 cm long;
midrib evident but furrowed above, prominent below; lateral veins 12–15 pairs, prominent
below, distant, arched and ascending; intercostal venation distantly subscalariform; petiole
1–1.2 cm long. Inflorescences terminal or axillary; rachis terete, to 11 cm long, unbranched
or singly branched, branchlets bearing few flowers; bracteoles minute, fugaceous. Flowers:
buds to 8 × 3.5 mm; petals brownish purple to dark yellow; stamens 15, anthers becoming
reflexed at anthesis, connectival appendage of same length as anther, ciliate towards apex;
ovary and stylopodium attenuateovoid, glabrous, tapering into glabrous style. Fruits
subsessile; calyx lobes subequal, broadly deltoid to ovate, rather thin, spreading not
clasping the nut, c. 0.7 × 0.7 cm. Nuts ellipsoid to ovoid or obovoid, to 4.5 × 1.7 cm,
glabrous, shiny, with short apical style remnant.
Vernacular name. Sarawak—lun paya (preferred name).
Distribution. Endemic in Borneo. Known in Sarawak from Kapit, Kuching, Marudi, Miri,
Samarahan, Simunjan, and Tatau districts (e.g., Richards 2441, S 6518, S 14400, S 19425,
and S 34356). Also occurring in Brunei (e.g., BRUN 109, FMS 35658 and SAN 17535) and
C and E Kalimantan (e.g., Suzuki K 11847).
Ecology. Scattered, often in local patches in mixed dipterocarp forest on humic soils, often
poorly drained but sometimes welldrained, on deep yellow sands; also in mixed peat
swamp and karapa forest at 400–1000 m altitude. Occurring in Lambir NP; elsewhere
vulnerable owing to its accessible habitat.
283
68. Shorea longisperma Roxb.

(Latin, longus = long, spermus = seed; the elongated nut)
Fl. Ind., ed. Carey 2 (1832) 618; Ashton op. cit. (1978) 43, op. cit. (1982) 481; PROSEA op. cit. 420;
Coode et al. (eds.) op. cit. 79; Newman et al. op. cit. (1996) 161. Type: Roxburgh s.n., Peninsular
Malaysia, Penang (‘Prince of Wales Is.’) (holotype CAL). Synonyms: Parashorea longisperma
(Roxb.) Kurz op. cit. 66; Shorea resinanigra Foxw., op. cit. (1932) 205, Ashton op. cit. (1968) 89,
Anderson op. cit. (1980) 123.
Huge tree, to 70 m tall, to 2.9 m diameter; bole tall, cylindrical; buttresses to 4 m tall and
out, stout; crown immense, diffuse, cauliflowershaped, pale from below. Bark tawny
brown, deeply cracked and thinly oblong flaky; dammar blackish oxidizing to cream, as
occasional small coxcombs. Leaf bud, stipule, inflorescence, bracteole, and parts of
perianth exposed in bud persistently fulvous scabridpuberulent; twig and fruit calyx
caducously so; ovary and nut evenly buffpuberulent; leaf undersurface buff lepidote. Twigs
slender, terete, 1–2 mm diameter apically. Leaf buds ovoid, acute, to 3 × 2 mm. Stipules
ovate to lorate, acute, to 5 × 2 mm, caducous. Leaves somewhat chartaceous, pale mauve
when opening, drying tawnybrown above, pale brownish grey below with the veins
distinctly darker; blade elliptic to ovate, 7–12 × 2.3–6 cm, base cuneate to obtuse, apex
with slender acumen to 1.5 cm long; midrib evident, flat to shallowly furrowed above,
prominent below; lateral veins 10–13 pairs, slender but distinctly raised below; intercostal
venation subscalariform; petiole 1–1.5 cm long. Inflorescences terminal or axillary; rachis
terete, slender, to 7 cm long, singly branched, branchlets bearing to 4 flowers; bracteoles
elliptic, to 3 × 2 mm. Flowers: buds to 9 × 3 mm; petals pale yellow; stamens 15,
connectival appendage c. 2x as long as anther; ovary and stylopodium pyriform, style as
long as both, columnar, glabrous. Fruits sessile; calyx lobes unequal, 3 longer lobes to 6 ×
4 cm, tapering to c. 3 mm above the narrowly saccate tuberculate base, 2 shorter ones
linearlobed, to 7 × 0.4 cm, similar at base. Nuts ellipsoid, to 2.3 × 1.2 cm, acute.
Vernacular name. Sarawak—lun meranti (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sarawak known from Belaga,
Kapit, Lawas, Marudi, Miri, and Tatau districts (e.g., S 16738, S 18279, S 19276, and S
69118). Also occurring in Brunei (BRUN 2539) and C and E Kalimantan (e.g., Arifin et al.
Berau 727).
Ecology. Rare, as scattered individuals in mixed dipterocarp forest on wellstructured deep

clay soils over shale, dacite and basalt, on low coastal hills and uplands, at altitudes to 1100
m. Vulnerable, becoming endangered owing to logging at higher altitudes.
69. Shorea lunduensis P.S.Ashton

(of Lundu, Sarawak)
Newman op. cit. (1998) 190. Type: Anderson et al. S 15396, Borneo, Sarawak, Lundu district, G.
Lundu (holotype K; isotypes KEP, L).
284
Large emergent tree, to 60 m tall, to 1.3 m diameter; bole straight, cylindrical; crown
irregularly hemispherical, with a few large ascending branches; buttresses to 4 m tall,
relatively stout. Bark greyish brown, irregularly flaky. Inflorescence, parts of perianth
exposed in bud, ovary, and nut densely buffpuberulent, persistent except on calyx; twigs
and leaf bud shortly evenly fugaceous buffpubescent; parts otherwise glabrous. Twigs at
first compressed, somewhat shiny, c. 3 × 2 mm apically; stipule scars pale, conspicuous,
ascending, c. 3 mm long. Leaf buds ovoid, acute, to 5 × 3 mm. Stipules unknown. Leaves
coriaceous, large, drying yellowish brown, satiny above; blade broadly ovate to elliptic, 14–
24 × 6–15 cm, base broadly cuneate to subcordate, apex with tapering acumen to 1 cm
long; midrib broad but hardly raised above, prominently so below; lateral veins 11–15
pairs, prominent below, ascending; intercostal venation densely scalariform, sinuate,
evident but hardly raised; petiole 2–3.5 cm long, geniculate. Inflorescences terminal or
axillary; rachis subterete, to 12 cm long, branchlets bearing to 5 flowers. Flowers: buds to
10 × 3 mm; corolla cream; stamens 47–52, filaments and anthers glabrous, connectival
appendage exceeding the length of anther, densely setose; ovary and stylopodium pyriform,
style glabrous, columnar. Fruits (mature fruit unknown): calyx lobes unequal. Nuts ovoid,
shortly apiculate.
Distribution. Endemic in W Borneo. In Sarawak, recorded from Bau, Kuching and Lundu
districts (e.g., S 59, S 7984, S 10172, S 15502, and S 25296) and in Kalimantan from
Sambas district (e.g., Suzuki K 9760).
Ecology. Very local, there frequent in mixed dipterocarp forest on clay soils, on
granodiorite, andesite plugs and a porphyry dike in limestone, at altitudes to 600 m.
Endangered by forest conversion.
70. Shorea macrantha Brandis

(Greek, makro = large, anthos = flower)
J. Linn. Soc. Bot. 31 (1895); Merrill op. cit. (1921) 405; Symington op. cit. (1933) 131, op. cit. (1943)
77; Masamune op. cit. 495; Browne op. cit. 148; Anderson op. cit. (1963) 110, op. cit. (1980) 126;
Ashton op. cit. (1968) 110, op. cit. (1982) 536; PROSEA op. cit. 397; Newman et al. op. cit. (1996)
162. Type: Haviland 2119, Borneo, Sarawak, Kuching (holotype K; isotype L).
Small canopy tree, to 40 m tall, to 60 cm diameter; bole moderately straight; crown with
pendent branches, twigs and leaves; buttresses short, stout. Bark dull purplish grey,
irregularly fissured; inner bark yellowish brown, pale at cambium; heartwood pale brown,
hard. Twig, leaf bud, stipule outside, inflorescence, parts of perianth exposed in bud,
petiole, and venation below densely persistently yellowish brown scabridpubescent; stipule
inside, midrib above, ovary, and nut evenly so; lateral veins above and fruit calyx sparsely
evenly so. Twigs terete, stout, 2–4 mm diameter apically; stipule scars short. Leaf buds
ovoid, subacute, 6 × 3 mm. Stipules lanceolate, ribbed, subacute, to 16 × 5 mm,
subpersistent. Leaves coriaceous, drying mauvebrown above, rufous below with yellowish
brown tomentum; blade bullate between veins, narrowly ovate, 6–17 × 2.5–8 cm, base
unequal, cordate, apex with slender acumen to 1.5 cm long; midrib, lateral and intercostal
venation more or less obscure, sunken above, prominent below; lateral veins 13–17 pairs,
arched; intercostal venation laxly scalariform; petiole very short, 0.5–0.6 cm long.
285
short, the flowers congested. Flowers: buds to 14 × 4 mm; petals dark red within, paler
outside; stamens 15, connectival appendage as long as anther; ovary ovoid, without
stylopodium, style glabrous, about twice as long as ovary. Fruits subsessile; calyx lobes
subequal, ovate, acute, saccate towards base, to 2.5 × 2 cm. Nuts ellipsoid, to 5.5 × 2.5 cm,
acute.
Vernacular name. Sarawak—engkabang bungkus (preferred name).
Distribution. Sumatra, Peninsular Malaysia and NW Borneo. In Borneo known only from
Sarawak and recorded from Kuching, Serian, Sibu, and Sri Aman districts (e.g., S 9781, S
13353 and S 35965).
Ecology. In small populations in the inland margins of the mixed peat swamp forest, and on
poorly drained podsols on terraces in kerangas. Endangered.
Uses. Collected as a lesser valued illipe nut, for the cocoa butter content.
71. Shorea macrobalanos P.S.Ashton

(Greek, makro = large, balanos = acorn; the huge nut)
Kessler & Sidiyasa op. cit. 105; Newman et al. op. cit. (1996) 163. Type: Smythies S 13192, Borneo,
Sarawak, Lundu district, G. Berumput (holotype K; isotypes KEP, L).
Emergent tree, to 45 m tall, to 1 m diameter; crown dense, dark, hemispherical; bole tall,
cylindrical; buttresses to 3 m tall, stout. Bark tawnybrown, irregularly flaky. Leaf bud and
stipule persistently buffpubescent; flower calyx sparsely so; parts otherwise glabrous.
Twigs terete, becoming prominently verrucose, stout, c. 4 mm diameter apically; stipule
scars obscure, ascending, c. 3 mm long. Leaf buds ovoid, acute, to 2 × 2 mm. Stipules
unknown. Leaves thinly coriaceous, drying greyish green above, tawny below with the
veins, midrib and petiole darker to blackish; blade oblong, 19–37 × 9–15 cm, base cordate,
margin revolute, apex obtuse or shortly broadly acuminate; midrib flat above, prominent
below; lateral veins 12–16 pairs, prominent below; intercostal venation slender, remotely
subreticulate; petiole stout, 1.8–3.8 cm long. Inflorescences terminal or axillary; rachis
terete or somewhat compressed, to 32 cm long. Flowers: buds large, to 9 × 3.5 mm;
stamens 10, connectival appendage as long as anther; ovary narrowly ovoid, tapering,
without stylopodium, style glabrous, short. Fruits sessile; calyx lobes subequal, ovate,
acute, subrotate, not appressed to the nut, to 0.8 × 0.8 cm, mounted on a to 8 mm deep
receptacle. Nuts oblong, to 5 × 2.5 cm, shortly apiculate.
Vernacular name. Sarawak—engkabang melapi (Iban).
Distribution. Endemic in Borneo; known in Sarawak from Kapit and Lundu districts (e.g.,
S 13712, S 19611, S 34481, and S 37292). Also occurring in NW and E Kalimantan (e.g.,
Suzuki K 9762).
286
Ecology. Locally frequent in forest on clay soils derived from shale and intermediate, and
basic igneous rocks; especially on low hills and moist valleys but also occurs at altitudes to
900 m. Conservation status uncertain; likely endangered by logging.
Notes. For differences from S. longiflora, see there.
72. Shorea macrophylla (de Vriese) P.S.Ashton Plates 4B–C.

(Greek, makro = large, phullon = leaf)
Wood op. cit. 156; Anderson op. cit. (1980) 127; PROSEA op. cit. 397; Coode et al. (eds.) op. cit. 79;
Newman et al. op. cit. (1996) 163. Basionym: Hopea macrophylla de Vriese, Minyak Tenkawang
(1861) 28. Lectotype (designated here): de Vriese s.n. (= RHL Sheet No. 902146589), ‘Borneo’
(hololectotype L). Synonyms: Shorea gysbertsiana Burck op. cit. (1886) 15, Merrill op. cit. (1921)
405, Browne op. cit. 139, Meijer & Wood op. cit. 108, Burgess op. cit. 182; S. bakeriana F.Heim op.
cit. (1891) 974, Merrill op. cit. (1921) 404; Pachychlamys gysbertsiana (Burck) Ridl., op. cit. (1922)
233 (sphalm. ghysbertsiana).
Stocky low emergent or main canopy tree, to 45 m tall, to 1.5 m diameter; crown dense,
oblong to hemispherical, branch endings pendent; bole usually not tall, often misshapen;
buttresses to 2 m tall, stout. Bark greenish grey to pale brown, at first smooth, hoop
marked, becoming thinly irregularly flaky, leaving scrollmarked new surfaces; inner bark
pale yellowish brown; heartwood rather light, pale brown. Young twig, leaf bud,
inflorescence, parts of perianth exposed in bud, ovary, nut, stipule, bracteole, petiole, leaf
blade below, and midrib above persistently densely evenly pale brown pubescent. Twigs
compressed, becoming smooth, 2.5–4 × 2–3 mm apically; stipule scar amplexicaul,
horizontal, c. 1 mm thick. Leaf buds hastate, narrowly obtuse, compressed, 12–18 × 4–6
mm. Stipules broadly hastate, obtuse, large, to 50 × 13 mm, not at first caducous. Leaves
thickly chartaceous, drying pale greyish tawny; blade ellipticoblong, 17–35 × 10–14 cm,
base obtuse or subcordate, apex with slender acumen to 1.5 cm long; midrib broad, flat
above, prominent below; lateral veins 11–20 pairs, prominent below, wellspaced;
intercostal venation scalariform, wellspaced, distinctly elevated below; petiole 1.5–3 cm
long. Inflorescences terminal or axillary; rachis compressed, pendent, to 17 cm long, singly
branched; bracteoles oblong, subacute, to 12 × 5 mm, not at first caducous. Flowers: buds
to 8 × 5 mm; petals pale pink; stamens 15, connectival appendage c. 2x as long as anther,
glabrous; ovary ovoid, densely pubescent in the distal half, stylopodium and style spindle
shaped, glabrous, as long as ovary. Fruits: calyx glabrescent; calyx lobes unequal, 3 longer
lobes oblong, coriaceous, to 11 × 3 cm, tapering to 15 mm above the saccate base, 2
shorter ones to 8 × 1.5 cm, similar at base. Nuts obovoid, to 6 × 4 cm, acute, large.
Vernacular names. Sabah—kawang jantung (preferred name). Sarawak—engkabang

jantong (Iban).
Distribution. Endemic in Borneo; widespread. In Sabah known from Beaufort, Keningau,

Kinabatangan, Sandakan, Sipitang, and Tawau districts (e.g., SAN 15073, SAN 15195, SAN
22864, SAN 23151, and SAN 38236) and in Sarawak from Belaga, Bintulu, Kapit, Kuching,
Lawas, Lundu, Miri, Mukah, Sarikei, and Song districts (e.g., S 554, S 1461, S 29218, S
42150, and S 77061). Also occurring in Brunei (e.g., BRUN 5280, FMS 35515 and Wong
287
WKM 1570) and Kalimantan (e.g., bb. 29058, Meijer 2218 and Wilkie 94275), north of the
equator. Widely planted.
Ecology. Locally abundant, in mixed dipterocarp forest on river banks and flood plains, also
less commonly on moist hillsides, at altitudes to 600 m. Most abundant from the Rajang
hinterland westwards. Occurring in Lambir and Mulu NPs; not vulnerable.
Uses. Besides a serviceable light hardwood, the tree produces the best and most plentiful of
the illipe nuts.
Notes. The indigenous form in E Sabah and NE Kalimantan differs in the fruit, with smaller
nut and longer narrower calyx lobes, and the leaf which is somewhat narrower, with cuneate
base. There may be hybridization with S. pinanga.
73. Shorea macroptera Dyer

(Greek, makro = large, pteron = a wing; the long auriculate fruit calyx lobes)
Fl. Brit. Ind. 1 (1874) 267; King op. cit. 113; Ridley op. cit. (1922) 225; Foxworthy op. cit. (1932)
195; Symington op. cit. (1943) 78; Ashton op. cit. (1963) 276, op. cit. (1964) 197, op. cit. (1968) 111,
op. cit. (1982) 532; Meijer & Wood op. cit. 116; Anderson op. cit. (1980) 127; PROSEA op. cit. 397;
Newman et al. op. cit. (1996) 165. Type: Maingay 1198 (= Kew Distr. No. 208), Peninsular Malaysia,
Malacca (holotype K). Synonyms: Shorea baillonii F.Heim op. cit. (1891) 973; S. sandakanensis
Symington op. cit. (1938) 343.
Main canopy to emergent tree, 40–50 m tall, 70–140 cm diameter; bole straight, cylindrical;
crown dense, hemispherical; buttresses to 2.5 m tall. Bark remaing smooth at first,
becoming shallowly vsection fissured, or patchily crumbly flaky; inner bark dark red to
pale brown at cambium; heartwood light, pinkish brown. Young twig, inflorescence, leaf
bud, parts of perianth exposed in bud, fruit calyx outside, stipule and bracteole outside
(sericeous within), and petiole sparsely tufted puberulent, glabrescent, or densely, evenly
persistent pubescent. Twig frequently somewhat compressed, becoming terete, c. 2 mm
diameter apically. Leaf buds ovoid, subacute, 4–6 × 2.5–4 mm. Stipules oblong, subacute,
to 8 × 3 mm, fugaceous. Leaves coriaceous, somewhat shiny, drying distinct pale orange
brown; blade narrowly elliptic, or narrowly ovate, or narrowly oblong, (8–)12–19(–23) ×
3.7–9.5 cm, base narrowly cuneate or obtuse, apex tapering or abruptly attenuate; midrib
obscure, shallowly sunken above, prominent below; lateral veins 10–15(–18) pairs,
prominent below, arched; intercostal venation densely scalariform, sinuate; petiole 1.5–2
cm long. Inflorescences terminal or axillary; rachis slightly compressed or terete, lax, 13–
16 cm long, singly or doubly branched, branchlets bearing to 7 distichous flowers;
bracteoles elliptic, obtuse, to 3.5 × 2.5 mm. Flowers: buds to 5 × 3 mm; petals cream with
pink base; stamens 15, connectival appendage short, becoming reflexed; ovary and
stylopodium conical, style half their length, glabrous. Fruits: calyx lobes unequal, 3 longer
lobes 12–14 × 2–3 cm, somewhat tapering to 1.2 cm wide at the incrassate concave hardly
auriculate base, 2 shorter ones loratelobed, to 6 × 0.6 cm, at base hardly auriculate to
prominently auriculate. Nuts ovoid, to 1.8 × 0.2 cm, with to 2 mm acute tapering style
remnant.
288
Vernacular names. Sabah—seraya melantai (preferred name). Sarawak—meranti melantai

(preferred name).
Ecology. In mixed dipterocarp forest on clay soils over sedimentary and igneous rocks, and
on sandy clay soils, at altitudes to 900 m. Not vulnerable.
Notes. Four subspecies, viz. subsp. baillonii, subsp. macroptera, subsp. macropterifolia, and
subsp. sandakanensis, are recognised. Whereas subsp. macroptera is confined to Peninsular
Thailand, Sumatra, Peninsular Malaysia, and Singapore, subsp. baillonii, subsp.
macropterifolia and subsp. sandakanensis are endemic in Borneo.
Key to subspecies
1. Inflorescences singly or doubly branched. Twigs, petiole and inflorescences sparsely

tuftedpuberulent, glabrescent. Two outer fruit calyx lobes hardly auriculate at base.
Leaf blade narrowly elliptic, base narrowly cuneate……………………………………..
subsp. baillonii (F.Heim) P.S.Ashton
(H.E. Baillon, 1827–1895, French botanist)
Gard. Bull. Sing. 20 (1963) 277, op. cit. (1968) 111, op. cit. (1982) 532; Brown op. cit. 137;
Anderson op. cit. (1980) 127, p.p.; PROSEA op. cit. 165; Newman et al. op. cit. (1996) 165.
Basionym: Shorea baillonii F.Heim op. cit. (1891) 973. Type: Beccari PB 2891, Borneo,
Sarawak, Mt. Matang (holotype P). Synonym: Shorea macroptera auct. non Dyer: Merrill
op. cit. (1921) 405, Masamune op. cit. 495, p.p.
Endemic in Borneo; recorded in Sarawak from Belaga, Bintulu, Kapit, Kuching,
Marudi, Miri, Simunjan, Song, and Tatau districts (e.g., S 1490, S 11090, S 15581,
S 25020, and S 43607). Abundant in mixed dipterocarp forest on clay and sandy
clay soils over sendimentary rocks, at altitudes to 600 m. Occurring in Bako,
Kubah and Lambir NPs; not vulnerable.
Inflorescences exclusively doubly branched. Twigs, petiole and inflorescences densely
persistently evenpubescent. Two outer fruit calyx lobes prominently auriculate at base.
Leaf blade narrowly ovate or narrowly oblong, base obtuse……………………………2
2. Leaf blade narrowly ovate, 8–16 × 4–6 cm, apex gradually tapering; petiole to 1.5 cm
long……………………………………………………………………………………….
subsp. macropterifolia P.S.Ashton
(Latin, with leaves resembling those of subsp. macroptera)
Gard. Bull. Sing. 20 (1963) 227, op. cit. (1964) 197, op. cit. (1968) 111, op. cit. (1982) 533;
Meijer & Wood op. cit. 116, p.p.; Burgess op. cit. 182, p.p.; Anderson op. cit. (1980) 127,
p.p.; PROSEA op. cit. 397; Coode et al. (eds.) op. cit. 79; Newman et al. op. cit. (1996)
166. Type: G.H.S. Wood SAN 16255, Borneo, Sabah, Sipitang district (holotype K; isotypes
KEP, SAN).
Endemic in Borneo. Known in Sabah from Beaufort, Kota Kinabalu,
Kinabatangan, Sipitang, and Tawau districts (e.g., SAN 16453, SAN 16598, SAN
72371, SAN 76270, and SAN 97510) and in Sarawak from Bintulu, Kapit, Lawas,
Limbang, and Miri districts (e.g., S 1514, S 10701, S 15888, S 22323, and S
66163). Also occurring in Brunei (e.g., BRUN 356, FMS 48102 and S 2151) and E
Kalimantan (e.g., Meijer 2555). Much overlapping in distribution with subsp.
baillonii but not on sandy clay soils and more frequent in the hills. Occurring in
Kinabalu, Lambir and Mulu NPs; not vulnerable.
289
Leaf blade narrowly oblong, (9–)18–23 × (4–)6.5–9.5 cm, apex abruptly tapering;
petiole to 2 cm long………………………………………………………………………
subsp. sandakanensis (Symington) P.S.Ashton
(of Sandakan, Sabah)
Gard. Bull. Sing. 20 (1963) 111, op. cit. (1982) 532; PROSEA op. cit. 397; Newman et al.
op. cit. (1996) 167. Basionym: Shorea sandakanensis Symington op. cit. (1938) 343. Type:
Agama FMS 38730, Borneo, Sabah, Sandakan district, Kabili FR (holotype KEP).
Synonym: Shorea macroptera auct. non Dyer: Slooten op. cit. (1929) 203, Masamune op.
cit. 495, p.p., Meijer & Wood op. cit. 116, p.p., Burgess op. cit. 182, p.p.
Endemic in Borneo. Known in Sabah from Beaufort, Kinabatangan, Kota
Kinabalu, Kudat, Labuk Sugut, Lahad Datu, Sandakan, and Tawau districts (e.g.,
SAN 23978, SAN 35341, SAN 38380, SAN 63791, and SAN 75971). Also occurring
in Brunei (e.g., FMS 35673) and E Kalimantan (e.g., Meijer 2043 and Meijer
2560). Occurring in Danum Valley Conservation Area and Sepilok FR; elsewhere
vulnerable owing to land conversion.
74. Shorea materialis Ridl.

(of the Materials, i.e., King’s (1893) Materials for a Flora of the Malay Peninsula)
Agr. Bull. Str. & F.M.S. 9 (1910) 183; Ashton op. cit. (1964) 140, op. cit. (1968) 74, op. cit. (1982)
456; Anderson op. cit. (1980) 120; PROSEA op. cit. 432; Coode et al. (eds.) op. cit. 79; Newman et
al. op. cit. (1998) 191. Lectotype (designated here): Ridley 15209 (Sanger Davies), Peninsular
Malaysia, Pahang, Kuala Baloh (hololectotype K). Synonym: Shorea glauca non King: Browne op.
cit. 168, p.p.
Emergent tree, to 40 m tall, to 1.2 m diameter; bole straight; crown rather flat, diffuse, with
a few large branches, pale from below; buttresses to 1.5 m tall, thin. Bark reddish to
yellowish brown, cracked and irregularly oblong flaky. Young twig, inflorescence, leaf bud,
stipule, bracteole, leaf below, and petiole cream lepidote; parts of calyx exposed in bud,
ovary, nut, and base of fruit calyx outside creambuff puberulent. Twigs terete, dark
chocolatebrown, c. 1.5 mm diameter apically; stipule scars pale, short, falcate. Leaf buds
linear, to 3 × 1 mm. Stipules linear, to 12 mm long, fugaceous. Leaves thinly coriaceous,
drying pale brown above, cream lepidote below with darker veins; blade broadly ovate, 8–
15 × 3.5–8.5 cm, base unequal, cuneate to subcordate, apex with slender acumen to 1.5 cm
long; midrib evident, flat above, broad but not prominently raised below; lateral veins 9–12
pairs, slender and elevated below; intercostal venation slender, densely scalariform,
unraised; petiole 1.2–2 cm long. Inflorescences terminal or axillary; rachis slender, to 15
cm long, singly branched, branchlets bearing to 12 flowers; bracteoles ovate, acute, to 2 mm
long. Flowers: buds to 10 × 4 mm; corolla cream; stamens c. 30, filament and anther
glabrous, connectival appendage short, setose; ovary and stylopodium ovoidconical, style
short, glabrous. Fruits: pedicel c. 2 mm long; calyx lobes unequal, 3 longer lobes to 9 × 3
cm, tapering to 8 mm above the saccate base, 2 shorter ones linearlobed, to 6 × 0.8 cm,
similar at base. Nuts broadly ovoid, to 1.2 × 1.2 cm, tapering to an acute 4 mm style
remnant.
Vernacular name. Sarawak—selangan batu pasir (preferred name).
290
Distribution. E Sumatra?, East coastal Peninsular Malaysia and Borneo. In Sarawak

recorded from Bintulu, Marudi and Miri districts (e.g., S 907, S 8254 and S 8661). Also
occurring in Brunei (e.g., BRUN 513 and FMS 30553).
Ecology. Very local, there sometimes quite common in kerangas forest on white sand
terraces and in mixed dipterocarp forest on leached greyish yellow sands on narrow ridges,
at altitudes to 800 m. Occurring in Lambir NP; elsewhere endangered.
75. Shorea maxwelliana King

(Sir George Maxwell, 1871–1959, onetime Resident of Perak, Peninsular Malaysia)
J. As. Soc. Beng. 62, 2 (1893) 114; Symington op. cit. (1933) 146, op. cit. (1934) 28, op. cit. (1938)
325, op. cit. (1943) 21; Masamune op. cit. 495; Browne op. cit. 169; Ashton op. cit. (1964) 141, op.
cit. (1968) 74, op. cit. (1982) 465; Meijer & Wood op. cit. 182; Burgess op. cit. 203; Anderson op. cit.
(1980) 120; PROSEA op. cit. 432; Coode et al. (eds.) op. cit. 79; Newman et al. op. cit. (1998) 193.
Lectotype (designated here): King’s Collectors 3744, Peninsular Malaysia, Perak (hololectotype K).
Synonyms: Shorea utilis King op. cit. 119; S. barbata Brandis op. cit. 81; S. alba Ridl., J. Str. Br.
Roy. As. Soc. 82 (1920) 171; Balanocarpus ovalifolius Ridl., J. Fed. Malay States Mus. 10 (1920)
130, p.p.
Large emergent tree, to 50 m tall, to 1.6 m diameter; bole tall, cylindrical; crown large,
diffuse, cauliflowershaped; buttresses to 4 m tall, thin, prominent. Bark reddish to greyish
brown, becoming shallowly densely cracked and small thin oblong flaked, eventually
sometimes shaggy. Leaf bud, parts of perianth exposed in bud, fruit calyx, inflorescence,
stipule, bracteole, ovary, and nut evenly buffpuberulent. Twigs slender, muchbranched,
terete, smooth, c. 0.7 mm diameter apically. Leaf buds globose to conical, minute, c. 1 mm
diameter. Stipules linear, to 4 mm long, fugaceous. Leaves thinly coraceous, sometimes
cream lepidote below, more or less shiny above, drying pale yellowish brown; blade ovate
lanceolate, 6–10 × 2.5–4 cm, base obtuse or broadly cuneate, apex caudate, acumen to 2
cm long; midrib raised below, evident, raised but shallowly furrowed above; lateral veins 8–
10 pairs, slender, raised but not prominent below, with or without minute axillary domatia;
intercostal venation very slender, scalariform; petiole 0.7–1 cm long, slender.
Inflorescences terminal or to 3axillary; rachis terete, straight, to 5 cm long, singly
branched, branchlets short bearing to 6 close flowers. Flowers: buds globose, to 2.5 mm
diameter; petals cream; stamens c. 30, filaments glabrous, anthers setose apically,
connectival appendage as long as anther, setose apically; ovary and stylopodium ovoid to
pyriform, style short glabrous. Fruits: calyx lobes unequal, 3 longer lobes to 10 × 1.5 cm,
tapering to 4 mm above the saccate base, 2 shorter ones to 6 × 0.6 cm, otherwise similar.
Nuts ovoid, to 2 × 1.5 cm, tapering, apiculate.
Vernacular names. Sabah—selangan batu asam (preferred name). Sarawak—kumus hitam

(preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah, recorded from Beaufort,
Kudat, Lahad Datu, Sandakan, Sipitang, Tawau, and Tenom districts (e.g., SAN 15221, SAN
16644, SAN 28972, and SAN 66156) and in Sarawak from Bau, Belaga, Kapit, Kuching,
Limbang, Lundu, Marudi, Miri, and Tatau districts (e.g., S 1816, S 7995, S 10195, S 19168,
and S 46549). Also occurring in Brunei (e.g., BRUN 3397 and BRUN 5774) and SE
Kalimantan (e.g., bb. 11212 and bb. 27761).
291
Ecology. Locally frequent in mixed dipterocarp forest and occasionally kerangas, on a wide
range of soils and substrates including basalt, shale, porphyry dykes, granodiorite and
sandstone, at altitudes to 600 m. Occurring in Bako, G. Gading, Kubah, Lambir and Mulu
NPs, and Sepilok FR; not vulnerable.
76. Shorea mecistopteryx Ridl.

(Greek, megisto = very big, pteron = wing; the fruit calyx lobes)
Bull. Misc. Inform. Kew (1925) 280, Slooten op. cit. (1929) 203; Symington op. cit. (1938) 348;
Masamune op. cit. 495; Ashton op. cit. (1964) 198, op. cit. (1968) 111, op. cit. (1982) 525; Meijer &
Wood op. cit. 119; Burgess op. cit. 154, 182; Anderson op. cit. (1980) 127; PROSEA op. cit. 397;
Coode et al. (eds.) op. cit. 79; Newman et al. op. cit. (1996) 168. Type: Taha s.n., Borneo, Sabah,
Kinabatangan (holotype K). Synonym: Shorea chrysophylla Ridl., Bull. Misc. Inform. Kew (1926)
202.
Large emergent tree, to 50 m tall, to 1.6 m diameter; bole tall, cylindrical or sometimes
sinuous; crown large, becoming irregularly hemispherical, golden from below; buttresses to
2 m tall, large, stout. Bark long remaining smooth, hoopmarked, greyish brown, eventually
cracking and thinly flaking, leaving a scrollmarked surface; inner bark dull rustbrown to
orangebrown at the cambium; heartwood pinkish brown, light. Young twig, inflorescence,
stipule and bracteole outside (sparsely so within), petiole, and leaf below evenly persistently
golden pubescent; nut yellowish buffpubescent. Twigs compressed, to 6 × 2.5 mm apically;
stipule scars cuneate, horizontal. Leaf buds ovoid, 5–8 × 2.5–5 mm, obtuse, compressed.
Stipules hastate, acute, to 25 × 7 mm, not immediately caducous. Leaves chartaceous,
brittle when dry and turning rich reddish brown below, paler above; blade oblong, 13–20(–
30) × 6–10(–12) cm, base cordate, apex with broad acumen to 0.8 cm long; midrib evident,
more or less flat or shallowly furrowed above, prominent and terete below; lateral veins
16–20 pairs, slender but prominent below, arched; intercostal venation densely scalariform,
elevated below; petiole 2.5–3.5 cm long, stout. Inflorescences terminal or axillary, rachis
terete or compressed, to 12 cm long, singly branched; bracteoles oblong, subacute, c. 7 × 2
mm. Flowers: buds large, to 12 × 4 mm; connectival appendage more than 2x the length of
anther; ovary ovoidconical, glabrous, stylopodium slightly longer than ovary, cylindrical,
thickened distally, style stout. Fruits: pedicel to 6 mm long, stout; calyx lobes unequal,
glabrescent or puberulent towards base, 3 longer lobes to 23 × 3.3 cm, tapering to 1.8 cm
above the saccate base, 2 shorter ones to 15 × 1.2 cm, otherwise similar. Nuts ovoid, to 4.2
× 2.5 cm, acute.
Vernacular names. Sabah—kawang burung (preferred name). Sarawak—engkabang

burung (preferred name), engkabang larai (Iban), kawang tikus (Malay).
Distribution. Endemic in Borneo; widespread but infrequent and local. In Sabah recorded
from Kinabatangan and Sandakan districts (e.g., SAN 4975, SAN 4977, SAN 5493, SAN
20811, and SAN A 3150) and in Sarawak from Kapit, Lundu, Marudi, and Miri districts
and S 1634) and C Kalimantan (e.g., Argent 9495, Newman et al. 625, Newman et al. 626,
and Peters 1035).
292
Ecology. Local but widespread in mixed dipterocarp forest on sandy clay soils, including on
intermediate igneous rocks, at altitudes to 400 m. Recorded from G. Gading NP; elsewhere
77. Shorea micans P.S.Ashton

(Latin, micans = gleaming; the shiny leaf)
Gard. Bull. Sing. 31 (1978) 38, op. cit. (1982) 463; Newman et al. op. cit. (1998) 193. Type: Meijer
SAN 39312, Borneo, NE Sabah, Karamuak, Bt. Meliau (holotype L; isotypes KEP, SAN).
Low emergent or main canopy tree. Bark flaky. Inflorescence and nut densely greyish
puberulent, fruit calyx sparsely so; other known parts glabrous, shiny. Twigs terete, much
branched, slender, c. 1 mm diameter apically. Leaf buds minute. Stipules unknown. Leaves
thinly coriaceous, shiny, drying dark tawnybrown below, greyish brown above; blade
ovatelanceolate, 5–10 × 1.8–4.7 cm, base subequal, broadly cuneate, margin narrowly
subrevolute, apex subcaudate, acumen to 1.3 cm long; midrib slender, evident and raised on
both surfaces; lateral veins 7–8 pairs, very slender, slightly elevated below; intercostal
venation scalariform, obscure; petiole very slender, 0.7–1.2 cm long. Inflorescences
terminal or subteminalaxillary; rachis slender, straight, shortly branched, to 7 cm long.
Flowers unknown. Fruits: pedicel c. 1 mm long, slender; calyx lobes unequal, 3 longer
lobes to 5 × 1.5 cm, tapering to c. 4 mm above the saccate base, 2 shorter ones to 2.5 × 0.4
cm, otherwise similar. Nuts ovoid, to 1.9 × 0.7 cm, prominently apiculate.
Distribution. Endemic in Borneo. Known only in Sabah from Kuala Penyu, Labuk Sugut
and Sandakan districts (e.g., SAN 24279, SAN 50994 and the type).
Ecology. In mixed dipterocarp forest on soils overlying ultrabasic rock, at altitudes to 500
m. Critically endangered.
78. Shorea monticola P.S.Ashton

(Latin: montis = mountain, cola = dweller; the habitat)
Newman et al. op. cit. (1996) 169. Type: Clemens 31885, Borneo, Sabah, Mt. Kinabalu, Penibukan
(holotype K; isotype L).
Low emergent tree to 40 m tall, to 1.2 m diameter; bole cylindrical, straight; buttresses to
1.5 m tall, stout; crown densely evenly hemispherical, golden suffused from below. Bark
pinkish to reddish brown, becoming shallowly cracked and thinly patchily oblong flaked;
inner bark reddish brown; heartwood dark red, hard. Young twig, leaf bud, petiole, venation,
inflorescence, and parts of perianth exposed in bud at first yellowish buffpubescent,
persistent on leaf bud, corolla and inflorescence, sparsely persisting in calyx base into fruit,
elsewhere caducous. Twigs terete, minutely warty lenticellate, stout, c. 3 mm diameter
apically. Leaf buds narrowly ovate, acute, falcate, 5–10 × 2–4 mm. Stipules unknown.
293
Leaves thickly coriaceous, drying golden to chocolatebrown below, tawnybrown above;

blade elliptic, 8–13 × 5–8 cm, base obtuse to broadly cuneate, apex with broad, short,
acumen to 0.5 cm long; midrib obscure, sunken above, prominent and sharply ridged below;
lateral veins 13–16 pairs, prominent below, arched; intercostal venation densely
scalariform, slender, hardly elevated or obscure; petiole 2–2.5 cm long, stout.
Inflorescences terminal or axillary; rachis terete, to 12 cm long, branchlets bearing to 12
flowers; bracteoles unknown. Flowers: buds to 7 × 3 mm; connectival appendage 2–3x as
long as anther; ovary and stylopodium pyriform, style as long as ovary, glabrous. Fruits:
calyx lobes unequal, 3 longer lobes to 7 × 1.5 cm, tapering only slightly above the saccate
base, 2 shorter ones narrowly oblong, to 3.5 × 0.6, similar at base. Nuts ovoid, to 1.6 × 1
cm, shortly acute.
Vernacular names. Sabah—seraya burung (preferred name). Sarawak—meranti gunung

(preferred name).
Distribution. Endemic in Borneo; throughout the northern mountain chain from W

Kalimantan and G. Penrissen, W Sarawak, to Kinabalu and Trus Madi Range in northern
Sabah. In Sabah recorded from Keningau, Kota Belud, Papar, Ranau, Sipitang, and
Tambunan districts (e.g., SAN 16704, SAN 16914, SAN 24058, SAN 50206, and SAN A
4353) and in Sarawak from Bintulu, Kapit, Kuching, Lawas, Miri, Sri Aman, and Tatau
districts (e.g., S 4525, S 8816, S 19169, S 26524, and S 67522).
Ecology. Locally common in the upper limits of upper dipterocarp forest, on organic soils
on various substrates including basalt, at 600–1500 m altitude. Common in Kinabalu NP,
and occurring in Mulu NP; probably not vulnerable.
79. Shorea mujongensis P.S.Ashton

(of the Mujong, Ulu Tinjar, Rajang, Sarawak)
Kessler & Sidiyasa op. cit. 106; Newman et al. op. cit. (1996) 170. Type: Ashton S 19038, Borneo,
Sarawak, Hose Mountains, Base of Bt. Temedu (holotype K; isotypes KEP, L, SAR).
Vast tree to 70 m tall, to 1.6 m diameter; crown dense, cauliflowershaped; bole tall,
cylindrical; buttresses stout, large, to 5 m tall and wide. Bark fawnbrown, deeply cracked
and small thin oblongflaky; dammar in dark brown coxcombs. Young twig, leaf bud, parts
of perianth exposed in bud, ovary, and nut buffpuberulent, caducous other than on
inflorescence, corolla and ovary. Twigs terete, rugulose, c. 2 mm diameter apically. Leaf
buds minute. Stipules unknown. Leaves glabrous, chartaceous, drying greenish grey; blade
ovate or elliptic, 6–14 × 2.5–5.5 cm, base obtuse or cuneate, margin narrowly revolute,
apex with acumen to 0.8 cm long; midrib evident, flat above, prominent and terete below;
lateral veins 8–13 pairs, slender but prominent below; petiole 1–1.6 cm long.
Inflorescences terminal or axillary; rachis terete, to 6 cm long, singly branched. Flowers:
buds to 6 × 4 mm; stamens 15, connectival appendage as long as anther, glabrous; ovary
and stylopodium pyriform, style short, glabrous. Fruits: pedicel to 2 mm long; calyx lobes
unequal, 3 longer lobes to 7 × 1.5 cm, tapering to 4 mm wide above the narrowly saccate
tuberculate base, 2 shorter ones to 4.5 × 0.7 cm, otherwise similar. Nuts narrowly ovoid, to
2.2 × 0.7 cm.
294
Distribution. Endemic in Borneo. Recorded in Sabah from Beaufort, Sandakan and Tawau
districts (e.g., SAN 4946, SAN 15429 and SAN 27425) and in Sarawak from Kapit and
Kuching districts (e.g., S 19038, S 19993 and S 29569). Also occurring in E Kalimantan.
Ecology. Frequent, but very local in mixed dipterocarp forest, and apparently mostly on
basic to weakly acid volcanic rocks, on clay soils, at altitudes to 1000 m. Endangered by
land conversion.
80. Shorea multiflora (Burck) Symington Plate 4D.

(Latin, multi = many, flos = a flower; the manyflowered inflorescence)
Gard. Bull. S. S. 9 (1938) 330, op. cit. (1943) 54; Slooten op. cit. 3 (1956) 320; Browne op. cit. 63;
op. cit. 218; Anderson op. cit. (1980) 122; PROSEA op. cit. 420; Coode et al. (eds.) op. cit. 80;
Newman et al. op. cit. (1996) 171. Basionym: Doona multiflora Burck op. cit. (1887) 234. Type:
Teijsmann HB 12063, Sumatra (holotype BO). Synonyms: Richetia latifolia F.Heim op. cit. (1891)
976; R. acuminata F.Heim op. cit. (1891) 979; R. oblongifolia F.Heim op. cit. (1891) 979; R.
penangiana F.Heim op. cit. (1891) 980; Balanocarpus penangianus (F.Heim) King op. cit. 131;
Hopea multiflora (Burck) Brandis op. cit. 60; H. multiflora (Burck) Brandis var. venosa Boerl., Cat.
Hort. Bog. 2 (1901) 102; Balanocarpus latifolius (F.Heim) Brandis op. cit. 112, Merrill op. cit. (1921)
407; B. acuminatus (F.Heim) Brandis op. cit. 115, Merrill op. cit. (1921) 407; B. sibogae Boerl., op.
cit. 112; B. multiflorus (Burck) Symington op. cit. (1933) 153.
Low emergent or main canopy tree, to 40 m tall, to 1 m diameter; bole straight, cylindrical;
crown diffusely hemispherical, with many ascending branches; buttresses short, stout. Bark
dappled fawn, becoming densely cracked and thin oblongflaky. Young twig, petiole, stipule
outside, inflorescence, leaf bud, parts of perianth exposed in bud, and usually ovary and
nut, densely greyish buffpuberulent, persistent except on twig, petiole and calyx; ovary and
nut sometimes glabrous. Twigs slender, terete, smooth, muchbranched, c. 1 mm diameter
apically. Leaf buds small, ovoid, c. 2 × 1.5 mm. Stipules narrowly lanceolate, to 6 × 2 mm,
fugaceous. Leaves thinly coriaceous, drying yellowish green to yellowish brown; blade
ovatelanceolate, 4.5–7.5 × 2–3.5 cm, base cuneate, equal or subequal, sometimes with
paired basal domatia, apex caudate, acumen to 1.5 cm long; midrib narrowly shallowly
sunken above, somewhat raised below; lateral veins 8–10 pairs, very slender, barely
elevated below, distant; intercostal venation subreticulate; petiole slender, 0.7–1 cm long.
Inflorescences terminal or axillary; rachis terete, lax, to 16 cm, branchlets short, bearing to
9 flowers; bracteoles minute, fugaceous. Flowers: buds small, to 2.5 mm long; petals pale
yellow, linear; stamens (10–)15, connectival appendage 1½–2x as long as anther, ciliate
distally; ovary and stylopodium ovoidtapering, style glabrous, with stylopodium as long as
ovary. Fruits: calyx lobes subequal, deltoid, acute, saccate, thickened, to 0.5 × 0.4 cm. Nuts
obovoid, to 2 × 1.2 cm, mucronate, pubescent or glabrous and shiny.
Vernacular names. Sabah—banjutan (preferred name). Sarawak—lun jantan (preferred

name).
Distribution. Sumatra, Peninsular Malaysia and Borneo (common throughout the island). In
Sabah known from Beaufort, Kota Kinabalu, Labuk Sugut, Lahad Datu, Semporna,
Sipitang, and Tawau districts (e.g., SAN 15081, SAN 15503, SAN 16464, SAN 38705, and
SAN 131979) and in Sarawak from Belaga, Bintulu, Kapit, Kuching, Limbang, Lubok Antu,
295
Lundu, Marudi, Miri, and Tatau districts (e.g., S 13761, S 23812, S 37845, S 46588, and S
69656). Also occurring in Brunei (e.g., BRUN 2630 and BRUN 5649) and SE Kalimantan
(e.g., bb. 34427 and Church et al. 427).
Ecology. The most abundant of the yellow merantis: in kerangas, and mixed dipterocarp
forest on the yellow sands of the coastal hills and the inland shale ridges; also on organic
soil over karst limestone; but rare or absent from moist clay slopes and the deep friable
clays of basic volcanic rocks and calcareous shale, at altitudes to 900 m. In Sabah, common
in Sepilok FR and recorded from Kinabalu NP, and in Sarawak, common in Bako, Kubah,
Lambir and Mulu NPs; not vulnerable.
Notes. Shorea hopeifolia can be confused with this species (see there).
81. Shorea myrionerva Symington ex P.S.Ashton

(Greek, myrio = countless; Latin, nervus = vein; the manyveined leaf blade)
Newman et al. op. cit. (1996) 173. Type: Drahman S 1519, Borneo, Sarawak, Lawas district,
Merapok (holotype KEP; isotype L).
Canopy, occasionally emergent tree, to 45 m tall, to 1.2 m diameter; bole often leaning;
crown loose, generally oblong, the branches pendent with upturned twig endings and
hanging leaves; buttresses to 1.5 m but usually less, stout. Bark at first smooth, becoming
shallowly vsection fissured then powdery flaky, often mosscovered; inner bark yellowish
brown; heartwood pinkish brown. Twig and petiole persistently sparsely coarsely pale
fulvous hispid; venation below and stipule more shortly so; inflorescence sparsely so
towards base; midrib above, stipule and bracteole outside puberulent; leaf blade
glabrescent; other parts glabrous. Twigs terete, straight, fewbranched, stout, c. 3 mm
diameter apically. Leaf buds imbedded in a loose group of bud scales, compressed, to 10 ×
7 mm. Stipules broadly hastate, subauriculate, subpersistent, to 17 × 7 mm. Leaves
chartaceous, drying reddish brown; blade narrowly oblong, 12–22 × 4.5–9 cm, base obtuse,
apex tapering abruptly, acumen to 0.8 cm long; midrib obscurely sunken above, prominent
below; lateral veins 24–28 pairs, prominent, arched towards margin; intercostal venation
slender, scalariform; petiole stout, 1.2–2 cm long. Inflorescences axillary (rarely terminal)
to ramiflorous; rachis terete to ribbed on drying, lax, singly branched, the branchlets short
bearing to 5 flowers; bracteoles ovate, obtuse, to 8 × 1.5 mm. Flowers: buds to 5 × 3 mm;
petals dark red with pale margin; stamens 15, connectival appendage short, becoming
reflexed; ovary and stylopodium narrowly conical to pyriform, style shorter than both,
columnar, glabrous. Fruits: calyx lobes unequal, 3 longer lobes oblong, obtuse,
chartaceous, to 17 × 2.5 cm, not tapering, the thickened base surrounded by prominent
lateral auricles, 2 shorter ones linearlobed, to 8 × 0.7 cm, saccate but not auriculate. Nuts
ellipsoid, to 2.5 × 1.3 cm, pruinose, with c. 1.5 cm tapering style remnant.
Vernacular names. Sabah—seraya urat banyak (preferred name). Sarawak—langgai sepit

udang (Iban), meranti sepit udang (preferred name).
296
Distribution. Endemic in Borneo. In Sabah known from Beaufort district (e.g., KEP 80268
and KEP 80269) and in Sarawak from Bintulu, Kapit, Lawas, Limbang, Marudi, and Miri
(e.g., BRUN 5200 and BRUN 5218) and Kalimantan (e.g., Argent et al. 9492).
Ecology. Locally common in mixed dipterocarp and riparian forest on moist clay soils on
lower hillsides and the banks of both fast and sluggish rivers, at altitudes to 400 m.
Occurring in Lambir and Mulu NPs; elsewhere vulnerable owing to the loss of its habitat.
82. Shorea obovoidea Slooten

(Latin, obovoideus = obovoid; the fruit)
Newman et al. op. cit. (1996) 477. Type: Egon SA 616, Borneo, Sarawak, Semengoh FR (holotype
KEP).
Main canopy tree, to 35 m tall, to 60 cm diameter; bole straight, cylindrical; crown small,
dense, hemispherical; buttresses low, stilt roots sometimes present. Bark dark tawnybrown,
smooth to shallowly patchily flaky. Leaf bud, twig, inflorescence, parts of perianth exposed
in bud, and petiole densely somewhat unevenly golden brown puberulent; venation below
sparsely so; ovary and nut evenly buffpubescent. Twigs terete, slightly rough, c. 2 mm
diameter apically. Leaf buds ovoid, acute, to 2 × 1 mm. Stipules unknown. Leaves
coriaceous, drying tawny brown below, greyish brown above; blade elliptic, 5–13 × 1.7–4.5
cm, base cuneate, margin minutely revolute, apex subcaudate, acumen to 1.2 cm long;
midrib narrowly evident but furrowed above, prominent below; lateral veins 8–10 pairs,
arched, slender but prominent below; intercostal venation subreticulate, elevated below;
petiole slender, 0.5–1.1 cm long. Inflorescences terminal or axillary; rachis terete, singly
branched, branchlets bearing to 5 flowers; bracteole minute, deltoid, acute, c. 1 mm long,
fugaceous. Flowers: buds small, to 3 × 1 mm; corolla cream; stamens 15, connectival
appendage slightly longer than anther; ovary ovoid, without stylopodium, style columnar,
stout, slightly shorter than ovary, puberulent in the basal half. Fruits: pedicel c. 1 mm long;
calyx lobes subequal, ovate, acute, relatively thin, hardly saccate, to 1.2 × 0.9 cm. Nuts
ellipsoid, to 2.3 × 1.6 cm, acute.
Distribution. Endemic in Borneo. Known in Sarawak from Kuching, Kapit and Samarahan
districts (e.g., S 9102, S 11072, S 14368, S 15225, and S 67572). Also occurring in SE
Ecology. Very local in mixed dipterocarp forest, on leached sandy clay soil on low hills, at
altitudes to 300 m. Endangered by forest conversion.
83. Shorea obscura Meijer

(Latin, obscurus = obscure; originally confused with S. atrinervosa)
297
Act. Bot. Neerl. 12 (1963) 333; Meijer & Wood op. cit. 184; Ashton op. cit. (1964) 142, op. cit. (1968)
Coode et al. (eds.) op. cit. 80; Newman et al. op. cit. (1998) 193. Lectotype (designated here): G.H.S.
Wood SAN 15166, Borneo, Sabah, Bt. Sipitang (hololectotype K; isolectotypes KEP, L, SAN).
Emergent tree, to 60 m tall, to 1.3 m diameter; bole tall, straight; crown diffuse cauliflower
shaped, pale from below; buttresses to 6 m tall, thin. Bark greyish to reddish brown, closely
cracked and evenly oblong flaky. Leaf venation below, petiole, twig, inflorescence, leaf bud,
parts of perianth exposed in bud, stipule, bracteole, ovary, and fruit densely pale buffbrown
puberulent, caducous except on inflorescence, buds, stipule and bracteole. Twigs terete,
slender, 1.5–2 mm diameter apically. Leaf buds ovoid to falcate, 2–4 × 1.5–2.5 mm.
Stipules oblong, acute, to 7 × 2.5 mm, caducous. Leaves coriaceous, cream lepidote below
with the veins drying darker brown, drying greyish brown above; blade ovatelanceolate, 7–
12 × 2.5–5 cm, base cuneate, apex with slender acumen to 1 cm long; midrib evident, flat or
elevated above, more prominently so below; lateral veins 7–9 pairs, arched, distinctly
raised below, intercostal venation scalariform, slender, hardly elevated, obscure; petiole
slender, 1.2–2 cm long. Inflorescences terminal or axillary; rachis terete, to 12 cm long,
singly branched, branchlets bearing to 5 flowers; bracteoles ellipticovate, acute, c. 3 mm
long, fugaceous. Flowers: buds to 6 × 2 mm; petals bright pink, cream at margin; stamens
35–45, filaments and anthers glabrous, connectival appendage almost as long as anther,
setose; ovary and stylopodium ovoid, tapering into the short glabrous style. Fruits: calyx
lobes unequal, 3 longer lobes to 10 × 2 cm, tapering 5 mm broad above the saccate base, 2
shorter ones oblong, lobed, to 5 × 0.5 cm, similar at base. Nuts globose to ellipsoid, to 1.2
× 0.9 cm, abruptly apiculate.
Vernacular names. Sabah—selangan batu tanduk (preferred name). Sarawak—selangan

batu padi (preferred name).
Distribution. Endemic in Borneo. In Sabah known from Kota Kinabalu (Gaya Is.), Ranau,
Sipitang, Tawau, and Tuaran districts (e.g., SAN 15166, SAN 16772, SAN 20651, SAN
22370, and SAN 99400) and in Sarawak from Belaga, Limbang, Lundu, and Tatau districts
BRUN 5438 and Prance 30589) and SE Kalimantan (e.g., bb. 16966 and bb. 22911).
Ecology. Locally frequent in mixed dipterocarp forest on leached sandy clay and well
drained clay soils, especially on spurs and ridges, probably most abundant at 600–800 m
altitude, occasionally to 1400 m in upper dipterocarp forest. Occurring in Kinabalu and
Mulu NPs; elsewhere vulnerable.
84. Shorea ochracea Symington Fig. 26.

(Latin, ochraceus = the colour of ochre, warm yellow; the leaf undersurface)
Gard. Bull. S. S. 8 (1935) 285; Masamune op. cit. 495; Browne op. cit. 158; Ashton op. cit. (1964)
165, op. cit. (1968) 94, op. cit. (1982) 492; Meijer & Wood op. cit. 57; Burgess op. cit. 159; Anderson
op. cit. (1980) 123; PROSEA op. cit. 412; Coode et al. (eds.) op. cit. 80; Newman et al. op. cit. (1996)
174. Lectotype (designated here): Spurway S 167, Borneo, Sarawak, Baleh, Merirai (hololectotype
KEP).
298
Fig. 26. Shorea ochracea. A, leafy twig with young shoots; B, abaxial view of bract; C,
adaxial view of bract; D, abaxial view of bracteole; E, adaxial view of bracteole; F, flower
bud; G, adaxial view of outer sepal; H, adaxial view of inner sepal; I, abaxial view of petal;
J, adaxial view of petal; K, gynoecium and stamens; L, adaxial view of stamens; M, abaxial
view of stamens; N, fruit; O, fruit with the nut partially exposed. (A from S 32359, B–O
from BRUN 3274.)
299
Fig. 27. Shorea ovalis, subsp. ovalis (N–O) and subsp. sarawakensis (A–M). A, flowering
leafy twig; B, detail of venation and indumentum on lower leaf surface; C, portion of
inflorescence; D, opening flower; E, longitudinal section of open flower; F, abaxial view of
bract; G, adaxial view of outer sepal; H, adaxial view of inner sepal; I, adaxial view of petal;
J, abaxial view of petal; K, adaxial view of stamens; L, abaxial view of stamens; M,
gynoecium; N, fruit; O, longitudinal section of fruit. (A–M from S 15159, N–O from SAN
16474.)
300
Large emergent tree, to 50 m tall, to 1.3 m diameter; bole straight or malformed; crown
irregularly hemispherical, dense, largeleaved, pale yellowish from below; buttresses to 1.5
m tall, very stout. Bark mottled pale brown, becoming deeply prominently irregularsection
fissured and eventually crumbly flaky, dark chocolatebrown; inner bark pale yellowish
brown and creamyellow laminated; dammar pale creamyellow, as smears on bole. Twig,
leaf bud, inflorescence, parts of perianth exposed in bud, fruit calyx, stipule, bracteole, and
petiole densely persistently rufous brown or greenish yellow powdery pubescent, partially
caducous, sparse on leaf below; ovary and nut densely evenly cream pubescent. Twigs
terete, with horizontal or ascending stipule scars, stout, 4–5 mm diameter apically. Leaf
buds globose to ovoid, obtuse, 3–6 × 3–5 mm. Stipules elliptic, obtuse, to 20 × 15 mm,
caducous. Leaves thickly coriaceous and somewhat turned up along the midrib, bright
yellow lepidote below (mature trees); blade broadly ellipticoblong, 12–18 × 7–10 cm
(larger in juveniles), base cordate, apex with broad acumen to 0.5 cm long; midrib obscure,
sunken, above, prominently terete below; lateral veins 25–30 pairs, dense, arched,
prominent below; intercostal venation densely scalariform, obscure; petiole stout, 1.3–2 cm
long. Inflorescences terminal or axillary; rachis terete, lax, to 10 cm long, branchlets
bearing to 8 flowers; bracteoles large, elliptic, obtuse, subpersistent, to 20 × 10 mm.
Flowers: buds to 8 × 3 mm; stamens 15, anther oblong, tapering, connectival appendage
somewhat longer than anther; ovary ovoid, without stylopodium, style glabrous except
towards base, as long as ovary. Fruits: calyx lobes unequal, 3 longer lobes to 10 × 2 cm,
oblong, tapering to 1 cm above the narrowly auriculate centrally thickened base, 2 shorter
ones to 4.5 × 0.5 cm, unequal, otherwise similar. Nuts ovoid, to 1.5 × 1 cm, with to 2.5 mm
style remnant.
Vernacular names. Sabah—melapi daun besar (preferred name). Sarawak—raruk (Iban).
Distribution. Endemic in Borneo. In Sabah recorded from Beaufort, Kinabatangan,

Sipitang, and Tawau districts (e.g., SAN 15229, SAN 16813, SAN 19537, SAN 27974, and
SAN 93773) and in Sarawak from Belaga, Bintulu, Kapit, Limbang, and Miri districts (e.g.,
S 15117, S 29685, S 32359, S 46447, and S 69173). Also occurring in Brunei (e.g., BRUN
3274 and BRUN 5434) and W and NE Kalimantan (e.g., Arifin Berau 1092 and Kessler et
al. Berau 891).
Ecology. Scattered, usually as isolated individuals, in mixed dipterocarp forest on

undulating land and hills, on sandy clay and clay soils, at altitudes to 750 m. Occurring in
Lambir and Mulu NPs; elsewhere vulnerable owing to both land conversion and logging.
85. Shorea ovalis (Korth.) Blume Fig. 27.

(Latin, ovalis = oval; the leaf shape)
(sect. Ovalis, red meranti)
Mus. Bot. Lugd.Bat. 2 (1852) 33; Merrill op. cit. (1921) 406; Symington op. cit. (1939) 370, op. cit.
(1943) 80; Masamune op. cit. 495; Browne op. cit. 132; Ashton op. cit. (1963) 274, op. cit. (1982)
548; Meijer & Wood op. cit. 125 p.p.; Burgess op. cit. 154, 183; PROSEA op. cit. 398; Kessler &
Sidiyasa op. cit. 106; Newman et al. op. cit. (1996) 176. Basionym: Vatica ovalis Korth., Kruidk.
(1841) 73. Type: Korthals s.n. (= RHL Sheet No. 902146618), Borneo, Kalimantan, Prarawing
(holotype L). Synonyms: Vatica eximia Miq., op. cit. 486; V. sublacunosa Miq., op. cit. 486; Hopea
aspera de Vriese op. cit. 28; Shorea eximia (Miq.) Scheff., Nat. Tijd. Ned. Ind. 31 (1870) 349; S.
eximia (Miq.) Scheff. var. angustifolia Burck op. cit. (1887) 218; S. sublacunosa (Miq.) Scheff., op.
cit. 350.
301
hemispherical, rather even, with many ascending branches; buttresses to 1.5 m tall, stout.
Bark pale pinkish brown and grey mottled, becoming shallowly vsection fissured,
eventually patchily powdery flaky; inner bark pale reddish brown; heartwood pink,
relatively light. Twig, inflorescence, parts of perianth exposed in bud, stipule, bracteole,
petiole, and leaf below densely persistently scabridpuberulent to coarsely pubescent or
tufted tomentose or sparsely so; ovary and nut densely evenly pubescent; fruit calyx sparsely
puberulent. Twigs terete, straight, sparingly branched, stout, c. 3 mm diameter apically;
stipule scars cuneate, slightly ascending. Leaf buds ovoid, obtuse, to 8 × 6 mm. Stipules
ovate, acute, prominently veined, to 13 × 7 mm, subpersistent. Leaves coriaceous, weakly
blistered in juveniles, drying rich reddish brown below, pinkmauve above; blade narrowly
oblongovate, 10–18 × 3–7 cm, or narrowly oblong, 12–17 × 2–4.5 cm, base obtuse,
margin more or less narrowly revolute, apex with narrow acumen, to 0.8 cm long; midrib
narrowly evident but furrowed above, prominent below; lateral veins (20–)22–25 pairs,
prominent below, arched, dense; intercostal venation distinctly elevated, scalariform;
petiole 0.7–0.9 cm long. Inflorescences terminal or axillary; rachis rigid, to 18 cm long,
branchlets compact bearing to 8 flowers; bracteole broadly elliptic, obtuse, to 8 × 6 mm,
concave round bud. Flowers: buds broadly ovoid to subglobose, to 5 mm long; petals
cream, pinktinged at base; stamens 50–70, connectival appendage short, vestigial; ovary
and stylopodium narrowly conical, style short, cylindrical, glabrous. Fruits: calyx lobes
ones linearlobed, to 6 × 0.4 cm, similar at base. Nuts ovoid, to 2.2 × 1.3 cm, with to 2.5
mm tapering style remnant.
Vernacular names. Sabah—seraya kepong (preferred name), Sarawak—meranti kepong

(preferred name).
Ecology. Frequent, sometimes common in mixed dipterocarp forest on fertile well

structured clay soils, on low hills, at altitudes to 500 m.
Notes. Two species, viz. subsp. ovalis and subsp. sarawakensis, are recognised.
Key to subspecies
Twigs and inflorescences densely persistently scabridpuberulent to coarsely pubescent.

Leaf blade narrowly oblongovate, 10–18 × 3–7 cm, densely scabridpubescent below;
margin more or less revolute…………………………………………………………………..
subsp. ovalis
Absent from Brunei, Sarawak and W Sabah but widespread in E Sabah and known
from Kinabatangan, Sandakan and Tawau districts (e.g., SAN 16474, SAN 20906, SAN
23973, SAN 39114, and SAN 63079). Also occurring in E and SE Kalimantan (e.g.,
Ambriansyah & Arifin Berau 1072, bb. 34377, Meijer 2198, and Meijer 2222). Ecology
as the species. Occurring in Sepilok FR; elsewhere vulnerable owing to land
conversion.
Twigs and inflorescences prominently tufted tomentose, tufts to 3 mm long. Leaf blade
narrowly oblong, 12–17 × 2–4.5 cm, glabrous below; margin prominently revolute………...
subsp. sarawakensis P.S.Ashton
(of Sarawak)
302
Gard. Bull. Sing. 20 (1963) 275, op. cit. (1968) 111, op. cit. (1982) 549; Meijer & Wood op. cit.
126, p.p.; Burgess op. cit. 154, 183; Anderson op. cit. (1980) 127; PROSEA op. cit. 398; Coode
et al. (eds.) op. cit. 80; Newman et al. op. cit. (1996) 177. Type: Ashton BRUN 3281, Borneo,
Brunei, Andulau FR (holotype K; isotype KEP).
Endemic in Borneo; recorded in Sabah from Beaufort, Kota Merudu, Lahad Datu,
Ranau, Sandakan, Semporna, Sipitang, and Tawau districts (e.g., SAN 15182, SAN
16364, SAN 17850, SAN 18301, and SAN 63754) and in Sarawak from Kuching, Lundu
and Miri districts (e.g., S 15159, S 29472, S 37800, S 46492, and S 49957). Also
occurring in Brunei (e.g., BRUN 3281 and S 1652) and NW Kalimantan (e.g., Suzuki K
9758). As subsp. ovalis, but usually as scattered individuals, on leached clay and sandy
clay soils. Occurring in Kubah, Lambir and Mulu NPs; not vulnerable.
86. Shorea ovata Dyer ex Brandis

(Latin, ovatus = ovate; the leaf shape)
J. Linn. Soc. Bot. 31 (1895) 91; Merrill op. cit. (1921) 406; Symington op. cit. (1933) 140, op. cit.
127; PROSEA op. cit. 398; Coode et al. (eds.) op. cit. 179. Lectotype (designated here): Curtis 201,
Peninsular Malaysia, Penang Hill (hololectotype K). Synonyms: Shorea plagata Foxw., op. cit.
(1918) 192; S. agsaboensis W.L. Stern, Brittonia 17 (1965) 36.
Main canopy or low emergent tree, occasionally to 50 m tall, to 1.4 m diameter but usually
shorter; bole straight, cylindrical; crown hemispherical to cauliflowershaped, usually with
many radiating branches; buttresses to 1 m tall, usually short, stout. Bark at first mottled
fawn, dark brown and ochreous reminiscent of Calophyllum, becoming deeply closely v
section fissured; inner bark deep pink to dark meatred; heartwood dark reddish brown.
Young twig, inflorescence, parts of calyx exposed in bud, leaf bud, stipule, bracteole,
petiole, midrib above, and leaf below densely persistently evenly ochreous to rust
puberulent, scabrous on inflorescence and venation below, sparsely so on fruit calyx; nut
yellowish brown pubescent. Twigs muchbranched, terete, slender, c. 1 mm diameter
apically; stipule scars short, descending. Leaf buds ovoid, obtuse, 3–6 × 1.5–3 mm. Stipules
ovate, subacute, to 13 × 5 mm, caducous. Leaves coriaceous, drying ashenbrown below,
mauvebrown above; blade broadly ovate, 4–8 × 2.5–4.5 cm, base subequal, obtuse, margin
subrevolute, apex with narrow acumen to 1 cm long; midrib obscurely sunken above,
prominent below; lateral veins 8–10 pairs, prominent below, arched, with small pilose
axillary domatia; intercostal venation hardly elevated, slender, densely scalariform; petiole
1–1.5 cm long. Inflorescences terminal or axillary; rachis terete or slightly compressed, to
11 cm long, branchlets bearing to 8 flowers; bracteoles oblong, obtuse, to 3.5 × 2 mm.
Flowers: petals pale pink with cream margin; stamens 15, or 10 by abortion of outer 5,
connectival appendage short, becoming reflexed; ovary ovoid, without stylopodium,
tapering into the short glabrous style. Fruits: calyx lobes unequal, 3 longer lobes to 5.5 × 1
cm, tapering to 4 mm above the saccate base, 2 shorter ones linearlobed, to 3 × 0.3 cm,
otherwise similar. Nuts broadly ovoid, to 1.1 × 0.8 cm; style remnant tapering, to 2.5 mm
long.
Vernacular names. Sabah—seraya punai bukit (preferred name). Sarawak—meranti pitis

(preferred name).
303
Distribution. Sumatra, Peninsular Malaysia, Borneo, and the Philippines (Mindanao).

Widespread in Borneo though not in the south. In Sabah recorded from Kota Belud,
Sipitang and Tambunan districts (e.g., KEP 80407, KEP 80433, SAN 15130, and SAN
16263) and in Sarawak from Bintulu, Kuching, Lundu, Miri, Samarahan, and Simunjan
(e.g., BRUN 551 and Niga NN 227).
Ecology. Locally common, in mixed dipterocarp forests on yellow sandy soil in the
lowlands; also in the ecotone to kerangas, and in upper dipterocarp forest, at 600–1000 m
altitude. Common in Bako NP and frequent in Kinabalu, Kubah, Lambir and Mulu NPs;
elsewhere vulnerable in the lowlands owing to land conversion.
87. Shorea pachyphylla Ridl. ex Symington

(Greek, pachy = thick, phullon = leaf; the leathery leaf)
(sect. Brachyptera, red meranti)
J. Malay. Br. Roy. As. Soc. 19 (1941) 163; Browne op. cit. 148; Anderson op. cit. (1963) 159, op. cit.
(1980) 128; Ashton op. cit. (1964) 205, op. cit. (1968) 112, op. cit. (1982) 511; PROSEA op. cit. 399;
Coode et al. (eds.) op. cit. 81; Newman et al. op. cit. (1996) 180. Type: Haviland 2228, Borneo,
Sarawak, near Kuching (holotype SING; isotype L).
Emergent tree, to 45 m tall, to 1.5 m diameter; bole cylindrical, often tapering; crown dense,
irregularly hemispherical with a few large twisted branches; buttresses to 2.5 m tall,
prominent, stout. Bark pinkish to mauvebrown, becoming deeply vsection fissured, the
intervening ridges thinly flaky; inner bark yellowish brown; heartwood coffeecoloured,
rather hard. Exposed young parts evenly densely tawny puberulent, caducous on all but leaf
bud, inflorescence and petal outside, sparse on stipule and bracteole; other parts glabrous.
Twigs terete to slightly compressed, ridged, stout, c. 4 mm diameter apically; stipule scars
horizontal, long to amplexicaul. Leaf buds narrowly ovoid to broadly falcate, 5–8 × 2–4
mm. Stipules lanceolate, to 20 × 7 mm, fugaceous. Leaves thickly coriaceous, shiny and
drying rich tawny below, greyish brown above; blade broadly ovate to suborbicular, 10–20
× 9–16 cm, base obtuse or broadly cuneate, margin slightly but distinctly revolute, apex
obtuse or with broad acumen to 0.5 cm long; midrib obscure, sunken above, stoutly
prominent below; lateral veins 7–9 pairs, wellspaced, arched at margin, stoutly prominent
below; intercostal venation slender, hardly elevated, densely scalariform; petiole 4–6 cm
long, stout, hardly geniculate. Inflorescences terminal or axillary; rachis compressed, to 11
cm long, doubly branched, branchlets bearing to 5 flowers; bracteoles ovate, acute, to 3 × 2
mm. Flowers: buds to 6 × 4 mm; stamens 15, connectival appendage 3–4x as long as
anther; ovary ovoid, without distinct stylopodium, style as long as ovary, filiform. Fruits:
calyx glabrous, lobes unequal, 3 longer lobes to 16 × 3.5 cm, tapering to c. 7 mm broad
above the to 2 × 1.5 cm saccate base, 2 shorter ones to 8 × 0.6 cm, otherwise similar. Nuts
ovoid, pruinose, to 3.3 × 1.8 cm, with c. 1.5 mm long tapering style remnant.
Vernacular names. Sarawak—kerukup (Malay), meranti kerukup ( preferred name).
Distribution. Endemic in Borneo. Widespread where its habitat occurs; unknown from
Sabah. In Sarawak recorded from Bintulu, Kuching, Lundu, Marudi, Miri, Samarahan, Sibu
and Tatau districts (e.g., FMS 7173, S 9316, S 11245, and S 11753). Also occurring in
304
Brunei (e.g., BRUN 695, BRUN 5531, FMS 30556, Niga NN 116, and S 7882) and
Ecology. Locally common, in mixed peat swamp forest and in kerangas forest on poorly
drained podsols. Endangered by forest conversion.
88. Shorea palembanica Miq.

(of Palembang, Sumatra)
(sect. Brachyptera, red meranti)
Fl. Ned. Ind., Suppl. (1861) 487; Symington op. cit. (1933) 141, op. cit. (1943) 83; Masamune op. cit.
496; Browne op. cit. 141; Ashton op. cit. (1968) 112, op. cit. (1982) 513; Anderson op. cit. (1980)
128; PROSEA op. cit. 181; Newman et al. op. cit. (1996) 181. Type: Teijsmann s.n. (= Herb. Utrecht
no. 35907), Sumatra, Palembang (holotype U). Synonyms: Shorea aptera Burck op. cit. 210; S.
brachyptera F.Heim op. cit. (1891) 571; Pachychlamys beccarianus Dyer ex Brandis op. cit. 77; P.
brachypterus Dyer ex Brandis op. cit. 77.
Main canopy tree, rarely large, to 45 m tall, to 1.3 m diameter; bole often leaning or
misshapen, often mossy; crown dense, irregularly oblong to hemispherical; buttresses stout.
Bark dark greyish brown, overall smooth, at first hoopmarked, becoming irregularly
cracked and shallowly patchily flaky; inner bark pinkish brown to cream at cambium;
heartwood pale pink, fairly soft. Young twig, inflorescence, leaf bud, parts of perianth
exposed in bud, stipule and bracteole outside, ovary and nut, and midrib above densely
evenly pale buffpuberulent; stipule and bracteole inside, petiole and leaf venation below
sparsely so, more or less glabrescent. Twigs compressed, c. 2 mm diameter apically. Leaf
buds lanceolatefalcate, to 13 × 6 mm. Stipules lanceolate, to 15 × 3 mm, caducous. Leaves
chartaceous, drying dark chocolatebrown below, purplish brown above, undulating; blade
ovate to oblong, 8–25 × 4–10 cm, base broadly cuneate to subcordate, apex with broad
acumen to 1 cm long; midrib slender but evident, shallowly furrowed or flat above,
prominent below; lateral veins 12–16 pairs, slender but prominent below; intercostal
venation scalariform, slender, dense; petiole 1.4–2.5 cm long. Inflorescences terminal or
axillary; rachis terete, doubly branched, branchlets bearing to 10 flowers; bracteoles ovate,
acute, to 4 × 2 mm, caducous. Flowers: buds to 7 × 3 mm; petals pale yellow; stamens 15,
connectival appendage c. 2x as long as anther; ovary ovoid, without stylopodium, style c. 2x
as long as ovary, filiform, pubescent in the basal quarter. Fruits: pedicel to 2 mm long;
calyx lobes unequal but short, 3 longer lobes lorate, to 5 × 0.7 cm above the saccate base, 2
shorter ones to 3 cm long, otherwise similar. Nuts ovoid, to 3.5 × 2.5 cm, apiculate.
Vernacular name. Sarawak—engkabang asu (preferred name).
Distribution. E Sumatra, Peninsular Malaysia and Borneo. In Sabah known from Beluran,
Sandakan, Semporna, and Tawau districts (e.g., SAN 9264, SAN 24288 and SAN 32503) and
in Sarawak from Belaga, Bintulu, Kapit, Kuching, Lundu, Marudi, Miri, Samarahan, Sibu,
Simunjan, and Tatau districts (e.g., S 15587, S 22289, S 23384, S 53529, and S 76331). Also
occurring in Kalimantan (e.g., bb. 29273, Endert 4935 and Kostermans 12686). Vulnerable
owing to forest conversion.
305
Ecology. Locally frequent on silty floodplains and hanging over the banks of sluggish
rivers. Occurring in Kubah NP; elsewhere vulnerable owing to land conversion.
Uses. A minor source of illipe nuts.
89. Shorea pallidifolia P.S.Ashton

(Latin, pallidus = pale, folius = leaf; the pale leaf undersurface)
Newman et al. op. cit. (1986) 182. Type: Rashid S 16054, Borneo, Sarawak, Bako NP (holotype K;
isotypes KEP, L).
Canopy tree, to 40 m tall, to 80 cm diameter; bole cylindrical, short; crown conical to

hemispherical, pale from below, dense; buttresses to 1.5 m long, stout. Bark pale pinkish to
greyish brown, becoming shallowly vsection fissured; inner bark pale yellowish brown,
faintly laminated; heartwood pinkish brown, quite hard. Twig, leaf bud, stipule,
inflorescence, parts of calyx exposed in bud, petiole, and venation below persistently pale
yellowish brown scabridpubescent; outside of petal and nut evenly creambuff pubescent.
Twigs compressed and ribbed at first, stout, c. 6 × 3 mm apically. Leaf buds broadly ovoid
conical, compressed, to 10 × 8 mm. Stipules oblong, obtuse, to 7 × 5 mm. Leaves thickly
coriaceous, cream lepidote below (mature trees); blade broadly oblong to ovate or obovate,
13–22 × 7–13 cm, base obtuse or subcordate, apex with short acumen; midrib obscure and
sunken above, prominent below; lateral veins 12–19 pairs, prominent below, arched;
intercostal venation densely scalariform, hardly elevated below; petiole stout, 2–3.5 cm
long. Inflorescences axillary; rachis terete, shortly singly branched, branchlets bearing to 4
flowers. Flowers: buds to 6 × 4 mm; petals pinkish yellow; stamens 15; ovary ovoid,
without stylopodium, glabrous, with somewhat longer columnar glabrous style. Fruits:
pedicels c. 1 mm long; calyx lobes unequal, 3 longer lobes to 7 × 1.5 cm, tapering to c. 7
mm above the saccate base, 2 shorter ones linearlobed, to 4 × 0.3 cm, similar but smaller
at base. Nuts ovoid, to 1.2 × 0.9 cm, shortly apiculate.
Distribution. Endemic in Borneo. Known in Sarawak from Kuching, Lundu and Serian
districts (e.g., KEP 79314, S 6161, S 9315, S 13357, and S 22768); also observed but not
collected in the low hills between the lower Mukah and Balingian rivers. Also occurring in
W Kalimantan north of Sg. Kapuas.
Ecology. Very local, rare overall, in kerangas on white sand terraces and over sandstone.
Locally frequent in Bako NP; endangered elsewhere.
90. Shorea parvifolia Dyer

(Latin, parvus = small, folius = leaf)
Fl. Br. Ind. 1 (1874) 306; Slooten op. cit. (1929) 203; Symington op. cit. (1933) 137, op. cit. (1943)
85; Masamune op. cit. 496; Browne op. cit. 141; Ashton op. cit. (1963) 277, op. cit. (1964) 206, op.
cit. (1968) 113, op. cit. (1982) 546; Meijer & Wood op. cit. 128; Burgess op. cit. 155, 183; Anderson
306
op. cit. (1980) 128; PROSEA op. cit. 399; Kessler & Sidiyasa op. cit. 107; Newman et al. op. cit.
(1996) 183. Type: Maingay 1577 (= Kew Distr. No. 206), Peninsular Malaysia, Malacca (holotype K;
isotype L). Synonyms: Shorea scutulata King op. cit. 110; S. gentilis Parijs op. cit. 244.
Large emergent tree, to 65 m tall, to 2 m diameter; bole tall, cylindrical; crown immense,
cauliflowershaped; buttresses to 4 m tall, stout. Bark at first remaining smooth, dappled
dark brown mottled with grey, becoming shallowly or deeply vsection fissured, chocolate
brown, eventually crumbly flaky towards the base; inner bark rich reddish brown;
heartwood deep pink to reddish brown; dammar smears cream, abundant. Young twig,
inflorescence, leaf bud, parts of perianth exposed in bud, stipule, bracteole, petiole and
midrib above evenly pale brown puberulent, persistent or glabrous; ovary and nut buff
pubescent. Twigs terete, c. 2 mm diameter apically. Leaf buds ovoid, obtuse, to 3 × 2 mm.
Stipules broadly ovate, obtuse, to 6 × 4 mm, caducous. Leaves thinly coriaceous, drying
chocolatebrown below, mauvebrown above, veins glabrous or sparsely scabridpubescent
below; blade broadly ovate, ovate, or elliptic, 5–11 × 2.5–6 cm, base obtuse, cordate, or
cuneate, sometimes with one or more pairs of opposite glabrous pale pit or scalelike basal
domatia, apex with acumen to 1 cm long; midrib obscure, sunken above, slender but
prominent below; lateral veins 10–13 pairs, slender, hardly elevated or prominent below;
intercostal venation slender, densely scalariform, hardly elevated; petiole 1–1.5 cm long.
Inflorescences terminal or axillary; rachis slender, terete or slightly compressed, to 12 cm
long, branchlets short, bearing to 8 dense flowers; bracteole ellipticoblong, obtuse, to 6 × 3
mm. Flowers: buds to 7 × 5 mm; petals cream suffused with pink at base; stamens 15,
connectival appendage short, slender, becoming reflexed; ovary and stylopodium ovoid
conical, surmounted by short glabrous style. Fruits: calyx lobes unequal, 3 longer lobes
chartaceous, to 9 × 1.5 cm, tapering to c. 4 mm above the saccate base, 2 shorter ones
linearlobed, to 3.5 × 0.2 cm, similar at base. Nuts ovoid, to 1.4 × 0.7 cm, with to 2.5 mm
tapering style remnant.
Vernacular names. Sabah—seraya punai (preferred name). Sarawak—meranti sarang

punai (preferred name).
Distribution. Peninsular Thailand (Pattani), Peninsular Malaysia, Sumatra, and Borneo.
Ecology. Common where it occurs, in mixed dipterocarp forest on yellow clay soils, usually
welldrained, on shales and intermediate and acid volcanic rocks, but rare on base rich rocks
such as basalt.
Notes. Two subspecies, viz. subsp. parvifolia and subsp. velutina, are recognised.
Key to subspecies
Bark becoming shallowly vsection fissured. Leaf blade broadly ovate, base obtuse or
cordate, margin slightly revolute; veins glabrous and hardly elevated below………………...
subsp. parvifolia
Sumatra, Peninsular Malaysia and Borneo. In Borneo occurring throughout the uplands,
but confined to Tawau district in Sabah (e.g., Elmer 21401 and Elmer 21530) and
Belaga and Kapit districts in Sarawak (e.g., S 29354 and S 29509). Vulnerable owing to
land conversion.
307
Bark becoming deeply vsection fissured. Leaf blade ovate or elliptic, base obtuse or
cuneate, margin frequently narrowly revolute; veins sparsely scabridpubescent and
prominent below……………………………………………………………………………….
subsp. velutinata P.S.Ashton
(Latin, velutinatus = somewhat velvety; the indumentum)
Gard. Bull. Sing. 20 (1963) 278, op. cit. (1982) 547; Newman et al. op. cit. (1996) 185. Type:
Awang Lela FMS 4502, Peninsular Malaysia, Pahang, Temerloh, Belingo FR (holotype KEP).
E Sumatra, Peninsular Malaysia east of the main range (known there as the “Pahang
Form”) and Borneo. In Borneo, widespread, and the prevailing form in Sabah (e.g.,
SAN 21627 and SAN 36950) and Sarawak (e.g., S 9612 and S 29673).
With S. leprosula the commonest red meranti in the mixed dipterocarp forest; with a
similar ecological range and cooccurring with that species, but tending to be prevalent
on slightly more leached soils, particularly in the lowlands near to the coast. In Sabah,
common in Danum Valley Conservation Area and Sepilok FR and occurring in
Kinabalu NP, and in Sarawak, common in G. Gading, Lambir and Mulu NPs; not
vulnerable.
91. Shorea parvistipulata F.Heim

(Latin, parvus = small, stipulatus = with stipules; the small stipules)
Bull. Mens. Soc. Linn. Paris 2 (1891) 974; Merrill op. cit. (1921) 406 (‘parvistipula’); Masamune op.
cit. (‘parvistipula’); Meijer & Wood op. cit. 132; Burgess op. cit. 184; Ashton op. cit. (1968) 114, op.
cit. (1978) 46, op. cit. (1982) 508; Anderson op. cit. (1980) 128; Kessler & Sidiyasa op. cit. 108;
Coode et al. (eds.) op. cit. 81; Newman et al. op. cit. (1996) 185. Type: Beccari PB 2547, Borneo,
Sarawak, Matang (holotype P). Synonyms: Shorea cristata Brandis op. cit. 97, Ashton op. cit. (1964)
184, op. cit. (1968) 106, Meijer & Wood op. cit. 102, Burgess op. cit. 166, Anderson op. cit. (1980)
125; S. nebulosa Meijer op. cit. 337, Meijer & Wood op. cit. 123, Burgess op. cit. 183.
Huge emergent tree, to 70 m tall, to 2 m diameter; bole tall, straight, cylindrical; crown vast,
diffuse but regularly hemispherical; buttresses to 5 m tall, prominent, stout. Bark mauve
brown, at first smooth, becoming shallowly cracked and flaky, eventually scalloped,
prominently pale lenticellate; inner bark pale chocolatebrown; heartwood pink; dammar
smears occasional, cream. Leaf bud, parts of perianth exposed in bud, twig, inflorescence,
stipule and bracteole outside, petiole and leaf venation below persistently pale yellowish to
greyish brown scabridpubescent; stipule and bracteole inside and midrib above evenly so.
Twigs terete, c. 2 mm diameter apically; stipule scars pale, short, horizontal or ascending.
Leaf buds ovoid, acute, to 6 × 4 mm. Stipules ovate, subacute, to 14 × 7 mm. Leaves
chartaceous, drying rust to chocolatebrown below, mauvebrown above; blade variable,
more or less oblong, base cordate, sometimes obtuse or cuneate, apex with tapering acumen
to 1 cm long; midrib evident, flat or shallowly furrowed above, prominent below; lateral
veins 13–21 pairs, prominent below; intercostal venation scalariform, wellspaced; petiole
1.2–1.5 cm long. Inflorescences terminal or axillary; rachis terete, straight, lax, to 16 cm
long, branchlets to 4 cm long bearing to 11 flowers; bracteoles deltoid, acute, to 5 × 3 mm,
not at first caducous. Flowers: buds to 7 × 3 mm; petals cream suffused with pink at base;
stamens 15, connectival appendage c. 2x the length of anther; ovary ovoid, stylopodium
obscure, style equal to ovary, filiform, glabrous but for the base. Fruits: pedicels c. 2 mm
long; calyx lobes unequal, 3 longer lobes loratespatulate, to 20 × 2.5 cm, tapering to 11
mm above the saccate base, 2 shorter ones linearlobed, to 8 × 0.4 cm, similar at base.
Nuts ovoid, to 2.5 × 2 cm, often with persistent filiform style remnant.
308
Vernacular names. Sabah—kawang daun merah (Malay), seraya lupa (preferred name).
Sarawak—meranti kawang pinang (preferred name).
Ecology. Scattered in mixed and upper dipterocarp forest, particularly on hillsides, at

altitudes to 1300 m.
Notes. Three subspecies, viz. subsp. albifolia, subsp. nebulosa and subsp. parvistipulata, are
recognised, all occurring in Sabah and Sarawak.
Key to subspecies
1. Leaf blade silvery pink lepidote below…………………………………………………...

subsp. albifolia P.S.Ashton
(Latin, albus = white, folium = leaf; the silvery pink leaf undersurface)
Gard. Bull. Sing. 31 (1978) 46. Type: Wright S 29276, Borneo, Sarawak, Niah, Sg. Sekaloh
Occur in widely separated localities from Niah, NE Sarawak (e.g., the type), to
Brunei (e.g., BRUN 5630, S 1689 and S 1692) and SE and W Kalimantan (e.g., bb.
30193, bb. 34918 and Latupeirissa 95003). Locally common on periodically
flooded alluvium, and adjacent moist hillsides, on silty clay soil. Vulnerable owing
to suitability of its habitat for agriculture.
Leaf blade persistently pale yellowish to greyish brown scabridpubescent on the veins
below………….…………………………………………………………………………2
Tree at most 45 m tall, 1.2 m diameter. Leaf blade to 13 × 6 cm; lateral veins to 15 pairs.
Larger fruit calyx lobes to 9 × 1.8 cm……………………………………………...
subsp. nebulosa (Meijer) P.S.Ashton
(Latin, nebulosus = clouded with unevenly blended colours: the difficulty of
identification)
Gard. Bull. Sing. 31 (1978) 46; Newman et al. op. cit. (1996) 187. Basionym: Shorea
nebulosa Meijer op. cit. 337; Meijer & Wood op. cit. 123; Burgess op. cit. 183. Lectotype
(Ashton, 1978): G.H.S. Wood & Charrington SAN 16355, Borneo, Sabah, Ranau district,
Poring (hololectotype L).
Endemic in Borneo; recorded from Ranau, Sipitang, Tambunan, and Tawau
districts in Sabah (e.g., SAN 16273, SAN 16357, SAN 16435, SAN 16769, SAN
17010, and SAN 31438).
Locally frequent in upper dipterocarp forest, at 800–1300 m altitude. Occurring in
Kinabalu NP; elsewhere vulnerable.
Tree to 70 m tall, 2 m diameter. Leaf blade 6–20 × 3–9 cm; lateral veins 13–21 pairs.
Larger fruit calyx lobes to 20 × 2.5 cm…………………………………………………...
subsp. parvistipulata
Occurring throughout Borneo, except SW Kalimantan. In Sabah recorded from
Beluran, Beaufort, Kota Belud, Kota Merudu, Kudat, Sipitang, Tawau, and Tenom
districts (e.g., SAN 15108, SAN 17180, SAN 25601, SAN 43043, and SAN 99603)
and in Sarawak from Bau, Belaga, Kapit, Kuching, Lawas, Limbang, Lundu,
Marudi, Miri, and Tatau districts (e.g., S 15388, S 27550, SAN 29596, S 32376, and
S 49961). Also occurring in Brunei (e.g., BRUN 2005 and BRUN 3382) and E, C
and W Kalimantan (e.g., Ambriansyah & Arifin Berau 1045, bb. 29625, bb. 35337,
309
Church et al. 1660, and Laman et al. TL 1138). Scattered in mixed dipterocarp
forest on clay soils, particular on hillsides, most common on intermediate and basic
igneous rocks and porphyry dikes near limestone hills, at altitudes to 1200 m.
Occurring in G. Gading and Mulu NPs; not vulnerable.
92. Shorea patoiensis P.S.Ashton

(of Bt. Patoi, Brunei)
Wood op. cit. 78; Burgess op. cit. 218; Anderson op. cit. (1980) 123; Kessler & Sidiyasa op. cit. 109;
Coode et al. (eds.) op. cit. 81; Newman et al. op. cit. (1996) 188. Type: Ashton BRUN 3324, Borneo,
Brunei, Bt. Patoi (holotype K; isotype KEP).
Main canopy or low emergent tree, to 45 m tall, to 1 m diameter; crown hemispherical; bole
straight, cylindrical; buttresses to 2 m tall but usually less, somewhat slender. Bark tawny
brown, oblong flaky. Inflorescence and parts of perianth exposed in bud shortly densely
greyish puberulent; ovary and stylopodium cream puberulent; nut glabrescent; vegetative
parts glabrous but for fimbriate bud scales and stipules, and more or less waxy pruinose
young buds and twigs (mature trees). Twigs terete, smooth or rugulose, muchbranched,
very slender, c. 1 mm diameter apically. Leaf buds narrowly ovoid, c. 3 × 1 mm (to 4 × 2
mm in juveniles), with the bud scales typically somewhat patent. Stipules oblong, acute, c.
8 × 2 mm. Leaves more or less chartaceous, drying greyish green and curling; blade ovate,
5–8 × 2–3.5 cm, base cuneate, decurrent for c. 1 mm along the petiole, apex with slender
acumen to 1 cm long; midrib evident, flat or sightly elevated within a shallow groove above,
elevated but not prominent below; lateral veins 7–9 pairs, slender, elevated above, more so
below, arched; intercostal venation subscalariform, hardly elevated; petiole 0.7–1 cm long.
Inflorescences terminal or axillary; rachis very slender, terete, to 9 mm diameter, branchlets
bearing to 6 secund flowers; bracteoles minute, fugaceous. Flowers: buds small, to 3.5 ×
1.5 mm; corolla lemon yellow; stamens 15, connectival appendage equal to anther, ciliate
towards apex; ovary ovoid, with cylindrical stylopodium, style short, glabrous. Fruits: calyx
lobes subequal, shorter than nut, lobes broadly ovate, to 0.5 × 0.5 cm, appressed to nut and
united into a 0.5–0.9 × 0.4–0.5 cm receptacle at base. Nuts oblongellipsoid, to 1.8 × 1 cm,
with minute style remnant, subglabrous, striated.
Vernacular name. Sarawak—lun hitam padi (preferred name).
Distribution. Endemic in Borneo. Known in Sabah from Beaufort, Lahad Datu, Sandakan,
and Tawau districts (e.g., SAN 16462, SAN 21498, SAN 23556, SAN 31475, and SAN 63055)
and in Sarawak from Belaga, Bintulu, Kapit, Limbang, Miri and Tatau districts (e.g., S
3324, BRUN 5710, BRUN 5811, and Johns 7474) and E Kalimantan (e.g., Ambriansyah
Berau 885, bb. 18967 and Kostermans 13850).
310
Ecology. Frequent on moist wellstructured clay soils, especially on hill sides, over shale
and especially basic volcanic rocks; abundant for instance on the Bt. Mersing, Sarawak,
basalt; also on the Hose Mountain and Usun Apau dacite. Common in Lambir and Mulu
NPs; probably not vulnerable.
93. Shorea pauciflora King

(Latin, paucus = few, flos = flower; a supposed fewflowered inflorescence)
J. As. Soc. Beng. 62, 2, (1893) 116; Merrill op. cit. (1929) 203; Keith op. cit. 25; Masamune op. cit.
496; Symington op. cit. (1943) 87; Browne op. cit. 149; Ashton op. cit. (1964) 207, op. cit. (1968)
114, op. cit. (1978) 47, op. cit. (1982) 511; Meijer & Wood op. cit. 133; Burgess op. cit. 165;
Anderson op. cit. (1980) 128; PROSEA op. cit. 400; Kessler &. Sidiyasa op. cit. 109; Coode et al.
(eds.) op. cit. 114; Newman et al. op. cit. (1996) 189. Type: Curtis 1537, Peninsular Malaysia, Penang
Hill (holotype CAL; isotype K).
Huge emergent tree, to 60 m tall, to 2.2 m diameter; crown eventually immense,

cauliflowershaped; bole straight, tall, cylindrical; buttresses to 4 m tall, stout. Bark dark
purplish brown eventually fawnbrown, shallowly densely cracked becoming thinly oblong
flaky, flakes at first persisting and curling up, surface overall eventually appearing rather
smooth; dammar smears greyish brown; inner bark meatred; heartwood dark reddish
brown. Leaf bud, parts of perianth exposed in bud, inflorescence, stipule, and bracteole
persistently evenly golden brown pubescent; venation below, petiole, fruit calyx, and young
twig sparsely caducously so; ovary and nut buff pubescent. Twigs terete, 1.5–2.5 mm
diameter apically, becoming flaky; stipule scars pale, short, horizontal. Leaf buds ovoid,
acute, 4–8 × 2–3.5 mm. Stipules hastate, to 13 × 4 mm. Leaves thinly coriaceous, drying
brickred or purplish brown below; blade ovate, 9–15 × 4–5.5 cm, base frequently
subequal, obtuse to broadly cuneate, apex with slender acumen 0.6–1.2 cm long; midrib
obscurely sunken above, prominent below; lateral veins 8–9 pairs, slender but prominent
below, generally drying paler than blade, ascending and arched, sometimes with small
porelike domatia; intercostal venation densely scalariform, very slender, hardly raised;
petiole 1.3–1.8 cm long. Inflorescences terminal or axillary; rachis ribbed or slightly
compressed, lax, to 15 cm long, to trebly branched, branchlets bearing to 20 secund flowers;
bracteoles oblong, subacute, to 4 × 2.5 mm. Flowers: petals pale yellow; stamens 15,
connectival appendage c. 2x the length of anther, sericeous distally; ovary and stylopodium
ovoidconical, style slightly shorter than both, glabrous. Fruits: calyx lobes unequal, 3
longer lobes to 9 × 1.5 cm, tapering to c. 4 mm above the saccate base, 2 shorter ones
linearlobed, to 5 × 0.5 cm, similar at base. Nuts broadly ovoid, c. 1.4 × 1.2 cm, with short
apical style remnant.
Vernacular names. Sabah—oba suluk (preferred name). Sarawak—nemesu (preferred

name), perawan samak (Iban).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo widespread, recorded in

Sabah from Beaufort, Kinabatangan, Kuala Penyu, Lahad Datu, Sandakan, Sipitang, Tawau,
134945) and in Sarawak from Bau, Belaga, Bintulu, Kapit, Kuching, Lundu, Samarahan,
and Sri Aman districts (e.g., S 10183, S 22884, S 29242, S 36907, S 68724). Also occurring
in Brunei (e.g., BRUN 3374 and BRUN 5085) and Kalimantan (e.g., bb. 34256 and
Kostermans 6042).
311
Ecology. Ubiquitous, locally common, usually as scattered trees, in mixed dipterocarp forest
on yellowred sandy, sandy clay and clay soils, including on the plateau sandstone and
granodiorite of W Sarawak, the Arip rhyolite, and the inland sedimentaries. One of our most
widespread dipterocarps. Occurring in Bako, G. Gading, Kinabalu, Kubah, Lambir, and
Mulu NPs, and Danum Valley Conservation Area and Sepilok FR; not vulnerable.
94. Shorea peltata Symington

(Latin, peltatus = shieldshaped, the leaf)
(sect. Richetioides, subsection Richetioides, yellow meranti)
J. Malay. Br. Roy. As. Soc. 19 (1941) 158, op. cit (1943) 56; Slooten op. cit. (1956) 337; Ashton op.
cit. (1982) 473. Type: Abdul Jaffar KEP 49356, Peninsular Malaysia, Johor, Jemaluang FR (holotype
KEP).
Subcanopy or canopy hardly buttressed tree, to 40 m tall, 30 cm diameter. Bark smooth or

shallowly nobbly. Inflorescences petals outside and nut persistently densely buff pubescent,
caducous on calyx outside; parts otherwise glabrescent. Twigs terete, smooth, c. 2 mm
diameter apically. Leaf buds ovoidacute, minute. Stipules unknown. Leaves deeply peltate,
chartaceous to thinly coriaceous; blade oblong, 8–16 × 4–7.5 cm, apex with acumen to 1
cm long; midrib evident but slightly furrowed above, raised below; lateral veins 8–9 pairs,
the first 3 of which opposite and radiating from the petiole insertion; intercostal venation
subreticulate, elevated on both surfaces; petiole 1.8–3 cm long, slender. Inflorescence
terminal or axillary; branchlets to 1 cm long bearing to 5 flowers. Flower: buds lanceolate,
to 6 mm long; stamens 15, in 3 whorls, connectival appendage c. 1½x the length of anther,
slender, scabrous distally; ovary ovoid sericeous distally, without stylopodium, style
glabrous, columnar, equal to ovary. Fruits: subsessile; calyx lobes short, subequal, ovate,
tuberculateincrassate, to 1 × 0.8 cm. Nuts obovoid, to 3 × 2 cm, subacute.
Vernacular name. Sabah—seraya kuning keladi (preferred name).
Distribution. Sumatra, Peninsular Malaysia (SE Johore) and Borneo. In Borneo known
only from Labuk Sugut district, E Sabah (e.g., SAN 70329 and SAN 131964).
Ecology. Very local, in mixed lowland dipterocarp forerst on leached clay soil. Critically
endangered.
95. Shorea pilosa P.S.Ashton

(Latin, pilosus = long hairy; the indumentum)
Newman et al. op. cit. (1996) 191. Type: G.H.S. Wood SAN 16721, Borneo, Sabah, Sipitang district,
Ulu Mendalong (holotype K; isotypes KEP, L).
312
Mediumsized emergent tree, to 45 m tall, to 1.3 m diameter; crown golden suffused from
below, dense, hemispherical; bole straight, cylindrical; buttresses occasionally to 4 m tall,
stout. Bark pale yellowish brown, smooth, hoopmarked, eventually patchily cracked,
flaking leaving scrollmarked surfaces; inner bark pale coffeecoloured; heartwood pale
pink, fairly soft. Twig, leaf bud, inflorescence, parts of calyx exposed in bud, petiole, midrib
on both surfaces, and blade below persistently golden scabridtomentose; blade above and
fruit calyx sparsely pubescent, glabrescent; part of corolla exposed in bud and nut evenly
buff pubescent. Twigs slightly compressed at apex, muchbranched, 2–2.5 mm diameter
apically. Leaf buds ovoid, 3–7 × 1.5–3 mm. Stipules hastate, to 35 × 15 mm. Leaves thinly
coriaceous, drying goldentawny below, pinkish brown above; blade ovateelliptic, 10–17 ×
4–7.5 cm, base obtuse, margin usually narrowly revolute, apex with slender acumen to 1.3
cm long; midrib evident, slender but furrowed above, prominent below; lateral veins 12–15
pairs, arched, dense, with prominent axillary tufted tomentose domatia; intercostal venation
densely scalariform with reticulate finer veins, minutely raised below; petiole 1–1.3 cm
long. Inflorescences terminal or axillary, singly branched; rachis terete, to 14 cm long.
Flowers: buds to 8 × 3 mm; petals cream suffused with pink; stamens 15, connectival
appendage c. 4x as long as anther; ovary ovoid, glabrous, style and stylopodium spindle
shaped. Fruits: calyx lobes unequal, 3 longer lobes to 17 × 2.5 cm, tapering to c. 13 mm
above the saccate base, 2 shorter ones to 12 × 1 cm, otherwise similar. Nuts ovoid, to 2 ×
1.8 cm, with short acute style remnant.
Vernacular names. Sabah—kawang bulu (preferred name). Sarawak—kawang bulu (Iban),

meranti bulu (Malay).
Distribution. Endemic in Borneo. In Sabah known from Beaufort, Sipitang, Tawau, and
Tenom districts (e.g., SAN 24808, SAN 35890 and SAN A 1748) and in Sarawak from
Bintulu, Kapit, Lawas, Miri, and Tatau districts (e.g., S 1752, S 17792, S 18429, S 22235,
and S 25018). Also occurring in Brunei (e.g., BRUN 2636 and S 5777).
Ecology. Rare, local, in mixed dipterocarp forest on lowers slopes and undulating land, on
sandy clay and wellstructured clay soils, including the Bt. Mersing, Anap basalt. Occurring
in Lambir and Mulu NPs; elsewhere vulnerable owing to land conversion.
96. Shorea pinanga Scheff.

(from an indigenous name—pinang)
Nat. Tijd. Ned. Ind. 31 (1870) 350; Beccari op. cit. 570; Merrill op. cit. (1921) 406, op. cit. (1929)
203; Masamune op. cit. 496; Ashton op. cit. (1963) 281, op. cit. (1964) 210, op. cit. (1968) 115, op.
PROSEA op. cit. 400; Kessler & Sidiyasa op. cit. 109; Coode et al. (eds.) op. cit. 81; Newman et al.
op. cit. (1996) 192. Lectotype (designated here): s.c. s.n., cult. in Hort. Bogor (hololectotype BO;
isolectotypes L, U). Synonyms: Shorea gysbertsiana Burck var. scabra Burck op. cit. (1886) 17, p.p.;
S. compressa Burck op. cit. (1886) 26.
Mediumsized emergent tree, to 45 m tall, to 1.3 m diameter; bole straight, cylindrical;

crown loosely domeshaped to oblong, the branch endings pendent; buttresses to 1.5 m tall,
stout. Bark pale pinkish brown, smooth, hoopmarked, eventually becoming patchily
cracked and irregularly flaked; inner bark pale pinkish brown; heartwood pale pink,
relatively soft. Young twig, inflorescence, leaf bud, parts of calyx exposed in bud, stipule,
313
bracteole, petiole and sometimes blade below glabrous or fugaceously to persistently

golden brown pubescent; nut yellowish buff pubescent. Twigs compressed, 4–6 × 2–3 mm
apically; stipule scars prominent, falcate, descending. Leaf buds hastate, subacute, usually
hidden in subpersistent stipules, to 10 × 4 mm. Stipules hastate, to 10 × 4 mm,
subpersistent. Leaves thinly coriaceous, drying shiny coppery brown below (unless
tomentose); blade elliptic to narrowly ovate, 11–24 × 4–9 cm, base broadly cuneate to
subcordate, apex with tapering acumen to 1.5 cm long; midrib slender but evident, flat or
shallowly furrowed above, prominent below; lateral veins 10–20 pairs, slender but raised
below, arched; intercostal venation densely scalariform, hardly raised; petiole 1.5–2.3 cm
long, slender. Inflorescences terminal or axillary; rachis lax, compressed, to 24 cm long,
singly branched, branchlets bearing to 15 flowers; bracteoles broadly ovate, to 10 × 8 mm,
subpersistent. Flowers: petals deep pink; stamens 15, connectival appendage many times
longer than anther; ovary ovoid, stylopodium obscure, style twice the length of ovary,
columnar. Fruits: calyx lobes unequal, 3 longer lobes coriaceous, to 23 × 3.5 cm, tapering
to c. 13 mm above the saccate base, 2 shorter ones linearlobed, to 17 × 1.2 cm, similar at
base. Nuts broadly ovoid, to 2.3 × 2.3 cm, with 2 mm slender style remnant.
Vernacular names. Sabah—kawang pinang (preferred name). Sarawak—engkabang

langgai bukit (Iban), meranti langgai bukit (preferred name).
Distribution. Endemic in Borneo, occurring throughout the island. In Sabah recorded from
Beaufort, Keningau, Kinabatangan, Labuk Sugut, Ranau, Sandakan, Sipitang, and Tawau
Sarawak from Belaga, Bintulu, Kapit, Kuching, Lawas, Limbang, Miri, Samarahan, Serian,
and Sibu districts (e.g., S 14740, S 16429, S 22250, S 29060, and S 43616). Also occurring
in Brunei (e.g., BRUN 3344 and FMS 35682), and throughout Kalimantan (e.g., bb. 29665,
Kessler et al. Berau 704 and Meijer 2206).
Ecology. Locally common, in mixed dipterocarp forest on leached clay and sandy clay soils,
especially on ridges and low hills. Occurring in Mulu NP; probably not vulnerable.
Notes. A variable species, distinguished by its falcate stipule scar, many slender veins and
very long fruit sepals. Typically, the vegetative parts are glabrous but one form, with stout
twig and larger than normal leaf, is pubescent on twig and leaf below; in E Sabah the leaf is
very large, with markedly cuneate base, and shares characteristics with S. macrophylla (see
there).
97. Shorea platycarpa F.Heim

(Greek, platy = flat, karpos = fruit; perhaps referring to the fruit calyx)
(sect. Mutica subsect. Mutica, red meranti)
Bull. Mens. Soc. Linn. Paris 2 (1891) 956; Beccari op. cit. 510; Merrill op. cit. (1921) 406; Symington
op. cit. (1933) 133, op. cit. (1943) 88; Masamune op. cit. 496; Browne op. cit. 142; Anderson op. cit.
(1963) 159, op. cit. (1980) 128; Ashton op. cit. (1964) 211, op. cit. (1968) 115, op. cit. (1982) 541;
Meijer & Wood op. cit. 138; Burgess op. cit. 154; PROSEA op. cit. 400; Coode et al. (eds.) op. cit. 81;
Newman et al. op. cit. (1998) 194. Type: Beccari PB 3302, Borneo, Sarawak, Marop (holotype P).
Synonym: Shorea palustris Ridl., op. cit. (1922) 224.
314
Emergent tree, to 50 m tall, to 1.2 m diameter; bole tall, straight; crown shallowly
hemispherical, diffuse; buttresses to 4 m tall, stout. Bark greyish to pinkish brown,
becoming deeply vsection fissured, eventually crumbly flaky; inner bark dull pinkish
brown; heartwood pale reddish brown, relatively soft; dammar as pale yellowish cream
smears. Young twig, inflorescence, leaf bud, parts of calyx exposed in bud, and petiole
densely persistently rustbrown scabrid pubescent; sparsely so on stipule, bracteole, midrib
above and blade below, and base of fruit calyx; glabrescent on blade above; petals outside
and nut densely buff pubescent. Twigs ridged, verrucose, stout, c. 3 mm diameter apically.
Leaf buds broadly ovoid, obtuse, 4–6 × 3.5–5 mm. Stipules ovate, subacute, to 10 × 6 mm.
Leaves thinly coriaceous, drying reddish brown; blade ellipticoblong to broadly ovate, 9–
17 × 5.5–10 cm, base obtuse, apex with broad acumen to 1 cm long; midrib slender, more
or less flat above, prominent below, in juveniles furnished with pale scalelike domatia
along part of its length from the base; lateral veins 16–20 pairs, dense, straight until
approaching margin, slender but prominent, in juveniles with scalelike domatia; intercostal
venation laxly scalariform, slender but distinctly elevated; petiole 1.5–2 cm long, stout.
Inflorescences terminal or axillary; rachis rigid, ribbed, to 9 cm long; bracteoles broadly
ovate, acute, to 4 × 3 mm. Flowers: petals pale yellow; stamens 15, connectival appendage
as long as anther, becoming reflexed; ovary and stylopodium narrowly ovoid, subglabrous,
style as long as both, filiform, glabrous. Fruits subsessile; calyx lobes unequal, 3 longer
lobes chartaceous, to 6.5 × 1.2 cm, tapering to 4 mm above the saccate base, 2 shorter ones
linearlobed, to 2 cm long, similar at base. Nuts ovoid, to 0.8 × 0.6 cm, acute.
Vernacular name. Sarawak—meranti paya (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah known only from
Sipitang district (e.g., SAN 19001, SAN 27966 and SAN 130246) and in Sarawak from
Bintulu, Daro, Kuching, Sibu, Simunjan, and Sri Aman districts (e.g., S 261, S 452, S 7273,
S 12853, and S 13501). Also occurring in Brunei (e.g., BRUN 371 and FMS 28695) and
Kalimantan.
Ecology. Locally common, widespread, in mixed peat swamp forest, especially near the
coast, and on poorly drained podsols on the former beach terraces on the lower Medamit,
Ulu Limbang, Sarawak. Recorded from Mulu NP; elsewhere endangered by forest
conversion.
98. Shorea platyclados Slooten ex Foxw.

(Greek, platy = flat, clados = a shoot; the compressed twigs)
Malay. For Rec. 10 (1932) 214; Symington, J. Malay. Br. Roy. As. Soc. 14 (1936) 336, op. cit. (1939)
377, op. cit. (1943) 89; Slooten Bull. Jard. Bot. Buitenz. 3, 16 (1941) 110; Masamune op. cit. 496;
Ashton op. cit. (1964) 212, op. cit. (1968) 115, op. cit. (1982) 515; Meijer & Wood op. cit. 139;
81; Newman et al. op. cit. (1996) 195. Type: Symington KEP 24416, Peninsular Malaysia, Selangor,
Ulu Gombak (holotype KEP).
Large emergent tree, to 50 m tall, to 1.8 m diameter; bole tall, cylindrical, but also often a
small gnarled main canopy tree; crown large, hemispherical, eventually diffuse; buttresses
to 4 m tall, stout,. Bark dark warm chocolatebrown, deeply narrowly fissured and flaking;
dammar incrustations pale yellow, conspicuous on bole; inner bark reddish brown to
315
yellowish brown within; heartwood deep crimson red. Young parts pale grey sericeous,
fugaceous except on stipule, bracteole and panicle. Twigs compressed, glabrous, smooth, c.
3 × 1.5 mm apically. Leaf buds compressed, falcate, to 4 × 1.5 mm. Stipules hastate, to 13
× 3.5 mm. Leaves remaining distichous, thinly coraceous, satiny, drying dark chocolate
brown below; blade lanceolate, 6–9 × 2–3 cm, base cuneate owing to the revolute basal
margin, margin undulate, apex with slender acumen c. 1 cm long; midrib obscurely sunken
above, acute and sharply prominent below; lateral veins 15–18 pairs, very slender, hardly
elevated below, ascending, with short intermediate veins; intercostal venation densely
scalariform, very slender; petiole slender, laterally compressed, 1–1.5 cm long.
Inflorescences terminal or axillary; rachis compressed, slender, to 7 cm long, singly
branched, branchlets bearing to 7 flowers; bracteoles to 4 × 2 mm, elliptic, fugaceous.
Flowers: petals pale yellow; stamens 15, connectival appendage c. 4x the length of anther,
glabrous; ovary and stylopodium pyriform, sericeous towards apex, style as long as both,
glabrous. Fruits: calyx lobes unequal, 3 longer lobes to 10 × 1 cm, tapering to 4 mm above
the saccate base, 2 shorter ones linearlobed, to 5 × 0.4 cm, similar at base. Nuts ovoid, to
1.5 × 1 cm, acute, glabrescent.
Vernacular names. Sabah—seraya bukit (preferred name). Sarawak—meranti bukit

(preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. Throughout the mountain system
of Sabah and Sarawak, wherever the habitat occurs. In Sabah recorded from Keningau,
Ranau, Sipitang, and Tambunan districts (e.g., SAN 16581, SAN 17026, SAN 22091, SAN
69102, and SAN 132706) and in Sarawak from Kapit, Lawas and Limbang districts (e.g., S
12108, S 17754, S 22212, S 26549, and S 33073). Also occurring in Brunei (e.g., BRUN 410
and BRUN 2541) and SE and W Kalimantan (e.g., bb. 20666, bb. 30194 and Endert 3691).
Ecology. Locally common in upper dipterocarp forest, at 700–1300 m altitude, but only on
deep friable redbrown fertile clay soils; on shale and basalt. Occasional on similar soils in
valleys at the base of the mountains, at altitudes as low as 150 m. Occurring in Kinabalu and
Mulu NPs; elsewhere vulnerable owing to logging.
99. Shorea polyandra P.S.Ashton

(Greek, poly = many, andra = males; the many stamens)
(sect. Richetioides, subsect. Polyandrae, yellow meranti)
PROSEA op. cit. 420; Newman et al. op. cit. (1996) 196. Type: Bojeng S 10171, Borneo, Sarawak,
Lundu district, G. Gading FR (holotype K; isotype KEP).
Vast emergent tree, to 70 m tall, to 2 m diameter, with tall somewhat tapering bole and
immense cauliflowershaped crown pale from below; buttresses to 4 m tall, prominent,
somewhat stout. Bark becoming chocolatebrown, deeply cracked and thickly oblongflaky;
inner bark purplish brown, laminated. Twig, petiole and blade below persistently purplish
rufous lepidote fading to grey (mature trees); leaf bud, stipule, inflorescence, parts of
perianth exposed in bud, bracteole, ovary, style, and nut persistently shortly purplishrufous
pubescent; fruit calyx sparsely so towards base, glabrescent. Twigs terete, smooth or
rugulosestriated, slender, c. 1 mm diameter apically. Leaf buds ellipsoid, to 2 × 1 mm,
acute. Stipules lanceolate, to 5 × 2 mm, fugaceous. Leaves chartaceous, undulate, drying
316
purplish brown with the venation darker and curling; blade lanceolate, 8–13 × 3–5 cm,
base cuneate, margin minutely revolute, apex with slender acumen to 2 cm long; midrib and
veins narrowly sunken and obscure above, slender but distinctly raised below; lateral veins
11–14 pairs, arched, ascending; intercostal venation laxly scalariform, slender, more or
less obscure below; petiole 1.4–2 cm long, slender. Inflorescences terminal or axillary;
rachis terete, to 6 cm long, singly branched, branchlets bearing to 3 flowers; bracteoles
elliptic, subacute, to 4 × 3 mm, caducous. Flowers: buds broadly ovoid, to 5 × 4 mm; petals
oblong, obtuse; stamens 102–107, anthers narrowly oblong, length c. 4x breadth,
connectival appendage stout but filiform, tapering, somewhat shorter in length than anther,
densely ciliate apically, not reflexed; ovary ovoid, without stylopodium, surmounted by a
short broadly columnar trifid style. Fruits: pedicels c. 1 mm long; calyx lobes unequal, 3
longer lobes to 8 × 1.4 cm, tapering to 8 mm above the saccate tuberculate base, 2 shorter
ones linearlobed, to 5 × 0.5 cm, similar at base. Nuts narrowly ovoid, to 3 × 1.3 cm, acute.
Vernacular name. Sabah—seraya kuning Quoin (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from Tawau district (e.g., SAN 15265,
SAN 16478, SAN 63027, SAN A 3465, and SAN A 3959) and in Sarawak from Lundu and
Miri districts (e.g., S 7979, S 10174, S 10189, S 25298, and S 75394). Also occurring in SE
Kalimantan (e.g., bb. 23349 and Kostermans 13302).
Ecology. In clusters of immense trees, on wellstructured and wellwatered fertile clay

loams overlying calcareous shale, basic volcanic rock and granodiorite. Well represented in
G. Gading NP and occurring in Lambir NP; elsewhere endangered by land conversion.
100. Shorea praestans P.S.Ashton

(Latin, praestans = preeminent; the striking stipules and leaf shape)
Newman et al. op. cit. (1996) 198. Type: Brunig s.n., Borneo, Sarawak, Bintulu district, Nyabau FR
(holotype K).
Subcanopy, possibly canopy tree, to 30 m tall, to 15 cm diameter. Bark smooth, hoop

marked, ochreousbrown and greymottled; inner bark pinkish brown. All observed parts
glabrous. Twigs compressed, smooth, stout, c. 5 × 3 mm apically. Leaf buds falcate
lanceolate, to 25 × 7 mm. Stipules elliptic, subacute, very large, to 110 × 50 mm, not at first
caducous. Leaves thickly coriaceous, drying greyish brown above, reddish brown shiny
below; blade broadly oblong, 24–35 × 11–18 cm, base cordate, apex obtuse to shortly
broadly acuminate; midrib broad but flat above, prominent below; lateral veins 11–13
pairs, prominent below; intercostal venation remotely subscalariform, slightly elevated
below; petiole stout, weakly geniculate, 4–6.5 cm long. Inflorescences and flowers
unknown. Fruits: pedicels to 5 mm long and stout; calyx lobes unequal, 3 longer lobes to
15 × 3 cm, tapering to c. 1.5 cm above the saccate base, 2 shorter ones to 10.5 × 1.5 cm,
similar at base. Nuts ovoid, to 2 × 2 cm, glabrous, with to 4 mm long style remnant.
317
Distribution. Endemic in Borneo (C Sarawak); known from Bintulu and Mukah districts
(e.g., S 19598, S 22074, S 22075, and S 37863).
Ecology. Rare and confined to deep yellow sandy soil on low hills of Neogene sandstone, in
mixed dipterocarp forest; as scattered trees or small groups. Possibly extinct, otherwise
critically endangered.
Notes. Central Sarawak has other distinctive, apparently small, trees representing
undescribed entities in sect. Pachycarpae, the main section of red merantis yielding illipe
(engkabang, kawang, tengkawang) nuts. As such they represent important genetic resources;
yet all appear to be rare, and under present wide forest conversion gravely threatened with
extinction. None have been sufficiently collected in flower or fruit for scientific description.
One other such species is ‘63. Shorea sp.’ (Ashton op. cit. (1968) 122), from Pelagus rapids
and Bt. Alet, Nanga Bah, Kapit district.
101. Shorea pubistyla P.S.Ashton

(Latin, pubi = downy, stylus = the floral style; the pubescent stylopodium)
Newman et al. op. cit. (1996) 198. Type: Anderson S 14930, Borneo, Sarawak, Kuching district,
Semengoh FR (holotype K; isotypes KEP, L).
Large emergent tree, to 50 m tall, to 1.6 m diameter; bole straight, cylindrical; crown
loosely hemispherical, with a few large branches, golden suffused from below; buttresses to
1.5 m tall, stout. Bark pale chocolatebrown mottledgrey, deeply irregularly fissured and
chunkily flaky; dammar as frequent pale yellow incrustations; inner bark dark coffee
coloured; heartwood rich reddish brown. Twig, leaf bud, inflorescence, parts of calyx
exposed in bud, petiole, and midrib below densely dark fulvous scabridtomentose; more
shortly sparsely so on venation below, midrib above, and stipule outside; stipule within,
bracteole and fruit calyx densely pale fulvous pubescent, caducous on fruit; outside of petal,
ovary, stylopodium, and nut evenly pale creambrown pubescent. Twigs prominently ribbed
at first, stout, c. 7 mm diameter apically. Leaf buds ovoid, compressed, to 1.3 × 1.3 mm.
Stipules lanceolate, to 25 × 7 mm, caducous. Leaves thickly coriaceous, drying warm
chocolatebrown below; blade broadly oblong to obovate, 11–19 × 7–12 cm, base obtuse to
cordate, margin shallowly subrevolute, apex retuse, obtuse or shortly acuminate; midrib
slender and more or less evident, furrowed above, prominent below; lateral veins 14–18
pairs, stoutly prominent below, obscurely sunken above; intercostal venation remotely
scalariform; petiole 2.5–4 cm long, stout, weakly geniculate. Inflorescences axillary, stout;
rachis ribbed, lax, to 20 cm long, singly branched, branchlets bearing to 12 flowers;
bracteoles oblongovate, subacute, to 8 × 5 mm, fugaceous. Flowers: petals pink outside,
crimson within; stamens 15, connectival appendage 2–3x as long as anther, glabrous; ovary
ovoid, crowned by a slightly longer cylindrical stylopodium and short glabrous style.
Fruits: pedicels to 3 mm long and diameter, stout; calyx lobes unequal, 3 longer lobes to 14
× 2.5 cm, tapering to c. 8 mm above the saccate base, 2 shorter ones loratelobed, to 12 ×
1.2 cm, otherwise similar. Nuts narrowly ovoid, to 3 × 1.8 cm, with to 4 mm long apical
tapering stylopodium remnant.
318
Vernacular name. Sarawak—meranti bulu merah (preferred name).
Distribution. Endemic in Borneo (W and C Sarawak and presumably also in the Kapuas
valley, W Kalimantan). In Sarawak known from Kuching, Miri and Tatau districts (e.g., S
16473, S 19575, S 32422, S 42349, and S 68407).
Ecology. Locally frequent, on deep leached yellow sandy clay soils in mixed dipterocarp
forest, on low hills, at altitudes to 400 m, generally not far from the coast. Occurring in
Lambir NP; elsewhere vulnerable owing to its accessible lowland habitat being liable to
land conversion.
102. Shorea quadrinervis Slooten

(Latin, quadri = four, nervis = nerved; the fourveined leaf)
Bull. Jard. Bot. Buitenz. 3, 17 (1942) 220; Browne op. cit. 142; Ashton op. cit. (1964) 213, op. cit.
(1968) 116, op. cit. (1982) 534; Meijer & Wood op. cit. 140; Burgess op. cit. 155, 183; Anderson op.
cit. (1980) 129; PROSEA op. cit. 401; Coode et al. (eds.) op. cit. 82; Newman et al. op. cit. (1996)
199. Lectotype (designated here): bb. 29450, Borneo, W Kalimantan, Sambas, Sg. Akar
(hololectotype BO).
Mediumsized emergent tree, to 45 m tall, to 1.4 m diameter; bole straight, cylindrical,

frequently quite short; crown hemispherical to oblong, the branch endings pendent;
buttresses to 2.5 m tall, stout. Bark pinkish grey to pale brown, shallowly vsection fissured,
eventually deeply so and powdery chunkily flaking; dammar as pale yellow incrustations;
inner bark bright crimson, paler at cambium; heartwood pale reddish brown. Young parts
shortly densely tawny pubescent, more or less caducous on stipule, bracteole, leaf bud, and
blade below, becoming sparse on fruit calyx; base of midrib above, inflorescence, parts of
perianth exposed in bud, ovary, and nut persistently so. Twigs c. 3 mm diameter apically,
terete, stout, strongly ridged below stipule scars, becoming minutely cracked; stipule scars
pale, prominent, falcate or descendingamplexicaul. Leaf buds compressed, ovate
subcordate, to 20 × 13 mm, loosely enveloped in stipules. Stipules ovate to subcordate, to
25 × 13 mm, prominently ridged along their 5 veins, pairs sometimes united at base,
subpersistent. Leaves strongly concave, thinly coriaceous, drying warm reddish brown
below; blade broadly ovate to elliptic, 10–18 × 5–8 cm, base obtuse, margin more or less
revolute, apex with tapering acumen to 1.3 cm long; midrib obscure, sunken above,
prominent below; lateral veins c. 4 pairs, arched, ascending, prominent below with at least
one prominent but shorter intermediate vein; intercostal venation slender, scalariform;
petiole 0.8–1 cm long. Inflorescences terminal or axillary; rachis lax, spreading, terete or
angular, to 28 cm long, to trebly branched, branchlets bearing to 6 flowers; bracts as the
stipules, caducous. Flowers: petals pink, paler at margins; stamens 15, connectival
appendage shorter than anther, reflexed; ovary and stylopodium conical, style short, stigma
relatively prominent. Fruits: calyx lobes unequal, 3 longer lobes to 8 × 1.3 cm, tapering to
c. 3 mm above the saccate base, 2 shorter ones to 5.5 × 0.7 cm, otherwise similar. Nuts
narrowly ovoid, to 1.5 × 0.7 cm, with c. 1.5 mm long slender style remnant.
319
Vernacular names. Sabah—seraya sudu (preferred name). Sarawak—meranti sudu

(preferred name).
Distribution. Endemic in Borneo. Known in Sabah from Beaufort and Sipitang districts
from Belaga, Bintulu, Kapit, Kuching, Limbang, Lubok Antu, Lundu, Marudi, Miri, and
Simunjan districts (e.g., S 10067, S 27135, S 32271, S 37822, and S 69660). Also occurring
in Brunei (e.g., BRUN 1800 and BRUN 3066) and Kalimantan (e.g., bb. 30222 and Yamada
K 9525).
Ecology. Common in mixed dipterocarp forest on leached yellow shallowly sandy humic
soils in the lowlands, and also on skeletal soils on ridges, at altitude to 700 m. Common in
Bako, Kubah, Lambir and Mulu NPs; not vulnerable.
103. Shorea resinosa Foxw.

(Latin, resinosus = resinous; the resinous nut)
(1982) 496; Anderson op. cit. (1980) 124; PROSEA op. cit. 412; Newman et al. op. cit. (1996) 201.
Type: Symington FMS 17512, Peninsular Malaysia, Ulu Selangor, Serendah FR (holotype KEP).
hemispherical; buttresses small or tall, rather sharp. Bark dark grey, coarsely fissured and
eventually chunkily flaky; inner bark pale and dark yellow laminated. Young twig,
inflorescence, leaf bud, parts of perianth exposed in bud, stipule outside, and petiole buff
fugaceous puberulent. Twigs terete, smooth, c. 2 mm diameter apically. Leaf buds ellipsoid,
to 3 × 2 mm. Stipules elliptic, to 12 × 8 mm. Leaves thinly coriaceous, undulate, drying
tawny brown below; blade lanceolate to elliptic, base cuneate, apex with slender acumen to
1.2 cm long; midrib obscurely sunken above, slender but prominent below; lateral veins 10–
13 pairs, slender but prominent below, not drying paler than blade; intercostal venation
slender, hardly elevated below, scalariform; petiole 0.9–1.7 cm long. Inflorescences: rachis
terete, singly branched, to 8 cm long. Flowers: buds to 12 × 4 mm; corolla pale yellow;
stamens 15, connectival appendage c. 2½x as long as anther, glabrous; ovary ovoid,
without stylopodium, glabrous, style c. 3x the length of ovary, trifid towards apex. Fruits:
pedicels to 4 mm long, to 3 mm diameter; calyx lobes unequal, 3 longer lobes to 9.5 × 2.5
cm, tapering to c. 7 mm above the saccate base, 2 shorter ones linearlobed, to 7 × 0.6 cm,
similar at base. Nuts ovoid, to 1.5 × 1.5 cm, with to 6 mm long filiform style remnant,
frequently resincoated.
Vernacular name. Sarawak—meranti belang (preferred name).
Distribution. Sumatra, Peninsular Malaysia and NW Borneo. In Sarawak recorded from

Bau and Lundu districts (e.g., S 397, S 9616 and S 27005).
Ecology. Rare, apparently confined to mixed dipterocarp forest, on yellow sandy soils over
sandstone. Probably endangered.
320
104. Shorea retusa Meijer

(Latin, retusus = notched; the leaf apex)
(1968) 117, op. cit. (1982) 537; Anderson op. cit. (1980) 129; Newman et al. op. cit. (1996) 201.
Lectotype (designated here): bb. 29715, Borneo, C Kalimantan, Muara Tewe, Barito Ulu
(hololectotype BO).
Canopy to low emergent tree, to 40 m tall, to 80 cm diameter; bole straight, cylindrical;

crown becoming hemispherical, diffuse; buttresses low, rounded. Bark greyish brown,
becoming shallowly but distinctly vsection fissured; inner bark yellowish brown;
heartwood pale brown; dammar as pale creambrown exudations. Young parts greyish
sericeous, glabrescent except on leaf bud, parts of calyx and petals exposed in bud, ovary,
stylopodium, and stipule outside. Twigs terete, muchbranched, c. 1 mm diameter apically.
Leaf buds ellipsoid, to 3 × 2 mm. Stipules elliptic, to 7 × 3 mm, caducous. Leaves
coriaceous, shiny, drying pale chocolatebrown, darker above; blade elliptic, 3–9 × 1.5–5
cm, base cuneate, apex shallowly retuse; midrib obscurely sunken above, slender and raised
but not prominent below; lateral veins 7–10 pairs, arched, ascending, very slender and
hardly raised below; intercostal venation slender, scalariform, obscure below; petiole short,
0.6–0.9 cm long. Inflorescences terminal or axillary; rachis terete, to 10 cm long, branchlets
bearing to 7 flowers; bracteoles elliptic, to 3 × 2 mm. Flowers: buds to 8 × 3 mm; corolla
pale yellow; stamens 15, connectival appendage slender, equal or 2–4x the length of anther,
becoming reflexed; ovary and stylopodium narrowly conical, crowned with a short glabrous
style. Fruit: pedicels to 1 mm long; calyx lobes unequal, 3 longer lobes to 10 × 1.8 cm,
tapering to c. 4 mm above the saccate base, 2 shorter ones loratelobed, to 5.5 × 0.4 cm,
similar at base. Nuts ovoid, to 1.3 × 1.2 cm with 3 mm long style remnant.
Vernacular names. Sabah—seraya daun tumpul (preferred name). Sarawak—meranti telur

(preferred name).
Distribution. Endemic in Borneo. Known in Sabah from Tawau district (e.g., SAN 19569,
SAN 21504 and Villamal 87) and in Sarawak from Bintulu, Kuching, Lundu, and Serian
districts (e.g., S 952, S 7579, S 10268, S 13373, and S 47099). Also occurring in Kalimantan
(e.g., Ambriansyah & Arifin Berau 1067 and bb. 29716).
Ecology. Locally frequent in kerangas forest, mostly on shallow podsols over sandstone.
Occurring in Bako NP; overall conservation status uncertain.
105. Shorea revoluta P.S.Ashton

(Latin revolutus = turned in; the leaf margin)
Newman et al. op. cit. (1996) 202. Type: Smythies S 5914, Borneo, Sarawak, Limbang district, Bt.
Sagan (holotype K).
321
Fig. 28. Shorea richetia. A, flowering leafy twig; B, flower bud; C, opening flower; D,
adaxial view of outer sepal; E, adaxial view of inner sepal; F, adaxial view of petal with
stamens; G, abaxial view of petal; H, adaxial view of stamens; I, abaxial view of stamens; J,
longitudinal section of open flower; K, fruit; L, longitudinal section of fruit. (A from S 109,
B–J from S 563, K–L from S 15254.)
322
Low emergent tree, to 45 m tall, to 1 m diameter; bole cylindrical; crown hemispherical,

diffuse; buttresses to 2 m tall, stout. Bark pale to dark chocolatebrown, frequently mottled
with grey, prominently vsection fissured, the intervening ridges eventually crumbling in
chunky flakes; dammar smears pale creamyellow, common; inner bark and heartwood rust
red. Twig, inflorescence, leaf bud, calyx and stipule outside, and petiole densely pale brown
to fulvous scabridpuberulent; veins below and midrib above sparsely so; parts of petal
exposed in bud, nut and inside of stipules evenly pale buff puberulent. Twigs at first slightly
compressed and ribbed, c. 2–3 mm diameter apically. Leaf buds ovoid, compressed,
subacute, 4–7 × 3–5 mm. Stipules elliptic to oblong, obtuse, to 15 × 7 mm, caducous.
Leaves coriaceous, drying warm tanbrown; blade ovate, 10–15 × 5.5–10 cm, base obtuse
to subcordate, margin prominently revolute, apex with slender acumen to 1.5 cm long;
midrib more or less obscure and sunken above, prominent below; lateral veins 9–12 pairs,
prominent below, arched, wellspaced, with glabrous axillary domatia; intercostal venation
slender, scalariform, hardly elevated below; petiole 1.3–1.5 cm long. Inflorescences
terminal or axillary; rachis rigid, to 22 cm long. Flowers: corolla dark red; stamens 15,
connectival appendage slender, as long as anther, becoming reflexed; ovary and
stylopodium narrowly ovoid, style half their length, slender, glabrous. Fruits: pedicels to
1.5 mm long; calyx lobes unequal, 3 longer lobes to 7.5 × 1.3 cm, tapering to 3 mm above
the saccate base, 2 shorter ones linearlobed, to 3.5 × 0.2 cm, similar at base. Nuts ovoid,
to 1.5 × 0.8 cm, with c. 1.5 mm style remnant.
Vernacular names. Sabah—seraya daun tajam (preferred name). Sarawak—meranti

kerangas (Malay).
Distribution. Endemic in Borneo. In Sabah known from Beaufort and Sipitang districts
(e.g., SAN 15135, SAN 55656, SAN 110013, and SAN 131905) and in Sarawak from Lawas,
Limbang and Marudi districts (e.g., S 5618 and S 11248). Also occurring in Brunei (e.g.,
BRUN 5646, S 5782 and S 5814).
Ecology. Locally frequent in kerangas forest, both on terraces and sandstone plateaux, at
altitudes to 1200 m. Endangered by logging.
106. Shorea richetia Symington Fig. 28.

(after the former genus Richetia, now Shorea sect. Richetioides)
Gard. Bull. S. S. 9 (1938) 330; Slooten op. cit. (1956) 335; Ashton op. cit. (1968) 89, op. cit. (1982)
474; Anderson op. cit. (1980) 123; Newman et al. op. cit. (1996) 203. Type: Beccari PB 2888,
Borneo, Sarawak, Matang (holotype P; isotype L).
Main canopy tree, to 40 m tall, to 80 cm diameter; bole straight, cylindrical; crown dense,
rather small, hemispherical; buttresses to 1 m tall, small, stout. Bark dull greyish brown,
tardily cracking, then thinly oblongflaking eventually leaving a scrollmarked new surface.
Twig apices, inflorescence, leaf buds, and stipules puberulent; ovary and nut buff pubescent;
other parts glabrous. Twigs terete, smooth, 1–2 mm diameter apically. Leaf buds ovoid,
subacute, to 2 × 2 mm. Stipules lanceolate, to 5 × 2 mm. Leaves thickly coriaceous, drying
dark greyish green; blade broadly elliptic, 5–11 × 3–6.5 cm, base broadly cuneate to
occasionally subcordate, margin narrowly subrevolute, apex with broad, tapering acumen
to 1 cm long; midrib evident, flat to shallowly furrowed above, prominent below; lateral
323
veins 5–7 pairs, ascending, arched, prominent below; intercostal venation subscalariform;
petiole stout, 0.6–1 cm long. Inflorescences terminal or axillary; rachis terete or
compressed, glabrescent, lax, to 8 cm long, singly branched, branchlets bearing to 5 flowers;
bracteoles deltoid, to 2 × 1 mm, fugaceous. Flowers: buds to 7 × 2 mm; petals limeyellow;
stamens 16–17, connectival appendage at least 1½x the length of anther; ovary ovoid,
without stylopodium, style 2–3x the length of ovary, columnar, tapering, pubescent in basal
half. Fruits: pedicels to 2 mm long, stout; calyx lobes subequal, narrowly ovate, to 1.7 × 1
cm, acute, saccate, tuberculate. Nuts obovoid, acute, to 2.5 × 1.5 cm.
Vernacular name. Sarawak—lun melapi (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from Tenom district (e.g., SAN 64311)
and in Sarawak from Kuching and Lundu districts (e.g., S 6404, S 15525, S 25451, S 32409,
and S 32520).
Ecology. Very local, in mixed dipterocarp forest on leached sandy clay soils, on low hills, at
altitudes to 200 m. Recorded from Kubah NP; elsewhere endangered by land conversion.
107. Shorea rotundifolia P.S.Ashton Plate 4E.

(Latin, rotundus = round, folius = leaf; the distinctive leaf shape)
Newman et al. op. cit. (1996) 204. Type: Smythies S 9470, Borneo, Sarawak, Kapit district, Pelagus,
Sg. Iran (holotype K; isotype KEP).
Low emergent tree, to 40 m tall, to 70 cm diameter; bole straight, cylindrical; buttresses to

80 cm tall and out, stout. Bark greyish brown and chocolatemottled with a crimson
‘scrape’, smooth, hoopmarked, inner bark pinkish brown with pale patches; sapwood pink,
relatively soft. Young twig, petiole and stipule pruinose; petals shortly pubescent on parts
exposed in bud; parts otherwise glabrous. Twigs terete or slightly compressed at first, c. 3 ×
2 mm apically; stipule scars amplexicaul, prominent. Leaf buds lanceolate, compressed,
large, to 20 × 8 mm. Stipules lanceolatefalcate, obtuse, to 60 × 20 mm, subpersistent.
Leaves remaining distichous, coriaceous, shiny, drying dark purplish brown below; blade
broadly ovate to orbicular, 9–21 × 8–14 cm, base obtuse to cordate, apex cuspidate,
acumen to 1 cm long; midrib evident above, flat, prominent below; lateral veins 9–11 pairs,
arched, slender but prominent below; intercostal venation remotely scalariform, elevated
below; petiole 3–4 cm long, somewhat geniculate. Inflorescences terminal or axillary;
rachis compressed, lax, to 15 cm long, regularly singly branched, branchlets bearing remote
flowers; bracteoles ellipticlanceolate, acute, to 10 × 6 mm. Flowers: buds to 10 × 4 mm;
stamens 15, connectival appendage c. 3x the length of anther; ovary ovoid, glabrous, style
and stylopodium spindleshaped. Fruits: pedicels c. 6 mm long; calyx lobes unequal, 3
longer lobes spatulate, to 13 × 2.5 cm, tapering to c. 6 mm above the saccate base, 2
shorter ones narrowly lanceolate, to 8 × 0.7 cm, similar at base. Nuts ovoid, to 2.5 × 1.1
cm, with to 7 mm long style remnant.
324
Vernacular name. Sarawak—langgai (Iban).
Distribution. Endemic in Borneo. Confined to Bintulu and Kapit districts in C Sarawak

(e.g., S 22051 and S 22053).
Ecology. Local and rare, in scattered clumps on leached sandy clay soils along ridges, at
altitudes to 400 m. Vulnerable.
Notes. It is possible that this entity may merely represent one or more varieties of S.
amplexicaulis, with which it ofen grows. The small groups in which the species occur differ
somewhat from one another, but their characters are internally constant. This extreme
variation, unusual among dipterocarps, is nevertheless common among species in sect.
Pachycarpae (see, for instance, under S. macrophylla, S. pinanga and S. praestans).
108. Shorea rubella P.S.Ashton

(Latin, rubellus = reddish; the drying leaf blade)
Newman et al. op. cit. (1996) 205. Type: Ashton BRUN 3078, Borneo, Brunei, Andulau FR (holotype
Mediumsized emergent tree, to 45 m tall, to 1.4 m diameter; bole tall, cylindrical; crown
hemispherical, spreading, diffuse, becoming pale from below; buttresses to 2 m tall, stout,
prominent. Bark pinkish brown, becoming coarsely deeply vsection fissured, eventually
chunkily oblongflaked; dammar smears rare, cream; inner bark rustbrown; heartwood pale
pink. Young parts evenly pale creambuff puberulent, persistent only on inflorescence; leaf
bud, parts of perianth exposed in bud, ovary, nut, and fruit calyx glabrescent. Twigs 2–3
mm diameter apically, ridged and compressed on drying, becoming terete, stout; stipule
scars c. 1.5 mm long, narrow, horizontal. Leaf buds narrowly ovoid to subfalcate, acute, 6–9
× 2–3 mm. Stipules hastate to falcate, acute, to 18 × 6 mm, caducous. Leaves thickly
coriaceous, sparsely cream lepidote below (mature trees), drying rich pinkish brown; blade
broadly ovate but often somewhat irregularly shaped, 9.5–14 × 6–8 cm, base obtuse or
broadly cuneate, margin sometimes subrevolute, apex with tapering acumen to 1 cm long;
midrib evident, flat or shallowly furrowed above, prominent below; lateral veins 5–7 pairs,
arched, wellspaced, prominent below, sometimes with minute porelike axillary domatia;
intercostal venation densely scalariform, slender, more or less obscure; petiole 2.3–3.5 cm
long. Inflorescences terminal or axillary; rachis ribbed on drying, to 15 cm long, branchlets
lax, bearing to 5 flowers. Flowers: buds to 7 × 3 mm; petals pale pink, darker towards base;
stamens 15, subequal, filaments lorate, abruptly tapering below the large narrowly oblong
equal anthers, connectival appendage short, tapering, becoming somewhat recurved; ovary
small, ovoid, stylopodium indistinct, style filiform, c. 3x the length of ovary. Fruits: calyx
shorter ones linearlobed, to 5 × 0.6 cm, similar at base. Nuts ovoid, to 1.8 × 1.2 cm, acute.
Vernacular name. Sarawak—meranti laut putih (preferred name).
325
Distribution. Endemic in Borneo. In Sabah known from Beaufort and Papar districts (e.g.,
SAN 24812, SAN 24813 and SAN A 1750) and in Sarawak from Belaga, Bintulu, Kapit,
Miri, and Tatau districts (e.g., S 9474, S 15128 and S 19577). Also occurring in Brunei (e.g.,
FMS 30574, FMS 35667, S 1925, and SAN 17564).
Ecology. Locally frequent, on the deep yellow sandy soils of the subcoastal neogene
sedimentary hills, the northernmost Rajang series, and the Arip rhyolite. Well represented in
Lambir NP; elsewhere endangered.
109. Shorea rubra P.S.Ashton

(Latin, ruber = red; the leaf undersurface)
Newman et al. op. cit. (1996) 206. Type: Smythies S 9475, Borneo, Sarawak, Pelagus, Ulu Iran
(holotype K; isotypes KEP, L, SAR).
Large emergent tree, to 50 m tall, to 1.3 m diameter; bole, tall, cylindrical; crown dense,
cauliflowershaped, brownish from below; buttresses to 2.5 m tall, stout. Bark dark
chocolate to blackish, becoming deeply vsection fissured, eventually the ridges becoming
chunkily flaky; dammar smears bright pale yellow; inner bark yellowish brown; heartwood
deep purplish red. Twig, inflorescence, bud, calyx outside, stipule, bracteole, petiole, midrib
above, and leaf below densely persistently deep rufous brown scabrid puberulent. Twigs
terete, muchbranched, to 1.5 mm diameter apically. Leaf buds compressed, ovoid, acute, 4–
7 × 3–4.5 mm. Stipules hastate, to 16 × 5 mm. Leaves thinly coriaceous, drying chocolate
brown overlain by deep rufousbrown tomentum; blade broadly ovate, 8–13 × 4–7.5 cm,
base obtuse, margin narrowly revolute, apex with acumen to 1.3 cm long; midrib obscurely
sunken above, slender but prominent below; lateral veins 11–13 pairs, prominent below,
arched; intercostal venation densely scalariform, hardly elevated below; petiole 1–1.4 cm
long. Inflorescences terminal or axillary; rachis terete, rather rigid, to 14 cm long,
branchlets compact, bearing to 5 flowers; bracteoles broadly ovate, subacute, to 4 × 3.5 mm.
Flowers: buds to 7 × 4 mm; petals yellow; stamens 15, connectival appendage hardly
reflexed; ovary and stylopodium ovoidconical, with shorter glabrous style. Fruits: calyx
shorter ones linearlobed, to 4.5 × 0.4 cm, otherwise similar. Nuts broadly ovoid, to 1 × 0.9
cm, with c. 2 mm style remnant.
Vernacular names. Sabah—seraya bingkai (preferred name). Sarawak—meranti merah

kesumba (preferred name).
Distribution. Endemic in Borneo. In Sabah known from Beaufort, Keningau,

Kinabatangan, Ranau, Sandakan, Sipitang, and Tawau districts (e.g., SAN 15193, SAN
22354, SAN 50702, and SAN 76795) and in Sarawak from Belaga, Bintulu, Kapit, Kuching,
Lawas, Limbang, Marudi, Miri, Sibu and Tatau districts (e.g., S 3551, S 15875, S 22241, S
29499, and S 31723). Also occurring in Brunei (e.g., BRUN 784 and FMS 48492) and
Kalimantan (e.g., Jarvie et al. 5066).
326
Ecology. Scattered in mixed dipterocarp forest, usually on welldrained leached sandy and
clay soils, both on coastal hills, and on inland ridges, at altitudes to 800 m. Occurring in
Lambir and Mulu NPs; elsewhere vulnerable.
110. Shorea rugosa F.Heim

(Latin, rugosus = wrinkled; the appearance imparted by the rough tomentum)
Bull. Mens. Soc. Linn. Paris 2 (1891) 973; Merrill op. cit. (1921) 406; Symington op. cit. (1933) 132;
Masamune op. cit. 496; Ashton op. cit. (1964) 218, op. cit. (1968) 118, op. cit. (1982) 540; Meijer &
al. (eds.) op. cit. 82; Newman et al. op. cit. (1996) 207. Type: Beccari PB 2638, Borneo, Sarawak,
Matang (holotype P).
Emergent tree, to 50 m tall, to 1.3 m diameter; bole cylindrical, straight; crown dense,
cauliflowershaped; buttresses to 2.5 cm tall, prominent, rather stout. Bark dark chocolate
to blackish, in young trees mottled with fawn and ochre, becoming densely deeply vsection
fissured, eventually powdery chunkily flaky; dammar bright yellow, in bole incrustations;
inner bark yellowish brown; heartwood deep purplish red. Young twig, inflorescence, leaf
bud, flower calyx and stipule outside, petiole, and venation below densely purplish brown
coarsely scabridtomentose; stipule inside, midrib above and blade below shortly evenly so;
blade above and fruit calyx sparsely caducously so; parts of corolla exposed in bud, ovary,
stylopodium, and nut evenly fulvous pubescent. Twigs ribbed at first, becoming terete, to 2.5
mm diameter apically. Leaf buds ovoid, subacute, 4–6 × 3–5 mm. Stipules oblongelliptic,
subacute, to 14 × 5 mm. Leaves thickly coriaceous, drying chocolatebrown scabrous
below; blade oblongovate to oblongobovate, 9–17 × 4–9 cm, base obtuse, margin more or
less narrowly revolute, apex with broad acumen to 0.9 cm long; midrib obscurely sunken
above, prominent below; lateral veins 14–19 pairs, prominent below; intercostal venation
distantly scalariform, elevated below; petiole 1.3–2.3 cm long, stout. Inflorescences
terminal or axillary; rachis terete or slightly compressed, lax, to 12 cm long, branchlets
bearing to 6 flowers; bracteoles elliptic, subacute, to 4 × 3 mm. Flowers: buds to 7 × 4 mm;
petals yellow; stamens 15, connectival appendage short, reflexed; ovary and stylopodium
conical, style short glabrous. Fruits: calyx lobes unequal, 3 longer lobes to 10.5 × 2.5 cm,
tapering to 4–7 mm above the saccate base, 2 shorter ones linearlobed, to 5 × 0.4 cm,
similar at base. Nuts ovoid, to 2.3 × 1.5 cm, acute.
Vernacular names. Sabah—seraya buaya hantu (preferred name). Sarawak—meranti

buaya hantu (preferred name).
Distribution. Endemic in Borneo. Widespread but local in Sarawak and recorded from
Belaga, Bintulu, Kapit, Kuching, Lundu, Miri, and Sibu districts (e.g., S 7568, S 9466, S
15522, S 37854, and S 43398). Reported from Sabah in Meijer and Wood (op. cit.) but
collection now destroyed. Also occurring in Brunei (e.g., BRUN 3036 and S 2129) and
Kalimantan (e.g., bb. 19974 and bb. 29683).
Ecology. Scattered, rarely common, in mixed dipterocarp forest on deep yellow sandy
humic soils and grey leached sand in the ecotone to kerangas. Occurring in Bako, Lambir
and Mulu NPs; endangered outside parks system.
327
111. Shorea sagittata P.S.Ashton Fig. 29.

(Latin, sagittatus = with two basal lobes pointing downwards; the fruit calyx lobes)
Newman et al. op. cit. (1996) 209. Type: Sibat S 23616, Borneo, Sarawak, Simunjan district,
Balingian, Arip, Bt. Ijok (holotype K; isotype KEP).
Emergent tree, to 45 m tall, to 1.1 m diameter; bole straight, tall; crown evenly
hemispherical, diffuse; buttresses to 3 m tall, stout, prominent. Bark pinkish brown and
greymottled, becoming shallowly fissured, eventually chunkily oblongflaky; inner bark
pale yellow; heartwood reddish brown. Leaf blade below creambuff sericeous; corolla,
ovary and nut puberulent; other exposed fleshy parts densely pinkish brown scabrid
puberulent, persistent except on fruit calyx. Twigs ribbed, becoming terete, c. 2 mm
diameter apically; stipule scars short, ascending. Leaf buds broadly ovoid, to 7 × 7 mm.
Stipules ovate, subacute, to 10 × 8 mm, caducous. Leaves thinly coriaceous, drying pale
greyish brown; blade narrowly oblong to lanceolate, 7–15 × 3–6 cm; midrib obscurely
sunken above, prominent below; lateral veins 19–25 pairs, slender, dense, prominent below;
intercostal venation densely scalariform, obscure; petiole 0.9–1.7 cm long. Inflorescences
terminal or axillary; rachis terete, to 7 cm long, branchlets short bearing to 3 flowers;
bracteoles to 4 × 2 mm, narrow, acute. Flowers: buds to 6 × 3 mm; petals yellowish pink;
stamens 15; ovary and stylopodium conical, style short, glabrous. Fruits: calyx lobes
unequal, 3 longer lobes loratespatulate, to 12 × 2.5 cm, hardly tapering to the prominently
sagittateauriculate base with incrassate centre, 2 shorter ones linearlobed, to 2 × 0.3 cm,
not auriculate. Nuts ellipsoid, apiculate, to 1.5 × 0.8 cm, glabrescent.
Vernacular name. Sarawak—meranti daun mata lembing (preferred name).
Distribution. Endemic in Sarawak and known from Belaga, Bintulu, Kapit, Kuching,
Mukah, Simunjan, and Tatau districts (e.g., S 18316, S 23720, S 29166, S 53447, and S
60124); not yet recorded from Sabah.
Ecology. Locally frequent, in mixed dipterocarp forest on moist leached clay hillsides, also
floodplains, on sedimentaries and the Arip rhyolite. Endangered by land conversion.
112. Shorea scaberrima Burck Fig. 30.

(Latin, scaberrimus = very rough; the indumentum)
Med. Lands Pl. Tuin 3 (1886) 22; Merrill op. cit. (1921) 406; Masamune op. cit. 497; Browne op. cit.
82; Newman et al. op. cit. (1996) 210. Lectotype (designated here): s.c. s.n., Cult. Hort. Bog. VIII D
54 (hololectotype BO).
Main canopy tree, to 40 m tall, to 1.1 m diameter; bole often misshapen; crown dense, rather
irregular, more or less hemispherical; buttresses to 1.5 m tall, fairly stout. Bark at first
remaining smooth, hoopmarked, dappled grey, ochre and fawn brown, becoming shallowly
cracked and thinly patchily flaky; inner bark and heartwood coffeecoloured. Young twigs,
328
Fig. 29. Shorea sagittata. A, fruiting (young) leafy twig; B, detail of indumentum on lower
leaf surface; C, flower bud; D, longitudinal section of flower bud; E, gynoecium and
staments; F, adaxial view of outer sepal; G, adaxial view of inner sepal; H, abaxial view of
petal; I, adaxial view of petal; J, adaxial view of stamens; K, abaxial view of stamens; L,
fruit; M, fruit with the nut partially exposed. (A–B from S 7609, C–K from S 29166, L–M
from S 29652.)
329
Fig. 30. Shorea scaberrima. A, fruiting (very young) leafy twig; B, detail of indumentum on
lower leaf surface; C, side view of tufted hairs; D, flower bud; E, longitudinal section of
flower bud; F, gynoecium; G, adaxial view of outer sepal; H, adaxial view of inner sepal; I,
abaxial view of petal; J, adaxial view of petal; K, adaxial view of stamens; L, abaxial view
of stamens; M, mature fruit with the nut partly exposed; N, longitudinal section of mature
fruit. (A–B from S 3285, D–E from S 2132, F–N from S 36915.)
330
inflorescence, leaf bud, parts of calyx exposed in bud, stipule outside, petiole, leaf blade
below, and midrib above more or less sparsely persistently golden brown scabrid
pubescent; parts of corolla exposed in bud, ovary, stylopodium, and nut evenly densely so;
inside of stipule, bracteole and fruit calyx sparsely evenly so. Twigs ribbed at first,
becoming terete, 2–3 mm diameter apically. Leaf buds broadly ovoid, subacute, slightly
compressed, 4–6 × 4–5 mm. Stipules broadly hastate, to 18 × 8 mm, caducous. Leaves
thinly coriaceous to chartaceous, drying chocolatebrown below, purplish brown above;
blade oblongovate to oblongobovate, base obtuse, rarely subcordate, apex with tapering
acumen to 0.8 cm long; midrib evident, flat to shallowly furrowed above, prominent below;
lateral veins 14–17 pairs, slender but prominent below; intercostal venation scalariform,
distinctly elevated below; petiole 1.8–2.5 cm long. Inflorescences terminal or axillary;
rachis ribbed to terete, to 8 cm long, singly branched, branchlets bearing to 3 secund
flowers; bracteoles narrowly deltoid, to 4.5 × 3 mm. Flowers: buds to 8 × 5 mm; corolla
pink; stamens 15, connectival appendage at least 2x the length of anther; ovary ovoid,
small, stylopodium and style at least 3x the length of ovary, stoutly filiform with glabrous
apex. Fruits: calyx lobes unequal, 3 longer lobes nevertheless shorter than nut, to 4.5 × 1
cm, variable, tapering to c. 7 mm above the saccate base, 2 shorter ones linearlobed, to 3
× 0.3 cm, similar at base. Nuts obovoid, to 5 × 2.5 cm, acute.
Vernacular names. Sabah—seraya mempelas (preferred name). Sarawak—meranti paya

bersisik (preferred name).
Distribution. Endemic in Borneo. Recorded in Sabah from Beaufort, Labuk Sugut, Papar,
Penampang, Sipitang, and Tenom districts (e.g., SAN 15062, SAN 16339, SAN 24276, SAN
A 3016, and SAN A 3285) and in Sarawak from Bau, Belaga, Bintulu, Kapit, Kuching,
Lawas, Limbang, Lubok Antu, Lundu, Marudi, Miri, Mukah, Samarahan, and Tatau districts
BRUN 2001 and KEP 80085) and Kalimantan (e.g., Suzuki K 9757 and the type).
Ecology. Locally common in mixed dipterocarp forest, on shallowly humic leached sandy
clay, and welldrained skeletal clay soils; throughout the lowlands and along inland ridges,
at altitude to 850 m. Common in Bako, Lambir and Mulu NPs; not vulnerable.
113. Shorea scabrida Symington

(Latin, scabridus = rough; the indumentum)
Gard. Bull. S. S. 8 (1935) 287; Masamune op. cit. 497; Browne op. cit. 143; Anderson op. cit. (1963)
159, op. cit. (1980) 130; Ashton op. cit. (1964) 221, op. cit. (1968) 119, op. cit. (1982) 543; Meijer &
Wood op. cit. 146; Burgess op. cit. 155, 183; PROSEA op. cit. 402; Coode et al. (eds.) op. cit. 82;
Newman et al. op. cit. (1996) 211. Type: Yusope S 171, Borneo, Sarawak, Marudi district, Bt. Liman
FR (holotype KEP).
Low emergent tree, to 45 m tall, to 1 m diameter; bole straight, cylindrical; crown becoming
shallowly cauliflowershaped, diffuse; buttresses to 1.5 m tall, stout. Bark at first mottled
purplish brown and grey, becoming pink to mauvebrown, widely vsection fissured; inner
bark rich reddish brown with paler wedges; heartwood pinkish brown. Young twig,
inflorescence, leaf bud, midrib above, and blade and venation below persistently more or
less sparsely tawnybrown scabridpuberulent. Twigs terete, much branched, c. 2 mm
331
diameter apically; stipule scars short, horizontal. Leaf buds ovoid, obtuse, compressed, 2.5–
5 × 1.5–3 mm. Stipules oblong, obtuse, to 6 × 3 mm, fugaceous. Leaves coriaceous, drying
orangebrown; blade obovate to elliptic, 5–9 × 3–5 cm, base broadly cuneate, occasionally
obtuse, apex retuse to shortly acuminate; midrib obscure and sunken above, prominent
below; lateral veins 8–11 pairs, slender, raised but hardly prominent below, arched,
occasionally with small porelike domatia; intercostal venation slender, scalariform, barely
elevated; petiole 0.9–1.1 cm long. Inflorescences terminal or axillary; rachis terete or
slightly compressed, to 8 cm long, branchlets short bearing to 7 flowers; bracteoles ovate,
subacute, to 3.5 × 2.5 mm. Flowers: buds to 8 × 3.5 mm; petals cream, pink at base;
stamens 15, connectival appendage slightly longer than anther, becoming reflexed; ovary
and stylopodium narrowly conical, style almost as long as both, glabrous. Fruits: calyx
lobes unequal, 3 longer lobes to 7 × 1.5 cm, tapering to c. 3 mm above the saccate base, 2
shorter ones linearlobed, to 3 × 0.3 cm, similar at base. Nuts ovoid, to 1 × 0.8 cm, shortly
apiculate.
Vernacular names. Sabah—seraya lop (preferred name). Sarawak—meranti lop (preferred

name).
Distribution. Sumatra and Borneo. In Sabah widespread, known from Beaufort, Labuk
Sugut, Papar, and Sipitang districts (e.g., SAN 15854, SAN 16681, SAN 24343, SAN 25431,
and SAN 27969) and in Sarawak from Belaga, Betong, Bintulu, Kapit, Kuching, Lundu,
Marudi, Miri, Samarahan, Sibu, Simunjan, Sri Aman, and Tatau districts (e.g., S 14997, S
BRUN 5542) and Kalimantan (e.g., bb. 6399 and Jarvie & Ruskandi 5091).
Ecology. Locally frequent, in the edge of mixed peat swamp forest, in kerangas forest on
podsols, often in poor drainage, but also in mixed diptercarp forest on leached sandy and
sandy clay soils, at altitudes to 1200 m. Occurring in Bako, Lambir, Mulu NPs; elsewhere
Notes. A variable species. In dipterocarp forest the leaf is acuminate, the tomentum sparse,
in kerangas and peat swamp forest the leaf is smaller, retuse, and the tomentum more dense;
but there is much variation between the two.
114. Shorea scrobiculata Burck

(Latin, scrobiculatus = having the appearance of sawdust; the inflorescence indumentum)
Med. Lands. Pl. Tuin 3 (1886) 223; Merrill op. cit. (1921) 406; Masamune op. cit. 497; Browne op.
cit. 169; Ashton op. cit. (1963) 272, op. cit. (1964) 144, op. cit. (1968) 74, op. cit. (1982) 449;
cit. (1998) 195. Lectotype (designated here): Beccari PB 2538, Borneo, Sarawak, Matang
(hololectotype K). Synonyms: Shorea pierreana F.Heim op. cit. (1892) 48; S. meadiana Symington
op. cit. (1939) 366; S. leptoderma Meijer op. cit. 331 (Type: Meijer SAN 21711, Borneo, Sabah,
Sepilok FR; holotype SAN; isotypes K, KEP, L, SAR), syn. nov., Meijer & Wood op. cit. 180, Ashton
op. cit. (1982) 450.
332
Main canopy or low emergent tree, to 40 m tall, tall 1.2 m diameter; bole straight,
sometimes fluted; crown hemispherical, dense; buttresses to 0.7 m tall, thin, sometimes with
flying buttresses. Bark dark yellowish to reddish brown, becoming cracked, oblong flaky,
the flakes curling up from below, sometimes becoming shaggy. Twig, inflorescence, leaf
bud, parts of perianth exposed in bud, ovary, stylopodium, nut, bracteole, stipule, and
petiole densely buffpuberulent; fruit calyx, midrib on both surfaces and veins below
sparsely so. Twigs terete, muchbranched, slender, c. 1.5 mm diameter apically. Leaf buds
ovoid, more or less compressed, 2–3 × 1–2 mm apically. Stipules hastate, c. 7 × 3 mm.
Leaves thinly coriaceous, drying yellowish brown below, mauvegrey above; blade
narrowly ovate to oblonglanceolate, 5.5–11 × 2.5–4 cm, base broadly cuneate, apex with
narrow acumen to 1 cm long; midrib evident, elevated but somewhat furrowed above,
slender but prominent with the veins below; lateral veins 10–12 pairs, ascending, dense;
intercostal venation densely scalariform, hardly elevated; petiole c. 0.8 cm long.
Inflorescences terminal or axillary; rachis terete, slender, to 7 cm long, singly branched,
branchlets bearing to 9 flowers; bracteoles elliptic, subacute, to 3 mm long, fugaceous.
Flowers: buds to 6 × 2 mm; petals pink, cream at margin; stamens 20–30, filament and
anther glabrous, connectival appendage short, with 1(–2) apical bristle(s); ovary broadly
ovoid, stylopodium narrowly cylindrical, shorter than ovary, style as long as stylopodium,
tapering, columnar, glabrous. Fruits: calyx lobes unequal, 3 longer lobes to 5 × 1.2 cm,
tapering to 3 mm above the saccate base, 2 shorter ones to 3.5 × 0.4 cm, otherwise similar.
Nuts ovoid, to 1 × 0.7 cm, with c. 1.7 mm style remnant.
Vernacular names. Sabah—selangan batu kurap (preferred name). Sarawak—selangan

batu zang (preferred name).
Distribution. Peninsular Malaysia and Borneo. In Sabah recorded from Beaufort,

Keningau, Kinabatangan, Lahad Datu, Ranau, Sandakan, and Tawau districts (e.g., SAN
15012, SAN 21237, SAN 22848, SAN 76949, and SAN 126639) and in Sarawak from Belaga,
Bintulu, Kuching, Limbang, Lundu, and Miri districts (e.g., S 9641, S 24694, S 32212, S
43462, and S 52753). Also occurring in Brunei (e.g., BRUN 740 and SAN 17530) and
Kalimantan (e.g., bb. 29700 and Endert 2540).
Ecology. In mixed dipterocarp forest on leached welldrained sandy and clay soils, on low
hills and along shale ridges, at altitude to 700 m; locally frequent. Occurring in Bako,
Kubah, Lambir and Mulu NPs; though its habitat is gradually reduced by land conversion,
the species is not yet vulnerable.
115. Shorea seminis (de Vriese) Slooten Plate 4F.

(of Seminis, W Kalimantan)
In Merrill, PEB (1929) 204, op. cit. (1941) 117; Keith op. cit. 44; Masamune op. cit. 497; Browne op.
cit. 170; Ashton op. cit. (1963) 272, op. cit. (1964) 145, op. cit. (1968) 75, op. cit. (1982) 451; Meijer
& Wood op. cit. 185; Burgess op. cit. 203, 211; Anderson op. cit. (1980) 120; PROSEA op. cit. 433;
Kessler & Sidiyasa op. cit. 110; Coode et al. (eds.) op. cit. 83; Newman et al. op. cit. (1998) 196.
Basionym: Hopea seminis de Vriese op. cit. 32. Type: de Vriese s.n., Borneo, W Kalimantan,
Sambas, Seminis (holotype L). Synonyms: Hopea lanceolata de Vriese op. cit. 32; Shorea
schefferiana Hance op. cit. 303; Isoptera borneensis Scheff. ex Burck op. cit. (1886) 27; S. borneensis
(Scheff. ex Burck) Pierre, For. Fl. Coch. 3 (1889) t. 234; Ridleyinda borneensis (Scheff. ex Burck)
Kuntze op. cit. 65; I. seminis (Scheff. ex Burck) Burkill, Bull. Misc. Inform. Kew (1935) 317.
333
Large emergent tree, to 45 m tall, to 1.3 mm diameter, or more or less small leaning river
bank tree; bole straight, cylindrical, or bowed; crown dense, oblong to hemispherical;
buttresses to 2 m tall, thin. Twig, inflorescence, leaf bud, parts of perianth exposed in bud,
ovary, stylopodium, nut, stipule, bracteole, petiole, midrib on both surfaces, and veins below
evenly more or less densely greyish brown puberulent; fruit calyx sparsely so. Twigs at first
ribbed, becoming terete, sparingly branched, slender, c. 1.5 mm diameter apically; stipule
scars short, descending. Leaf buds ovoid to falcate, slightly compressed, 1.5–2.5 × 1–2 mm.
Stipules oblong, to 7 × 3.5 mm. Leaves thinly coriaceous, drying dull yellowish brown
below with paler veins, mauvegrey above; blade oblongovate to lanceolate, 9–18 × 2.5–8
cm, base obtuse or cuneate, apex with slender acumen 0.8–2 cm long; midrib evident, flat to
shallowly furrowed above, slender but prominent below; lateral veins 9–15 pairs,
ascending, slender but prominent below; intercostal venation densely scalariform, sinuate,
more or less obscure; petiole 1–1.5 cm long. Inflorescences terminal, axillary, or
ramiflorous; rachis terete, to 10 cm long, singly branched, branchlets bearing to 5 flowers.
Flowers: buds to 8 × 2 mm; petals cream, pink towards base; stamens 30–40, filaments with
a few long bristles, anthers glabrous, connectival appendage short, setose; ovary and
stylopodium conical to hourglassshaped, style short, glabrous. Fruits: calyx lobes
subequal, incrassate, orbicular, rotate, to 2 × 1.8 cm (usually smaller). Nuts ovoid to
globose, to 1 cm diameter, with to 2 mm long stout tapering style remnant.
Vernacular names. Sabah—selangan batu terendak (preferred name). Sarawak—

engkabang terendak (preferred name).
Distribution. Borneo and the Philippines. Known in Sabah from Kinabatangan, Kota Belud,
Pensiangan, Sandakan, and Tawau districts (e.g., SAN 16345, SAN 30438, SAN 36214, SAN
36652, and SAN 97082) and in Sarawak from Bau, Belaga, Bintulu, Kapit, Kuching, Lawas,
Limbang, Lundu, Marudi, Miri, Sibu, Sri Aman, and Tatau districts (e.g., S 13781, S 20883,
S 36684, S 43629, and S 68019). Also occurring in Brunei (e.g., BRUN 3359, Coode MC
6446 and Forman 1166) and Kalimantan (e.g., Ambriansyah AA 2137, bb. 29271 and
Kostermans 12640).
Ecology. Locally common, on clay and silt alluvium river banks along lowland rivers;
occasionally at altitude to 300 m. Occurring in Mulu NP; elsewhere vulnerable owing to
land conversion.
116. Shorea slootenii Wood ex P.S.Ashton

(D.F. van Slooten, 1891–1953, student of dipterocarps at Bogor, Indonesia)
Newman et al. op. cit. (1996) 213. Type: G.H.S. Wood SAN 15160, Borneo, Sabah, Sipitang district,
Mengalong FR (holotype K; isotypes KEP, L, SAN).
Low emergent or main canopy tree, to 40 m tall, to 1.6 m diameter; bole straight,
cylindrical; crown hemispherical, somewhat diffuse, the leaves hanging; buttresses to 1.5 m
tall, stout. Bark becoming dark reddish brown, relatively deeply vsection fissured; inner
bark yellowish brown; heartwood orangebrown. Twig, inflorescence, leaf bud, calyx and
stipule outside, petiole, and leaf blade below densely persistently pale fulvous scabrid
334
tomentose, the hair tufts short, hard and knoblike; midrib above, inside of stipule,
bracteole, parts of petal exposed in bud, ovary, stylopodium, and nut more or less densely
shortly pubescent. Twigs compressed and ribbed at first, becoming terete but remaining
verrucose, stout, to 5 × 3 mm apically; stipule scars long, cuneate, descending. Leaf buds
broadly ovate, 6–10 × 5–8 mm, obtuse, slightly compressed. Stipules oblong, to 25 × 8 mm,
caducous. Leaves coriaceous, deeply concave and more or less corrugated between the
veins, drying rustbrown below; blade oblong, 11–22 × 4–7 cm, base obtuse, apex with
abrupt, tapering acumen to 1.5 cm long; midrib evident but deeply furrowed above, stoutly
prominent below; lateral veins (25–)27–34 pairs, dense, shallowly furrowed above,
prominent below, arched near the margin; intercostal venation scalariform, elevated below;
petiole 1.7–2.3 cm long. Inflorescences terminal or axillary; rachis stout, rigid, brittle, more
or less compressed, to 18 cm long, branchlets short bearing to 4 flowers; bracteoles ovate to
deltoid, obtuse, to 3 × 2 mm. Flowers: buds to 12 × 7 mm; petals pale yellow; stamens 15,
connectival appendage reflexed on 5 inner stamens only; ovary and stylopodium narrowly
ovoid, style equal to ovary, broadly columnar, glabrous. Fruits: calyx shiny; calyx lobes
unequal, 3 longer lobes oblonglorate, to 17 × 1.8 cm, hardly tapering, the base forming a
central incrassate disk bordered by to 4 mm wide lateral auricles, tapering abruptly at the
pedicel, 2 shorter ones linearlobed, to 5.5 × 0.7 cm, hardly auriculate at base. Nuts ovoid,
to 3.5 × 1.3 cm, with to 4 mm long slender tapering style remnant.
Vernacular names. Sabah—seraya kepong kasar (preferred name). Sarawak—meranti

kepong kasar (preferred name).
Distribution. Endemic in Borneo. In Sabah known from Sipitang district (e.g., SAN 15160,
SAN 16922 and SAN A 4570) and in Sarawak from Bintulu, Kuching, Lawas, Lundu, and
Miri districts (e.g., S 1516, S 11053, S 15140, S 27119, and S 44067). Also occurring in
Brunei (e.g., BRUN 3070, BRUN 3175, BRUN 3270, BRUN 5436, and FMS 35560) and W
and E Kalimantan (e.g., bb. 7097 and bb. 14509).
Ecology. Locally frequent in mixed dipterocarp forest on leached thinly humic yellow sandy
and sandy clay soils, on low hills at altitudes to 400 m; mostly in the neogeosyncline (with
one record at Bt. Tanggi) in the central and northern part of its range. Occurring in Lambir
NP; endangered outside parks system.
117. Shorea smithiana Symington Fig. 31, Plates 5A–B.

(J.S. Smith, State Forest Officer in Brunei, 1933–1936)
Gard. Bull. S. S. 9 (1938) 345; Ashton op. cit. (1964) 223, op. cit. (1968) 119, op. cit. (1982) 503;
Meijer & Wood op. cit. 148; Burgess op. cit. 155, 183; Anderson op. cit. (1980) 130; PROSEA op. cit.
403; Coode et al. (eds.) op. cit. 83; Newman et al. op. cit. (1996) 213. Type: Smith FMS 30472,
Borneo, Brunei, Labi Hills (holotype KEP).
Large emergent tree, to 60 m tall, to 1.6 m diameter; bole tall, cylindrical; crown dense,
large, hemispherical; buttresses to 2 m tall and wide, stout, notably straightridged. Bark
dull greyish brown to greyish tawny, becoming vsection fissured and thinly oblong flaky;
dammar in pale creamyellow smears; inner bark dull pinkish brown; heartwood rich
335
Fig. 31. Shorea smithiana. A, flowering leafy twig; B, detail of indumentum on lower leaf
surface; C, flower bud; D, open flower; E, adaxial view of inner sepal; F, adaxial view of
outer sepal; G abaxial view of petal; H, adaxial view of petal with stamens; I, longitudinal
section of gynoecium with sepals and stamens; J, adaxial view of stamens; K, abaxial view
of stamens; L, fruit; M, longitudinal section of fruit. (A–K from S 46524, L–M from SAN
134957.)
336
pinkish brown, relatively soft. Young twig, inflorescence, parts of calyx exposed in bud, leaf
bud, stipule outside, petiole, leaf below, and midrib above persistently greyish brown to rust
scabridpubescent; stipule inside, bracteole, parts of corolla exposed in bud, ovary,
stylopodium, and nut evenly pubescent. Twigs ribbed, stout, 3–4 mm diameter apically;
stipule scars to 2 mm long initially, broad, falcate. Leaf buds ovoid, 5–8 × 3–4.5 mm,
slightly compressed. Stipules broadly hastate, to 20 × 6 mm, caducous. Leaves thickly
coriaceous, drying pinkish brown with greyish lepidote undersurface (mature trees); blade
broadly obovate to oblong, sometimes ovate, 12–20 × 6.5–11.5 cm, base obtuse to
subcordate, apex with broad, tapering acumen to 1.5 cm long; midrib broad, more or less
flat or shallowly furrowed above, prominent below; lateral veins 14–17 pairs, prominent
below, spreading, arched; intercostal venation scalariform, sinuate, elevated below; petiole
2.2–2.8 cm long. Inflorescences terminal or axillary; rachis terete, to 25 cm long, branchlets
bearing to 7 secund flowers; bracteoles ovate, subacute, to 5 × 4 mm, caducous. Flowers:
buds to 10 × 3 mm; petals pink; stamens 22–26, connectival appendage at least 2x the
length of anther, glabrous; ovary and stylopodium narrowly pyriform, style cylindrical,
equal in length to both, glabrous. Fruits: calyx glabrescent; calyx lobes unequal, 3 longer
lobes to 20 × 2.8 cm, tapering to c. 1.3 mm above the saccate base, 2 shorter ones linear
lobed, to 13 × 0.9 cm, similar at base. Nuts ovoid, to 2.7 × 1.8 cm, with to 3.5 mm tapering
style remnant.
Vernacular names. Sabah—seraya timbau (preferred name). Sarawak—meranti rambai

(preferred name).
Distribution. Endemic in Borneo. In Sabah widespread, known from Beaufort,

Kinabatangan, Kudat, Labuk Sugut, Lahad Datu, Ranau, Sandakan, Semporna, Sipitang,
Tawau, and Tenom districts (e.g., SAN 15168, SAN 15420, SAN 16427, SAN 19429, SAN
36587, and SAN 99912) and in Sarawak from Belaga, Bintulu, Lawas, Limbang, Lundu, and
Miri districts (e.g., S 1833, S 24970, S 46531, and S 46763). Also occurring in Brunei (e.g.,
BRUN 5170 and Niga NN 201) and E and S Kalimantan (e.g., Ambriansyah & Priyono AA
2192, bb. 22636, bb. 34403, and bb. 34463).
Ecology. Locally common in mixed dipterocarp forest, on deep welldrained sandy clay
soils, on hills at altitude to 400 m. Occurring in Lambir and Mulu NPs; Danum Valley
Conservation Area and Sepilok FR; elsewhere vulnerable owing to land conversion.
118. Shorea splendida (de Vriese) P.S.Ashton

(Latin, splendidus = splendid)
PROSEA op. cit. 403; Newman et al. op. cit. (1996) 215. Basionym: Hopea splendida de Vriese op.
cit. 28. Lectotype (designated here): de Vriese s.n., ‘Borneo’, Sambas (hololectotype L). Synonym:
Shorea martiniana Scheff. op. cit. 408.
Main canopy tree, to 30 m tall, to 80 cm diameter; bole frequently misshapen; crown oblong
to narrow hemispherical, with pendent branches and distichous hanging leaves; buttresses
low, rounded. Bark smooth, hoopmarked, pale greyish brown; inner bark and heartwood
pale pinkish brown. Parts of corolla and nut densely cream pubescent; all parts otherwise
glabrous. Twigs compressed, ribbed, c. 3 × 1 mm apically; stipule scars to 1 mm thick,
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prominent, amplexicaul. Leaf buds c. 8 × 6 mm, comprising loose aggregation of young

leaves and stipules. Stipules ovate with cordate base, acute, subpersistent, prominent, to 25
× 15 mm. Leaves chartaceous, undulate, drying reddish brown, somewhat shiny; blade
oblong, 8.5–23 × 4.2–11 cm, base typically cordate, sometimes obtuse or cuneate, apex
with slender and tapering acumen to 1 cm long; midrib very slender, evident but shallowly
sunken above, prominent below; lateral veins 9–12 pairs, slender but prominent below,
wellspaced; intercostal venation remotely scalariform, sinuate, elevated below; petiole 1.1–
2.2 cm long. Inflorescences terminal or axillary; rachis terete, to 20 cm long, singly
branched, branchlets to 7 cm long, bearing to 10 flowers; bracts to 18 × 8 mm, caducous.
Flowers: buds to 10 × 3 mm; petals pale pink; filaments connate along ¾ their length,
connectival appendage 4–5x as long as anther; ovary ovoid, glabrous, style and stylopodium
spindleshaped. Fruits: pedicel to 7 mm long, to 4 mm diameter; calyx lobes unequal but
hardly longer than nut, 3 longer lobes to 7.5 × 2.3 cm, tapering to c. 12 mm above the
saccate base, 2 shorter ones to 6.5 × 1.2 cm, otherwise similar. Nuts ovoid, to 5.5 × 3 cm,
acute.
Vernacular names. Sarawak—engkabang bintang (preferred name), engkabang layar

(Iban).
Distribution. Endemic in Borneo. Known in Sarawak from Bau, Kuching, Lundu,

Simunjan, and Sri Aman districts (e.g., S 10307, S 22769, S 27347, and S 37048). Also
occurring in W Kalimantan (e.g., bb. 29663, bb. 29717 and bb. 31419).
Ecology. Locally common, in dense clumps, on floodplains with sweet water. Occurring in
Kubah NP; not vulnerable.
Uses. The secondmost important source of illipe nuts in NW Borneo, after S. macrophylla.
119. Shorea stenoptera Burck

(Greek, steno = narrow, pteron = wing; the short narrow fruit calyx lobes)
Med. Lands Pl. Tuin 3 (1886) 11; Merrill op. cit. (1921) 407; Masamune op. cit. 497; Browne op. cit.
144; Ashton op. cit. (1968) 120, op. cit. (1982) 523; Anderson op. cit. (1980) 103; PROSEA op. cit.
403; Newman et al. op. cit. (1996) 216. Lectotype (designated here): de Vriese s.n., Borneo, W
Kalimantan, Sintang (hololectotype L).
Main canopy tree, to 30 m tall, to 70 cm diameter; crown conical to narrowly hemispherical,

dense, with somewhat pendulous branches; bole straight, tapering, often branching low;
buttresses low, concave, relatively thin. Bark purplish brown and greymottled, smooth,
hoopmarked; inner bark thin, yellowish brown; heartwood pinkish brown. Twig apex, base
of inflorescence, petiole and midrib above densely ochreous puberulent in young trees,
becoming glabrous in mature trees; stipules occasionally sparsely sericeous; nut densely
creambrown puberulent. Twigs somewhat compressed to terete, smooth, 3–4 mm diameter
apically; stipule scars pale, prominent, descendingamplexicaul. Leaf buds ellipsoid, to 4 ×
3 mm, usually obscured within stipules. Stipules ovatedeltoid, obtuse, subauriculate at
base, to 20 × 10 mm, subpersistent. Leaves thickly coriaceous, drying dark reddish
chocolate; blade oblong, 18–40 × 8–22 cm, base broadly cuneate to cordate, apex with
prominent, tapering acumen to 2 cm long; midrib evident, more or less flat above, stoutly
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prominent below as also the lateral veins; lateral veins 10–14 pairs; intercostal venation
remotely scalariform, elevated below; petiole 2.3–4.5 cm long, stout. Inflorescences
terminal or axillary, borne between leaf flushes at the axils of fugaceous rudimentary leaves
along the twig, the twig there with short internodes (visible when not reproducing);
bracteoles ovate, acute, to 6 × 4 mm, not at first caducous. Flowers: buds to 7 × 3 mm;
petals deep pink; stamens 15, filaments connate along ¾ of their length, connectival
appendage 3–4x as long as anther; style and stylopodium spindleshaped. Fruits: pedicel to
4 mm long and diameter; calyx lobes unequal but hardly longer than nut, 3 longer lobes to
7.5 × 2 cm, tapering to 10 mm above the saccate base, 2 shorter ones to 5.5 × 0.8 cm,
similar at base. Nuts ovoid, to 5 × 3 cm, apiculate.
Vernacular name. Sarawak—engkabang rusa (Iban, preferred name).
Distribution. Endemic in Borneo. Recorded in Sarawak from Betong, Kuching, Lundu,

Simunjan, and Sri Aman districts (e.g., S 6506, S 10292, S 18010, S 27349, and S 34397).
Also occurring in W Kalimantan (e.g., the type and bb. 29664).
Ecology. Locally common, on humic soils on seldom flooded sandy alluvium, and in
kerangas forest on more or less poorly drained podsols. Occurring in Bako and Kubah NPs;
probably not vulnerable.
Use. An important source of illipe nuts.
120. Shorea subcylindrica Slooten

(Latin, sub = more or less, cylindricus = cylindrical; the nut shape)
Newman et al. op. cit. (1996) 217. Type: Egon SA 615, Borneo, Sarawak, Semengoh FR (holotype
KEP).
Main canopy tree, to 30 m tall, to 50 cm diameter; bole straight, cylindrical; crown oblong,
dense, irregular, small; buttresses low, rather thin. Bark chocolatebrown mottled with grey,
hoopmarked. Young twigs, leaf bud, inflorescence, parts of calyx exposed in bud, stipule
and bracteole outside, and petiole sparsely caducous pale greyish brown puberulent; parts
of corolla exposed in bud, ovary and nut densely persistently creambrown sericeous to
pubescent. Twigs terete or somewhat compressed, rugulose, c. 2 mm diameter apically. Leaf
buds ovoid, obtuse, c. 3 × 2 mm. Stipules lanceolate, to 10 × 2 mm, fugaceous. Leaves
coriaceous, somewhat shiny, drying tawnybrown below, greenish grey above; blade
narrowly elliptic to lanceolate, 9–24 × 4–12 cm, base cuneate, rarely obtuse, apex
subcaudate, acumen to 1.5 cm long; midrib evident but shallowly furrowed above, stout and
elevated but not prominent below; lateral veins 8–11 pairs, ascending; intercostal venation
subreticulate; petiole 0.9–1.8 cm long. Inflorescences terminal or axillary; rachis terete, lax,
to 12 cm long, to trebly branched, branchlets bearing to 5 flowers; bracteoles deltoid, to 1
mm long, fugaceous. Flowers: buds to 3 × 1 mm; petals cream; connectival appendage
somewhat longer than anther, glabrous; ovary ovoid, without stylopodium, style somewhat
shorter than ovary, sericeous in the basal half. Fruits: pedicel c. 1 mm long; calyx lobes
subequal, ovate, acute, relatively thin, hardly saccate, to 1.2 × 0.9 cm. Nuts ellipsoid, to 2.3
× 1.6 cm, acute.
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Distribution. Endemic in Borneo. Known in Sabah from Beaufort district (e.g., SAN 43270)
and in Sarawak from Kapit, Kuching and Lundu districts (e.g., S 13708, S 29467, S 37781, S
44071, and S 68774). Also occurring in W Kalimantan (e.g., Suzuki 9669 and Suzuki 9690).
Ecology. Very local but common in mixed dipterocarp forest on leached sandy clay soils on
low hills, at altitudes to 200 m. Critically endangered by forest conversion.
121. Shorea superba Symington

(Latin, superbus = magnificent; the stature and elegance of the tree)
Gard. Bull. Sing. 17 (1960) 491; Ashton op. cit. (1964) 146, op. cit. (1968) 75, op. cit. (1982) 459;
Meijer & Wood op. cit. 187; Burgess op. cit. 203, 211; Anderson op. cit. (1980) 121; PROSEA op. cit.
434; Coode et al. (eds.) op. cit. 83; Newman et al. op. cit. (1998) 197. Type: Puasa FMS 38853,
Borneo, Sabah, Sandakan, Betotan (holotype KEP; isotypes L, SAN, SING).
Vast emergent tree, to 75 m tall, to 3.1 m diameter; bole tall, cylindrical; crown vast
cauliflowershaped, diffuse, pale from below; buttresses initially thin, extending to 4 m tall,
concave, stout. Bark yellowish brown, becoming cracked and peeling with small oblong
thin flakes, retaining an overall smooth appearance. Twig, inflorescence, leaf bud, calyx
outside, stipule, bracteole, petiole, veins and midrib on both surfaces more or less densely
evenly pinkish brown puberulent; fruit calyx sparsely so; leaf blade below silvery lepidote.
Twigs compressed and ridged at first, becoming terete, muchbranched, c. 2.5 × 1 mm
apically; stipule scars pale, linear, horizontal. Leaf buds ovoid, to 4 × 3 mm, subacute,
compressed. Stipules ovatelanceolate, to 12 × 5 mm, fugaceous. Leaves thinly coriaceous,
conspicuously silvery lepidote below; blade oblong, 7–12 × 4–7 cm, base broadly cuneate,
apex with broad, tapering acumen to 1 cm long; midrib and veins slender but evident, more
or less narrowly furrowed above, slender but prominent below; lateral veins 16–24 pairs,
dense, straight, appearing like a fish skeleton; intercostal venation densely scalariform,
obscure; petiole 1–1.5 cm long. Inflorescences terminal or axillary; rachis somewhat
compressed and ribbed, to 8 cm long, branchlets bearing to 8 flowers; bracteoles lanceolate,
to 5 mm long, caducous. Flowers: buds to 7 × 2 mm; corolla cream; stamens c. 30,
filaments and anthers glabrous, connectival appendage short, setose; ovary and
stylopodium cylindrical to conical, style short, glabrous. Fruits: calyx lobes unequal, 3
longer lobes to 6 × 1.2 cm, tapering to c. 8 mm at the saccate base, 2 shorter ones linear
lobed, to 1.2 × 0.7 cm, similar at base. Nuts ovoid, to 1.2 × 0.7 cm, apiculate.
Vernacular names. Sabah—selangan batu daun halus (preferred name). Sarawak—

selangan batu tulang ikan (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from Beaufort, Kinabatangan, Labuk

Sugut, Lahad Datu, Sandakan, Sipitang, and Tawau districts (e.g., SAN 16910, SAN 19253,
SAN 27146, SAN 33442, and SAN 61069) and in Sarawak from Lawas, Limbang, Marudi,
and Miri districts (e.g., S 1538, S 1765, S 21411, S 24968, and S 26155). Also occurring in
Brunei (e.g., KEP 80136, S 1660 and S 1661) and W and E Kalimantan.
340
Ecology. Scattered in mixed dipterocarp forest on wellstructured clay soils in moist lower
slope, low hill and floodplain sites. Occurring in Lambir and Mulu NPs, Danum Valley
Conservation Area and Sepilok FR; elsewhere endangered owing to land conversion.
122. Shorea symingtonii Wood

(C.F. Symington, 1905–1943, master of the dipterocarps)
Gard. Bull. Sing. 17 (1960) 493; Meijer & Wood op. cit. 57; Burgess op. cit. 159; Ashton op. cit.
(1982) 495; Newman et al. op. cit. (1996) 218. Type: G.H.S. Wood & Charington FMS 35608 (= SH
16522), Borneo, Sabah, Sandakan district, Sepilok FR (holotype SAN; isotypes KEP, L).
hemispherical, somewhat irregular; buttresses large, stout. Bark greyish brown, cracked and
crumbly flaky; inner bark cream and yellow laminated. Twig apice, leaf bud, stipule,
inflorescence, bracteole, flower calyx outside, and ovary sparsely caducously tawny
puberulent; parts otherwise glabrous. Twigs compressed, stout, c. 4 × 2 mm apically. Leaf
buds ovoidfalcate, acute, c. 4 × 3 mm. Stipules lanceolate, to 12 × 5 mm, fugaceous.
Leaves coriaceous, corrugated between lateral veins, drying yellowish tawny; blade oblong
to narrowly obovate (9–)10–18 × (4–)5–8 cm, base obtuse to cordate, apex with tapering
acumen to 1 cm long; midrib and lateral veins evident but furrowed above, slender but
prominent below; lateral veins dense, 18–22 pairs; intercostal venation densely scalariform,
evident below; petiole stout, 1–2 cm long. Inflorescences terminal or axillary; rachis singly
branched, lax, to 17 cm long, branchlets bearing to 5 flowers; bracteoles ovateelliptic,
acute, to 3 × 2 mm, fugaceous. Flowers: buds large, to 10 × 4 mm; stamens 15, connectival
appendage c. 2½x the length of anther, scarious towards apex; ovary ovoid, without
stylopodium, style stoutly columnar, obscurely trifid apically. Fruits: pedicel c. 5 mm
diameter, stout, expanding into the trumpetshaped receptacle; calyx lobes unequal, 3
longer lobes to 18 × 2.5 cm, tapering into the concave incrassate base, 2 shorter lobes to 13
× 1.5 cm, narrower but otherwise similar. Nuts narrowly ellipsoidovoid, to 2.5 × 1 cm,
glabrous, prominently apiculate, hidden within calyx lobe bases.
Vernacular name. Sabah—melapi kuning (preferred name).
Distribution. Endemic in Borneo; confined to the eastern parts of Sabah, occurring in

Kinabatangan, Lahad Datu, Sandakan, Semporna, and Tawau districts (e.g., SAN 15539,
SAN 16208, 16348, SAN 23713, and SAN 39295).
Ecology. Scattered in mixed dipterocarp forest on undulating land, on clay rich soils.
Occurring in Sepilok FR; possibly endangered.
123. Shorea tenuiramulosa P.S.Ashton

(Latin, tenuis = slender, ramulus = small twig; the slender twig)
Gard. Bull. Sing. 31 (1978) 42, op. cit. (1982) 478; Newman et al. op. cit. (1996) 219. Type: Meijer
SAN 39306, Borneo, Sabah, Karamuak, Bt. Meliau (holotype K; isotypes KEP, SAN).
341
Small canopy tree, to 30 m tall, to 30 cm diameter. Bark greyish brown, smooth.

Inflorescence, stipule, bracteole, calyx outside, ovary, and nut greyish puberulent; caducous
on twig, becoming sparse on fruit calyx lobes. Twigs straight, sparsely branching, terete,
rugulose, slender, pale, to 2 mm diameter apically. Leaf buds and stipule not seen. Leaves
thinly coriaceous, drying pale greyish brown; blade elliptic to lanceolate 9–24 × 4–11 cm,
base broadly cuneate to obtuse, margin prominently undulate, apex shortly broadly
acuminate; midrib prominent on both surfaces; lateral veins 8–9(–11) pairs, very slender
but distinctly elevated below, slightly so above, arched; intercostal venation lax, reticulate;
petiole 1.1–2 cm long, 1.5 mm diameter, drying black but cream towards the end.
Inflorescences terminal, or in axillary cluster of 3, or ramiflorous; rachis slender, many
flowered; bracteoles elliptic, to 2 mm long, fugaceous. Flowers: buds to 5 × 2 mm; stamens
15, anthers narrowly ellipsoid, connectival appendage c. 1½x the length of anther; ovary
ovoid, without stylopodium, tapering into the somewhat shorter columnar glabrous style.
Fruits: subsessile; calyx lobes subequal, oblong, obtuse, incrassate, saccate, patent, to 0.4
× 0.3 cm. Nuts ellipsoidcylindric, to 2.5 × 1.4 cm, apiculate.
Distribution. Endemic in Borneo. Known in Sabah from Lahad Datu, Sandakan and Tawau
Sarawak from Kapit and Lundu district (e.g., S 29666 and S 37815).
Ecology. In mixed dipterocarp forest, on yellow sandy soil in Sarawak, and on shallow
ultrabasic soil near the coast in Sabah. Rare and probably endangered.
124. Shorea teysmanniana Dyer ex Brandis

(J.E. Teijsmann, 1808–1882, Curator of Buitenzorg (now Bogor) Botanic Garden and the
paramount collector of living plants in tropical Asia)
J. Linn. Soc. Bot. 31 (1895) 100; Symington op. cit. (1933) 134, op. cit. (1943) 94; Masamune op. cit.
497; Browne op. cit. 144; Anderson op. cit. (1963) 159, op. cit. (1980) 130; Ashton op. cit. (1964) 225
p.p., op. cit. (1968) 121, op. cit. (1982) 538; Meijer & Wood op. cit. 152; Burgess op. cit. 155, 183;
PROSEA op. cit. 403; Coode et al. (eds.) op. cit. 83; Newman et al. op. cit. (1996) 220. Type:
Teijsmann s.n., Sumatra, Bangka (holotype K). Synonyms: Shorea cochinchinensis Pierre var.
oligoneura Boerl., op. cit. 107; S. balangeroides Boerl., op. cit. 107; S. paludosa Foxw., op. cit.
(1932) 277.
Emergent tree, to 45 m tall, to 1.3 m diameter; bole straight, cylindrical; crown cauliflower
shaped, diffuse; buttresses to 1.5 m tall, somewhat slender. Bark pinkish to chocolate
brown, densely vsection fissured, hardly flaking; inner bark reddish brown; heartwood deep
pinkish red. Young twig, leaf bud, inflorescence, bracteole, and flower calyx outside
persistently greyish brown puberulent; petiole caducously so; ovary, nut and parts of
corolla exposed in bud densely buffpubescent. Twigs compressed at first, smooth, much
branched, c. 2 mm diameter apically; stipule scars pale, prominent, cuneate, 2 mm long.
Leaf buds ovoid, subacute, depressed, 4–8 × 2–5 mm. Stipules ovate to oblong, subacute, to
14 × 5 mm, fugaceous. Leaves coriaceous, glabrous, shiny above, drying rich reddish
brown below; blade ovate, 7.5–11 × 3.5–7 cm, base obtuse, apex with tapering acumen to
0.8 cm long; midrib and lateral veins obscurely sunken above, slender but prominent below;
lateral veins 8–11 pairs, arched, with or without to 3 pairs of small axillary porelike
domatia; intercostal venation scalariform, hardly elevated, obscure; petiole 1.2–1.8 cm
long, slender. Inflorescences terminal or axillary; rachis terete, to 8 cm long, singly
342
branched. Flowers: buds to 14 × 5 mm; stamens 15, connectival appendage short, becoming
reflexed; ovary and stylopodium narrowly conical, style c. ½x the length of ovary and
stylopodium, glabrous. Fruits: calyx lobes unequal, 3 longer lobes to 8 × 1 cm, tapering to
c. 8 mm above the saccate base, 2 shorter ones linearlobed, to 3 × 0.5 cm, similar at base.
Nuts ovoid, to 1.2 × 0.8 cm, with c. 2 mm slender style remnant.
Vernacular names. Sabah—seraya bunga (preferred name). Sarawak—meranti lilin

(preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Sabah rare and confined to
Sipitang district (e.g., SAN 19003 and SAN 27969) but in Sarawak widespread and known
from Kuching, Lawas, Lundu, Marudi, Miri, Sarikei, Sibu, Simunjan, and Sri Aman districts
and FMS 35661).
Ecology. Very local, but often frequent, in mixed peat swamp forest; rare in alan (S. albida)
forest; once recorded at 900 m from the sandstone plateau on G. Bubong Rumah, Lawas,
Sarawak. Recorded from Mulu NP; elsewhere endangered by deforestation.
125. Shorea uliginosa Foxw.

(Latin, uliginosus = growing in swamps; the habitat)
Malay. For. Rec. 10 (1932) 210; Ashton op. cit. (1967) 294, op. cit. (1968) 121, op. cit. (1982) 539;
Anderson op. cit. (1980) 130; PROSEA op. cit. 404; Newman et al. op. cit. (1996) 221. Type:
Foxworthy CF 7938, Peninsular Malaysia, Selangor, Bt. Ceraka FR (holotype KEP). Synonym:
Shorea rugosa (non F.Heim) Symington var. uliginosa Symington op. cit. (1939) 372, op. cit. (1943)
91, Anderson op. cit. (1963) 159.
Large emergent tree, to 45 m tall, to 1.5 m diameter; bole tall, straight, cylindrical; crown
irregularly hemispherical, dense, the twigs and leaves pendent; buttresses to 4 m tall,
spreading, stout. Bark chocolate to purplish brown mottled with fawn and grey, deeply v
section fissured, the ridges eventually becoming powdery chunkily flaky; dammar
incrustations pale yellow; inner bark rich reddish brown; heartwood purplish red. Twig, leaf
bud, stipule outside, inflorescence, calyx outside, bracteole outside, petiole, and blade
below persistently pale chocolatebrown scabridpubescent; stipule within, midrib and veins
above puberulent. Twigs at first prominently ribbed below the petiole insertions, stout, 3–4
mm diameter apically; stipule scars short, pale, horizontal. Leaf buds ovoid, subacute,
compressed, to 6 × 4 mm. Stipules elliptic, subacute, to 14 × 5 mm. Leaves chartaceous,
prominently concave; blade ellipticoblong, 12–22 × 6–12 cm, base broadly cuneate to
subcordate, apex with acumen to 1 cm long; midrib barely evident, furrowed above,
prominent below; lateral veins 16–21 pairs, slender but prominent below, arched near
margins, dense; intercostal venation scalariform, elevated below; petiole 2.2–3.5 cm long.
Inflorescences terminal or axillary; rachis ribbed, to 16 cm long, doubly branched,
branchlets bearing to 6 flowers; bracteole elliptic, subacute to 3 × 2 mm. Flowers: buds to 5
× 3 mm; petals pale yellow; stamens 15, connectival appendage about as long as anther,
becoming reflexed; ovary and stylopodium conical, style the same length, columnar,
glabrous. Fruits: calyx lobes unequal, 3 longer lobes to 6 × 1.2 cm, tapering to 4 mm at the
343
saccate base, 2 shorter ones linearlobed, to 2 cm long, similar at base. Nuts ovoid, to 0.8 ×
0.6 cm, acute.
Vernacular name. Sarawak—meranti buaya (preferred name).
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo known in Sarawak from
Bintulu, Marudi, Miri, Sarikei, Sibu, and Sri Aman districts (e.g., S 7908, S 14507, S 23287,
S 27901, and S 32001). Also occurring in W Kalimantan.
Ecology. Formerly locally abundant in the mixed peat swamp forests of the Lower Rajang,
local and rare elsewhere; rare in alan (S. albida) forest. Endangered.
126. Shorea venulosa Wood ex Meijer

(Latin, venulosus = with tiny veins; the slender leaf venation)
Act. Bot. Neerl. 12 (1963) 342; Wood & Meijer op. cit. 153; Burgess op. cit. 166; Ashton op. cit.
(1964) 510, op. cit. (1968) 122, op. cit. (1982) 510; Anderson op. cit. (1980) 130; PROSEA op. cit.
404; Coode et al. (eds.) op. cit. 83; Newman et al. op. cit. (1996) 222. Lectotype (designated here):
G.H.S. Wood SAN 15134, Borneo, Sabah, Sipitang district, Menggalong FR (hololectotype K;
isolectotypes KEP, SAN).
Large emergent tree, to 55 m tall, to 1.3 m diameter; bole straight, often somewhat tapering;
crown cauliflowershaped, diffuse; buttresses to 3 m tall, stout, prominent. Bark becoming
dark chocolatebrown, deeply vsection fissured, the ridges becoming oblong flaky; dammar
incrustations pale yellow; inner bark yellowish brown; heartwood dark brown. Young parts
at first grey sericeous, fugaceous except on leaf bud and stipule outside; inflorescence, parts
of perianth exposed in bud, bracteole, ovary, and nut towards apex persistently cream
puberulent. Twigs terete, muchbranched, slender, c. 1 mm diameter apically; stipule scars
pale, more or less horizontal, amplexicaul. Leaf buds lanceolatefalcate, to 7 × 2 mm.
Stipules hastate, to 20 × 3.5 mm, fugaceous. Leaves coriaceous, drying chocolatebrown,
often shiny above; blade ovate, 6–10 × 3–5 cm (sometimes smaller on exposed peaks, larger
on E Sabah ultrabasics), base obtuse, margin sometimes narrowly revolute, apex with
slender acumen to 0.6 cm long; midrib obscure, sunken above, slender but prominent below;
lateral veins 15–18 pairs, very slender, hardly elevated as also the scalariform intercostal
venation, often with small glabrous porelike, sometimes prominent and pustular, axillary
domatia; petiole 1.7–3 cm long, slender, geniculate. Inflorescences terminal or axillary;
rachis terete, slender, lax, to 11 cm long, branchlets short, bearing to 10 flowers; bracteoles
ovate, obtuse, to 3.5 × 2 mm. Flowers: buds to 5 × 2.5 mm; corolla pink; stamens 15,
connectival appendage more than 2x the length of anther; ovary ovoid, without
stylopodium, style somewhat shorter, filiform, puberulent at base. Fruits: calyx lobes
unequal, chartaceous, 3 longer lobes to 6 × 1.3 cm, tapering to c. 3.5 mm above the saccate
base, 2 shorter lobes linearlobed, to 3 × 0.3 cm, similar at base. Nuts narrowly ovoid, to 2
× 0.8 cm, acute.
344
Vernacular names. Sabah—seraya kerangas (preferred name). Sarawak—meranti tangkai

panjang padi (preferred name).
Distribution. Endemic in Borneo. In Sabah widespread and recorded from Keningau,

Kinabatangan, Kota Marudu, Labuk Sugut, Lahad Datu, Ranau, Sandakan, Sipitang,
Tambunan, and Tawau districts (e.g., SAN 15501, SAN 16359, SAN 26168, SAN 37855, and
SAN 39323) and in Sarawak from Bau, Kapit, Kuching, Lawas, Limbang, Lundu, Marudi,
and Mukah districts (e.g., S 10128, S 23287, S 26552, S 28977, and S 60087).
Ecology. Locally common, on podsols in kerangas forest, and forest on ultrabasic rock
throughout E Sabah; also common in lower montane kerangas at 1000–1600 m altitude,
including on the Batu Tibang andesite. Occurring in Bako, Kinabalu and Mulu NPs;
vulnerable elsewhere in the lowlands, less so on the mountains.
127. Shorea virescens Parijs

(Latin, virescens = becoming green; inference unclear)
In Fedde, Rep. 33 (1933) 244; Slooten op. cit. (1949) 240; Rojo, Kalikasan 5 (1976) 99; Ashton op.
cit. (1978) 44, op. cit. (1982) 492; Anderson op. cit. (1980) 124; PROSEA op. cit. 413; Coode et al.
(eds.) op. cit. 83; Newman et al. op. cit. (1996) 223. Type: Frijd bb. 13898, Borneo, W Kalimantan,
Sanggau (holotype L; isotype BO).
Large emergent tree, to 50 m tall, to 1.6 m diameter; crown hemispherical, diffuse, large;
bole tall, cylindrical; buttresses to 3 m tall, stout, prominent. Bark greyish tawny, becoming
shaggily flaky, the flakes adhering and thick; dammar incrustations cream; inner bark cream
and yellow laminated. Twig, inflorescence, leaf bud, parts of perianth exposed in bud,
bracteole, stipule, petiole, midrib on both surfaces, and venation below persistently densely
greyish buffpubescent; ovary minutely puberulent; nut glabrescent. Twigs compressed,
stout, 2.4–4 × 1.5–3 mm apically; stipule scars to 2.5 mm long, horizontal. Leaf buds
globose to stoutly ovoid, 2–3 × 4 mm. Stipules linear, acute, to 25 × 4 mm, caducous.
Leaves coriaceous, drying dull tawnygrey; blade obovate, 8–15 × 4–8 cm, base obtuse to
subcordate, apex with broad acumen to 7.5 mm long; midrib evident but deeply furrowed
above, slender but prominent below as also the lateral veins; lateral veins 20–26 pairs,
dense, ascending; intercostal venation very slender, scalariform; petiole 1.5–2 cm long.
Inflorescences terminal or axillary; rachis more or less compressed, to 10 cm long, singly
branched; bracteoles narrowly lanceolate, to 8 mm long. Flowers: buds to 6 × 3 mm;
stamens 15, connectival appendage c. 2x the length of anther; ovary ovoid, without
stylopodium, style filiform, equal to ovary. Fruits: calyx lobes unequal, 3 longer lobes to 8
× 1.3 cm, tapering to 6 mm above the saccate base, 2 shorter ones linearlobed, to 5.5 × 0.5
cm, similar at base. Nuts ovoid, to 2.2 × 1.2 cm, with c. 3 mm long tapering style remnant.
Vernacular name. Sarawak—meranti sulang sulang (preferred name).
Distribution. Borneo and the Philippines. In Sabah uncommon and known only from Lahad
Datu, Sandakan and Tawau districts (e.g., SAN 19573, SAN 34788 and SAN 62950) and in
Sarawak recorded from Kapit, Lawas, Marudi, Miri, Serian, and Tatau districts (e.g., S
29661 and S 41149). Also occurring in Brunei (e.g., BRUN 323 and SAN 17067).
345
Ecology. Rare and scattered in mixed dipterocarp forest on clay soils, at altitude below 600
m, especially on low hills and slopes. Occurring in Lambir and Mulu NPs; highly vulnerable
elsewhere owing to land conversion.
Note. The species was formerly confused with S. confusa and S. lamellata by me (Ashton
op. cit. (1964) 164, 167, Meijer & Wood op. cit. 60, 66 and Burgess op. cit. 159).
128. Shorea waltonii Wood ex Meijer

(A.B. Walton, 1904–?, Conservator of Forest at Sandakan, 1952–1954)
Act. Bot. Neerl. 12 (1963) 344; Wood & Meijer op. cit. 155; Burgess op. cit. 182; Ashton op. cit.
(1982) 510; Newman et al. op. cit. (1996) 224. Type: G.H.S. Wood SAN 16508, Borneo, Sabah,
Sandakan district, Sepilok FR (holotype K; isotypes A, KEP, SAN).
Large emergent tree; bole straight; crown spreading; buttresses large, stout. Bark deeply
fissured and thinly oblong flaky; inner bark pale reddish brown; heartwood pink. Twigs, leaf
bud, stipule, petiole, venation below, and inflorescence more or less densely cream lepidote;
flower calyx caducously so; ovary, nut, parts of petals exposed in bud, midrib, and
sometimes veins above persistently creambrown puberulent. Twigs terete, smooth, stout, c.
3 mm diameter apically. Leaf buds small, conical. Stipules ellipticlanceolate, acute, to 20 ×
8 mm. Leaves coriaceous, drying cream lepidote to mauvebrown below; blade oblong to
ovate or obovate, 10–25 × 5.5–12 cm, base obtuse or shallowly cordate, apex abruptly
acuminate, acumen to 1 cm long; midrib very slender but evident, deeply furrowed above,
slender but prominent below as also the lateral veins; lateral veins 18–22 pairs, ascending
except at base; intercostal venation remotely scalariform, hardly elevated below; petiole
stout, 2.5–4.5 cm long. Inflorescences stout, terminal or axillary; rachis to 12 cm long,
branchlets to 5 cm long; bracteoles to 10 cm long, amplexicaul. Flowers: buds to 8 × 3 mm;
stamens 15, connectival appendage c. 2x the length of anther; ovary broadly ovoid, without
stylopodium, style of equal length, puberulent in the basal half. Fruits: pedicel c. 3 mm
long, stout; calyx lobes unequal, 3 major lobes to 14 × 2.3 cm, tapering to c. 11 mm wide
above the saccate base, 2 shorter ones loratelobed, to 10 × 0.8 cm, similar at base. Nuts
ovoid, to 2.5 × 1.8 cm, apiculate.
Vernacular name. Sabah—seraya kelabu (preferred name).
Distribution. Endemic in Borneo. In Sabah recorded from to Kinabatangan, Lahad Datu

and Sandakan districts (e.g., SAN 16903, SAN 17848, SAN 34486, SAN 36207, and SAN A
2985). Also occurring in E Kalimantan (e.g., Kostermans 13816).
Ecology. Locally common in mixed dipterocarp forest on clay soils on low hills. Occurring
in Sepilok FR; elsewhere endangered by forest clearance.
346
129. Shorea woodii P.S.Ashton

(G.H.S. Wood, forest botanist in Sabah 1953–1957 and a dipterocarp specialist)
TFSS 5 (2004) 480, op. cit. (1968) 122 (‘63. Shorea sp.’). Type: s.c. A 56/36, Sarawak (holotype
FHO).
Mediumsized tree, to 40 m tall, to 60 cm diameter; bole frequently branching low. Bark at

first smooth, hoopmarked, eventually shallowly patchily fissured, with bright yellow
dammar smears. Young twig and petiole evenly densely golden brown caducous sericeous;
parts otherwise glabrous. Twigs 3–4 mm diameter apically, terete to somewhat compressed,
becoming pale brown, smooth; internodes to 9 cm long; stipule scars amplexicaul, down
curved, prominent. Leaf buds ovoid, to 3 × 2 mm. Stipules lanceolate, acute, to 40 × 13
mm, rich crimsonred. Leaves rich crimsonred when young, coriaceous, drying greyish
brown; blade ovateelliptic, 20–37 × 11–20 cm, base to 1 cm peltate, apex tapering to 1.5
cm prominent slender acumen; midrib slightly raised above, prominently terete below;
lateral veins 12–19 pairs, slender but prominent below; intercostal venation remotely
scalariform, slender; petiole 3–7 cm long, terete, distally rugose. Inflorescences and
flowers unknown. Fruits: 3 longer calyx lobes narrowly spatulate, to 20 × 1.5 cm, obtuse,
tapering to c. 5 mm above the saccate base; 2 sorter ones unequal, to 17 × 0.8 cm, otherwise
similar. Nuts ovoid, lustrons, to 3 × 2 cm, tapering to an apiculate style remnant.
Vernacular name. Sarawak—engkabang lajan (Iban).
Distribution. Endemic in Borneo; known only in Sarawak from Belaga and Kapit districts
(e.g., S 17801 and S 22318).
Ecology. Rare, as scattered individuals and small groups in mixed dipterocarp forest on
damp sandy clay soils, on hillsides, at altitudes below 200 m. Critically endangered by land
conversion.
Notes. Named after Geoffry Howarth Spencer Wood, who introduced me to the
dipterocarps but whose inspiring stay with me in Brunei ended in tragedy.
130. Shorea xanthophylla Symington

(Greek, xanthos = yellow, phullon = leaf; the yellowish dry leaf)
Gard. Bull. S. S. 9 (1938) 342; Browne op. cit. 164; Slooten op. cit. (1956) 344; Ashton op. cit. (1964)
160, op. cit. (1968) 91, op. cit. (1982) 479; Meijer & Wood op. cit. (1964) 79; Burgess op. cit. 218;
cit. (1996) 225. Type: Puasa FMS 36776 (= SH 3998), Borneo, Sabah, Sandakan district, Betotan
(holotype KEP).
Canopy tree, to 40 m tall, to 65 cm diameter; bole straight or often misshapen; crown dense,
irregular, twigs somewhat pendent; buttress to 1 m tall, somewhat stout. Bark reddish fawn,
smooth, hoopmarked, eventually vertically cracked and patchily thinly oblongflaky; with
small snaillike blackish dammar exudations. Young twig and petiole buff caducous
347
puberulent; inflorescence, stipule and bractole outside, leaf bud, parts of perianth exposed
in bud, ovary, and nut persistently cream puberulent. Twigs straight, terete, stout, drying
black, 2–3 mm diameter apically. Leaf buds conical, to 3 × 1.5 mm. Stipules narrowly
deltoid, concave, c. 8 × 3 mm. Leaves bright crimson on opening, thinly coriaceous, often
somewhat blistered between the veins, shiny, drying tawny brown; blade oblonglanceolate,
12–25 × 4–7 cm, base subcordate, obtuse or broadly cuneate, margin often narrowly
subrevolute, apex with tapering acumen to 1 cm long; midrib broadly evident, more or less
flat above, terete, prominent below; lateral veins 9–13 pairs, slender, raised below, arched
and continuing within the margin, sometimes anastomosing to form a looped intramarginal
vein; intercostal venation remotely subscalariform; petiole 0.8–1.5 cm long, drying black.
Inflorescences terminal, or in axillary cluster of 3, or ramiflorous; rachis angular, to 20 cm
long, branchlets zigzag bearing to 9 flowers; bracteoles minute, fugaceous. Flowers: buds to
4.5 × 1.5 mm; petals creamyellow; stamens 15, connectival appendage 1–2x the length of
anther, ciliate distally; ovary ovoidconical, tapering into stylopodium, style short, glabrous.
Fruits: calyx lobes subequal, broadly ovate, thickened, somewhat patent, saccate, c. 0.7 ×
0.7 cm. Nuts obovoid, c. 2 × 1.3 cm, subacute.
Vernacular names. Sabah—seraya kuning barun (preferred name). Sarawak—lun barun

(preferred name).
Distribution. Endemic in Borneo. In Sabah known from Beaufort, Kinabatangan, Kota

Merudu, Kudat, Labuk Sugut, Lahad Datu, Ranau, Sandakan, Tawau, and Tenom districts
from Belaga, Bintulu, Kapit, Miri, Sri Aman, and Tatau districts (e.g., S 21858, S 27543, S
29099, S 43639, and S 46514). Also occurring in Brunei (e.g., BRUN 2605 and BRUN
3126).
Ecology. Locally common in mixed dipterocarp forest on clay soils over shale and basic
volcanic rock at low altitude, rarely to 1000 m. Occurring in Lambir NP and Sepilok FR;
vulnerable elsewhere owing to land conversion.
8. UPUNA Symington
(Iban name—upun)
Bull. Jard. Bot. Buitenz. 3, 17 (1941) 88; Meijer & Wood, Sabah For. Rec. 5 (1964) 327; Ashton,
MDB (1964) 7, MDBS (1968) 3, FM 1, 9 (1982) 337; PROSEA 5, 1 (1993) 458.
Large main canopy trees with low stout buttresses in groups. Bark dark brown, finely flaky;
inner bark not laminated. Young parts caducous, inflorescence persistently multicellular
glandular tomentose and tufted tomentose. Stipules lineartapering, persistent and
conceiling the small leaf buds. Inflorescences cymose. Flowers: sepals imbricate but fused
into a shallow cup at base free from ovary; stamens 25–30; filaments compressed, dilated at
base and tapering to anthers; anthers oblongovoid, latrorse; connectival appendage aristate,
erect, many times the length of anther; ovary small, ovoid, without stylopodium; style
filiform, about twice as long as ovary, trifid towards apex; stigma minute. Fruits: calyx
with a distinct basal cup enclosing the base of but free from nut; lobes valvate, thinly
chartaceous, 2 much larger than the other 3. Nuts ellipsoid, triangular in crosssection,
tapering at base, splitting loculicidally into 3 valves at germination. Seeds with arillode.
Germination epigeal; cotyledons subequal, cordate.
348
Distribution. A monotypic genus endemic in Borneo.
Uses. A heavy hardwood similar to resak but available in larger sizes. The timber is greasy
and does not easily take glues.
Upuna borneensis Symington Fig. 32, Plates 5C–D.

(of Borneo)
Bull. Jard. Bot. Buitenz. 3, 17 (1941) 88; Browne, FTSB (1955) 171; Ashton op. cit. (1964) 7, op. cit.
(1968) 3, op. cit. (1982) 339; Meijer & Wood op. cit. 327; Burgess, TBS (1966) 228; Anderson, CLTS
(1980) 131; PROSEA op. cit. 458; Coode et al. (eds.), CLBD (1996) 84; Newman et al., MDFB
MHHW (1998) 201. Type: Flemmich FMS 48158, Borneo, Brunei, Sg. MauKargu headwaters
(holotype KEP).
Large main canopy, occasionally low emergent tree, to 45 m tall, to 1.5 m diameter; crown
dark, dense, hemispherical; buttresses low and concave but stout and grouped around the
base of the bole. Bark dark brown, densely cracked and flaking in narrow thin oblong
pieces; inner bark cream, homogeneous. Sapwood pale yellow, hard; heartwood coffee
brown; dammar exudations pale yellow, opaque. Young parts caducous multicellular
glandular tomentose; exposed living surfaces, leaf blade surfaces excepted, and ovary
densely persistently pale chocolatebrown tomentose; blade undersurface whitetomentose,
upper surface glabrescent. Twigs to 3.5 mm diameter apically; stipule scars small. Leaf
buds ovoidfalcate, to 12 × 6 mm, subacute. Stipules linear, to 12 mm long. Leaves thinly
coriaceous, frequently somewhat bullate; blade oblong to ovate, 9–17 × 4–9.5 cm, base
cordate, margin revolute, apex with tapering acumen to 0.5 cm long; midrib prominent
below, sunken above; lateral veins 16–20 pairs, wellspaced, arched towards margin,
prominent below, frequently with short intermediates; intercostal venation scalariform,
wellspaced, distinctly raised below; petiole 1–2.5 cm long, distinctly geniculate.
Inflorescences to 3axillary, to 15 cm long, muchbranched; bracts lanceolate, to 10 × 3.5
mm, caducous. Flowers: buds narrowly ovoid, to 5 × 2 mm; sepals subequal, imbricate, the
3 inner ones more attenuate, united at the base forming a shallow cup free from the ovary;
corolla deep purple with dark yellow margin, petals broadly ovate, subacute, becoming
reflexed apically at anthesis; stamens 25–30, filaments broad at base, tapering abruptly and
filiform at the subglobose anthers, connectival appendage c. 3x the length of anther,
aristate; ovary ovoid, without stylopodium, style c. 3x as long as ovary, filiform, more or
less trifurcate at apex. Fruits: calyx remaining sparsely pubescent towards base; lobes
chartaceous, united into a c. 1 cm wide and deep conical basal cup tapering into a short
pedicel, 2 outer lobes lanceolate, to 13 × 2.7 cm, tapering gradually to subacute apex, 3
inner ones to 7.5 × 1.7 cm, otherwise similar, subequal. Nuts narrowly ovoid, to 3.2 × 1.5
cm, triangular in crosssection, tapering to 5 mm slander style remnant.
Vernacular names. Sabah and Sarawak—upun (preferred name). Sarawak—penyau (Iban),

upun batu (Iban).
Distribution. Widespread in its chosen habitat, hence absent in most parts of Sabah and
coastal E Kalimantan. In Sabah confined to SW parts and known from Sipitang district (e.g.,
SAN 15184) and in Sarawak widespread and recorded from Bintulu, Kuching, Lawas,
Lundu, and Serian districts (e.g., Hotta 15808, S 1522, S 6876, and S 10064). Also occurring
in Brunei (e.g., BRUN 3091, FMS 37075 and FMS 48457) and W and SE Kalimantan (e.g.,
bb. 29206 and bb. 30145).
349
Fig. 32. Upuna borneensis. A, fruiting leafy twig; B, detail of indumentum on lower leaf
surface; C, flower bud; D, gynoecium and stamens; E, longitudinal section of gynoecium; F,
adaxial view of outer sepal; G, adaxial view of inner sepal; H, adaxial view of petal; I,
abaxial view of petal; J, abaxial view of stamens; K, crosssection of ovary; L, fruit; M, fruit
with exposed nut; N, nut; O, dissected nut to epose the embryo; P, crosssection of nut. (A–
B from KEP 19288, C–K from S 64513, L–N from S 26237, O–P from Niga NN 221.)
350
Ecology. Scattered, locally frequent in mixed dipterocarp forest and the ecotone to
kerangas, on deep yellow sandy soil on low hills, on sandstone and acid volcanics including
the Arip rhyolite, at altitudes to 400 m. Occurring locally in Bako NP and recorded from
Kubah NP; elsewhere vulnerable.
9. VATICA L.
(Latin, vates = prophet; alluding the use of dammar of one of the
Indian species in Hindu religions ceremony)
resak (preferred name)
Mant. 2 (1771)152; Brandis, J. Linn. Soc. Bot. 31 (1895) 128; Merrill, EB (1921) 408; Slooten, Bull.
Jard. Bot. Buitenz. 3, 9 (1927) 67; Masamune, EPB (1942) 297; Browne, FTSB (1955) 96; Ashton,
Gard. Bull. Sing. 20 (1963) 243; MDB (1964) 61, MDBS (1968) 25, FM 1, 9 (1982) 345; Meijer &
Wood, Sabah For. Rec. 5 (1964) 301; Burgess, TBS (1966) 226; Anderson, CLTS (1980) 131;
PROSEA 5, 1 (1993) 461; Coode et al. (eds.), CLBD (1996) 84. Synonyms: Retinodendron Korth.,
Kruidk. (1840) 55; Isauxis Reichb., Nom. (1841) 210; Pteranthera Blume, Mus. Bot. Lugd.Bat. 2
(1852) 30; Sunaptea Griff., Notul. 4 (1854) 516; Synaptea Kurz, J. As. Soc. Beng. 39, 2 (1870) 65;
Pachynocarpus Hook.f., Trans. Linn. Soc. 23 (1860) 159; Elaeogyne Miq., Fl. Ned. Ind., Suppl.
(1862) 460; Retinodendropsis F.Heim, C. R. Assoc. Fr. Pau 1892 (1893) 470; Perissandra Gagnep.,
Bull. Soc. Bot. Fr. 95 (1948) 27; Brachypodandra Gagnep. op. cit. 30.
Subcanopy to main canopy trees; bole frequently crooked; buttresses stout, usually low,
concave; crown usually irregular, diffuse, juveniles irregularly branched. Bark usually
greyish brown mottled, smooth and hoopmarked, in large trees becoming patchily flaked
and occasionally scrollmarked, with small lenticels hardly paler than bark surface; inner
bark pale creambrown, finegrained, homogeneous. Sapwood pale yellow, dense, fine
textured with ray ends not glistening on tangential surface, hard or rarely soft; heartwood
brown; dammar not generally present on bole, as whitish opaque smears if present. Young
parts usually more or less caducous powdery pubescent. Leaf buds generally small, ovoid.
Twigs usually pale greyish brown, smooth or rugose. Leaf buds generally small. Stipules
variable. Leaves variable; blade more or less coriaceous, flat or bullate; midrib evident,
usually more or less raised above though sometimes within a furrow (obscurely sunken in V.
rynchocarpa); lateral veins arched, distinctly raised but not usually prominent below, with
short or no intermediate veins, not or obscurely joined into an intramarginal vein;
intercostal venation usually reticulate but sometimes scalariform, more or less elevated on
both surfaces, not drying darker than the leaf blade; petioles variably but usually not
geniculate. Inflorescences irregularly branched, racemose or partly cymose. Flowers: buds
ovoid, spindleshaped to lanceolate; sepals more or less valvate, subequal; petals narrowly
oblong, usually cream often with a violet tinge, slightly contorted spreading at anthesis, not
connate and falling separately; stamens to 15 in 3 whorls, filaments short, dilated at base,
anthers broadly oblong, glabrous, connectival appendage at most 1½x the length of anther,
stout, deltoid; ovary ovoidconical, superior or semiinferior, pubescent, without
stylopodium, style stoutly columnar, less than 2x the length of ovary, somewhat expanded
apically with a conical 3lobed stigma. Fruits: calyx very variable; lobes valvate and free to
the base or united into a shallow cup enclosing less than 1/3 of the nut, if free 2 are longer
than the other 3, or all equal and revolute along the axis, or short, equal, reflexed to rotate,
chartaceous to woody. Nuts variable, small or large, globose to ovoid with no persistent
style remnant, round in crosssection, thinwalled or corky; pericarp splitting loculicidally
into 3 equal valves at germination. Germination epigeal, or hypogeal and cryptocotylar;
351
cotyledons, if free, foliose, subequal, hardly photosynthetic, if remaining in the nut more or
less fleshy, unequal.
Distribution. About 65 species, from South Asia and IndoBurma to New Guinea. Thirty
five species occur in Borneo, of which 23 are endemic and 33 occurring in Sabah and
Sarawak.
Ecology. From brackish river banks to the lower edge of the upper montane forest, at
altitudes to 1600 m; in all inland forest types except the alan peat swamps. Sometimes
scattered, but usually quite common in the subcanopy, one or a small number of species
common in any locality.
Uses. The larger hardwooded species are used locally in construction. The seeds of the
large fruited species were once used, though not apparently in Malaysia, for arresting the
fermentation of rice and coconut wine.
Notes. Vatica species are the most difficult dipterocarps to recognise, and this explains why
this common genus in our forests remains so poorly known. The smooth bark, absence of
dammar smears, irregularly branched diffuse crown, and often nondescript leaves with
reticulate intercostal venation can recall Aporosa or Hydnocarpus; but the scent of dammar
is always descernable in freshly rubbed living tissue.
Key to Vatica species

1. Fruit calyx lobes equal or subequal, chartaceous or coriaceous, often becoming reflexed
(sect. Vatica)……………………………………………………………………………2
Fruit calyx lobes unequal, 2 longer than the other 3, chartaceous, not becoming reflexed
(sect. Sunaptea)…………………………………………………………………….…15
2. Fruit calyx lobes thickened, corky, coalescing and adnate to nut forming a cup more or
less enclosing it…………………………………………………….…31. V. umbonata
Fruit calyx lobes free……………………………………………………………………3
3. Fruit calyx lobes revolute but not reflexed, foliaceous and concealing the nut,
subcordate................................................................................................................…4
Fruit calyx lobes more or less reflexed or spreading or rotate not concealing nut, not
subcordate…………………………………………………………………………….…6
4. Lateral veins at most 9 pairs, without distinct intermediates……………………………..

……………………………………………………..32. V. venulosa (subsp. venulosa)
Lateral veins at least 10 pairs, with distinct intermediates………………………….....5
5. Twig and inflorescence caducous buffpuberulent; petiole glabrous...….6. V. chartacea

Twig, petiole and inflorescence persistently fulvous somewhat flocculent tomentose;
petiole otherwise………………………………………………….…15. V. havilandii
352
6. Nut ovoid, verrucose, 2.5–4 cm diameter…………………………………………..…7

Nut more or less globose, smooth, at most 2.2 cm diameter………………………..…9
7. Nut asymmetric; petiole at least 2 cm long…………………………….....27. V. rassak

Nut symmetric; petiole at most 2 cm long………………………………………….…8
8. Leaf blade to 20 × 7 cm; lateral veins at most 14 pairs…………..….14. V. granulata

Leaf blade at least 22 × 7 cm; lateral veins at least 15 pairs….…30. V. sarawakensis
9. All parts glabrous……………………………………………………..…1. V. albiramis

Young parts and nut, at least, powdery tomentose to vinoussericeous……………....10
10. Tomentum rufous; leaves drying greyish to pinkish brown…………….13. V. globosa

Tomentum vinous; leaves at first drying bluish greyish green or mauvebrown……11
11. Twigs compressed……………………………………………….…21. V. oblongifolia

Twigs terete…………………………………………………………………….…12
12. Fruit calyx lobes chartaceous, hardly revolute…………………………10. V. dulitensis

Fruit calyx lobes incrassate, becoming revolute…………………………………….…13
13. Nut to 2.2 cm diameter………………………………………………25. V. pedicellata

Nut at most 1 cm diameter………………………………………………………….14
14. Lateral veins 9–11 pairs……………………………………………………28. V. rotata

Lateral veins at least 12 pairs…………………………………………..…33. V. vinosa
15. Fruit calyx lobes fused into a cup at base adnate to the nut……………………….…16
Fruit calyx lobes free………………………………………………………………...…17
16. Lateral veins 7–10 pairs………………………………………………18. V. maritima

Lateral veins 11–15 pairs…………………………………………….…22. V. odorata
17. Twigs at first compressed………………………………………………7. V. compressa

Twigs terete……………………………………………………………………….…18
18. Leaf blade obovate, thickly coriaceous, apex usually obtuse or retuse, margin revolute
................................................................................................................................…19
Leaf blade elliptic, lanceolate, oblanceolate, ovate or obovate, chartaceous, thinly
coriaceous, or coriaceous, apex acuminate to subcaudate, margin not revolute………20
19. Inflorescence at most 3 cm long, fascicled and congested; petiole 1.2–2.5 cm long…….
………………………………………………………………………….8. V. congesta
Inflorescence to 20 cm long, lax; petiole 1–1.5 cm long……………..…9. V. coriacea
20. Leaf undersurface persistently ochreous sericeous………………………5. V. brunigii

Leaf undersurface at most caducous pubescent………………………………………21
21. Lateral veins hardly or not more prominent below than above…………………….…22
Lateral veins distinctly more prominent below than above…………………………26
353
22. Petiole at least 1 cm long………………………………………………………….….23

Petiole at most 1 cm long………………………………………………………………24
23. Petiole, twig and leaf bud at first creampuberulent, glabrescent; leaf blade drying
greyish green to honeycoloured, often shiny……………………..…12. V. glabrata
Petiole, twig and leaf bud more or less densely persistently pinkish brown pubescent;
leaf blade drying dull rich pinkish brown………………………….…3. V. borneensis
24. Ripe nut narrowly ovoid, tapering, acute, glabrous……………..…29. V. rynchocarpa

Ripe nut subglobose or broadly ovoid, obtuse or shortly mucronate, densely pubescent..
……………………………………………………………………………………..…25
25. Midrib raised above; leaf base cuneate……………………………17. V. mangachapoi

Midrib flat or slightly furrowed above; leaf base obtuse………….…23. V. parvifolia
26. Inflorescence, petiole, lateral veins and midrib below persistently shortly pale brown
scabridpubescent………………………………………………………………….…27
Inflorescence and sometimes petiole pink to rufous tomentose; lateral veins and midrib
below caducously so, glabrescent……………………………………………………30
27. Inflorescence ramiflorous, fascicled………………………….……24. V. patentinervia

Inflorescence axillary, not fascicled……………………………………………..…28
28. Petiole to 2 cm long…………………………………………………..…11. V. endertii

Petiole at most 1 cm long……………………………………………………….…29
29. Leaf ellipticobovate, base narrowly cuneate……………………….…4. V. brevipes

Leaf ellipticoblong to lanceolate, base obtuse………………….…19. V. micrantha
30. Leaf blade narrowly oblongobovate, with many intermediate veins……...20. V. nitens
Leaf blade elliptic, lanceolate, oblanceolate, broadly elliptic, or obovate, with a few or
without intermediate veins………………………………………………………..…31
31. Lateral veins prominently raised below; tomentum flocculent, long.......16. V. maingayi
Lateral veins not prominent below; tomentum even, short………………………..…32
32. Petiole at least 2 cm long; leaf blade drying rich pinkish brown, glistening; fruit calyx
lobes to 8 × 2.5 cm……………………………………………………2. V. badiifolia
Petiole at most 2 cm long; leaf blade drying dull greyish brown; fruit calyx lobes to 6 ×
1.8 cm………………………………………………………………..26. V. perakensis
Key to Vatica species

1. Petiole glabrous or glabrescent, drying black………………………………………...…2

Petiole cinereous or tomentose, not drying black………………………………….…9
354
2. Lateral veins at most 6 pairs……………………………………………1. V. albiramis

Lateral veins at least 7 pairs……………………………………………………….…3
3. Leaf blade oblongobovate………………………………………………………..…4

Leaf blade ovate, elliptic or lanceolate………………………………………………5
4. Leaf blade narrowly obovate, thickly coriaceous, concave, apex obtuse to retuse,
margins revolute; petiole stout. In kerangas into the mixed dipterocarp forest
ecotone…………………………………………………….…9. V. coriacea (in part)
Leaf blade oblongobovate, chartaceous, flat, apex prominently acuminate; petiole
slender. In moist valleys and on river banks………………………..…6. V. chartacea
5. Midrib sunken above……………………………………29. V. rynchocarpa (in part)

Midrib flat or raised above……………………………………………………………6
6. Lateral veins raised on both surfaces. In upper dipterocarp forests at altitudes above
1000 m……………………………………………………………..…12. V. glabrata
Lateral veins unraised above. In lowland mixed dipterocarp forest………………..…7
7. Leaf blade drying rustbrown; lateral veins 9–11 pairs, sharply prominent below………
………………………………………………………………..16. V. maingayi (in part)
Leaf blade drying pale greyish or yellowishbrown; lateral veins 7–9 pairs, raised but
not prominent below……………………………………………………………………8
8. Leaf blade c. 12 × 5 cm; midrib drying hardly paler than petiole; intercostal venation
subscalariform; wood soft. On river banks and in mixed dipterocarp forest……………
……………………………………………………………………..31. V. umbonata
Leaf blade c. 9 × 4 cm, midrib drying paler than the dark petiole; intercostal venation
reticulate; wood hard. In kerangas and mixed peat samp forest………………………….
…………………………………………………………17. V. mangachapoi (in part)
9. Lateral veins hardly raised below, or no more than above…………………………10

Lateral veins distinctly raised below, more than above………………………….…19
10. Twig at first compressed…………………………………………….…7. V. compressa

Twig terete……………………………………………………………………………11
11. Tomentum vinoussericeous……………………………………………………….…12

Tomentum greyish, yellowish, pink, rufous, or ochreous, not sericeous………….…14
12. Leaf base cuneate………………………………………..…10. V. dulitensis (in part)

Leaf base obtuse…………………………………………………………………….…13
13. Leaf blade larger, 9–23 × 3.5–7.5 cm long, narrowly ellipticlanceolate………………...

………………………………………………………………25. V. pedicellata (in part)
Leaf blade smaller, 5.5–10 × 3–5 cm long, broadly ellipticovate…….…28. V. rotata
14. Leaf blade elliptic, coriaceous, drying dull, pale, pinkish brown; petiole at least 1.5 cm
long, slender. In mixed dipterocarp forest on yellow sandy soils.……...3. V. borneensis
Leaf blade drying dark, tawny, rufous, yellowish or greyishbrown; not coriaceous if
elliptic; petiole at most 1 cm long. In river banks, peat swamp or kerangas forest……15
355
15. Leaf blade narrowly obovate, apex obtuse to retuse, thickly coriaceous, concave,
margin revolute; petiole stout; tomentum ochreous. In kerangas into the mixed
dipterocarp forest ecotone………………………………….…9. V. coriacea (in part)
Leaf blade ovate, elliptic or lanceolate, apex acute, acuminate to caudate, coriaceous to
thinly coriaceous, flat, margin not revolute; or, if obovate, not coriaceous and with
slender petiole; tomentum otherwise. In different forest habitats.………………..…16
16. Tomentum pale pinkish to rufousbrown…………………………………………....17

Tomentum greyish brown or creamyellow……………………………………..…18
17. Lateral veins c. 8 pairs; midrib flat above; leaf blade narrowly ovate to lanceolate. In
kerangas forest…………………………………………………………23. V. parvifolia
Lateral veins 7–12 pairs; midrib sunken above; leaf blade elliptic to narrowly obovate.
In forest on river banks…………………………32. V. venulosa (subsp. venulosa)
18. Tomentum creamyellow, caducous; leaf blade at least 2.7 cm wide, broadly elliptic,
base cuneate; lateral veins at most 9 pairs; petiole at least 0.5 cm long. In kerangas and
mixed peat swamp forests…………………………….…17. V. mangachapoi (in part)
Tomentum pale greyish brown, not at first caducous; leaf blade at most 2.5 cm wide,
narrowly ellipticlanceolate, base obtuse; lateral veins at least 10 pairs; petiole to 0.4
cm long. In forest on alluvium along river banks……….....29. V. rynchocarpa (in part)
19. Petiole, midrib and veins below vinouscinereous; fresh leaves distinctly bluish green
below……………………………………………………………………………..…20
Petiole, midrib and veins below rufous, pinkish brown or pale greyish or yellowish
brown tomentose; fresh leaves not bluish green below…………………………......23
20. Twigs at first compressed……………………………………….…21. V. oblongifolia

Twigs terete………………………………………………………………………….…21
21. Petiole 2–3 mm diameter………………………………..…25. V. pedicellata (in part)

Petiole less than 1 mm diameter…………………………………………………..…22
22. Lateral veins 12–20 pairs, slender but more or less prominently raised below. In mixed
dipterocarp forest on clay soils, at altitudes to 600 m………………..…33. V. vinosa
Lateral veins 10–12 pairs, slender, hardly raised below. Mostly in upper dipterocarp
forests on ridges, at altitudes to 1350 m……………………10. V. dulitensis (in part)
23. Tomentum rufous or pinkishbrown…………………………………………….…24

Tomentum cream, pale greyish to yellowish brown, or yellowish………………….…30
24. Lateral veins 7–13 pairs, without or with a few short intermediate veins; leaf blade
broadly ovate, elliptic, lanceolate, oblanceolate or obovate; petiole long, slender…….25
Lateral veins 12–28 pairs, with many intermediate veins; leaf blade narrowly oblong to
obovate; petiole short and stout, or long and slender……………………………..…27
356
25. Lateral veins sharply prominent below; tomentum rufous, flocculent, caducous………..
………………………………………………………………..16. V. maingayi (in part)
Lateral veins terete below; tomentum even, pinkish brown to pale yellowish brown,
persistent………………………………………………………………………………26
26. Petiole at least 2 cm long; leaf blade drying rich pinkish brown, glistening……………..
…………………………………………………………………………2. V. badiifolia
Petiole at most 2 cm long; leaf blade drying pale greyish brown……..26. V. perakensis
27. Petiole at least 3 mm diameter, stout and short……….…30. V. sarawakensis (in part)
Petiole at most 2.5 mm diameter, slender………………………………………..…28
28. Tomentum short, even, persistent; lateral veins narrowly grooved above..13. V. globosa
Tomentum in part flocculent, caducous; lateral veins not grooved above……………29
29. Petiole to 1.2 cm long, hardly geniculate; leaf blade narrowly ellipticobovate;
intermediate and intercostal veins prominent below…………………15. V. havilandii
Petiole to 2 cm long, geniculate; leaf blade narrowly oblong; intermediate and
intercostal veins hardly raised below………………………………..…20. V. nitens
30. Lateral veins at least 15 pairs; petiole at least 3 mm diameter…………………..…31

Lateral veins less than 15 pairs; petiole less than 3 mm diameter……………….…32
31. Petiole more than 2 cm long………………………………………….…27. V. rassak

Petiole shorter than 2 cm…………………………………30. V. sarawakensis (in part)
32. Leaf blade below densely evenly ochreous sericeous. In kerangas forest...5. V. brunigii
Leaf blade below glabrous or sparsely tomentose on venation only. Not in kerangas
forest……………………………………………………………………………..…33
33. Leaf blade ellipticobovate………………………………………………………..…34

Leaf blade ovate, narrowly elliptic, lanceolate, oblong, or obovateoblong...………..38
34. Lateral veins at most 8 pairs; petiole at least 1.5 cm long……..…24. V. patentinervia
Lateral veins at least 10 pairs or, if less, then petiole at most 1.1 cm long.............…35
35. Leaf blade chartaceous………………………………………………………….……36

Leaf blade coriaceous to thickly coriaceous………………………………………..…37
36. Lateral veins at most 10 pairs……………………………………….…4. V. brevipes

Lateral veins at least 11 pairs……………………………………………11. V. endertii
37. Lateral veins 12–24 pairs; tomentum scabrid…………….…14. V. granulata (in part)
Lateral veins 10–13 pairs; tomentum even……………………………..…8. V. congesta
38. Intercostal venation prominent below; lateral veins and midrib below persistently
shortly scabridpubescent; lateral veins arching and anastomosing to form an
intramarginal vein……………………………………………………19. V. micrantha
Intercostal venation not prominent below; lateral veins and midrib glabrescent; lateral
veins otherwise..……………………………………………………………………39
357
39. Midrib raised above……………………………………………….…18. V. maritima

Midrib flat or furrowed above………………………………………………….…40
40. Leaf blade larger, 10–20 × 2.7–7 cm, narrowly obovate; twigs angular, peeling in small
flakes; lateral veins stoutly prominent below....................…14. V. granulata (in part)
Leaf blade smaller, 8–16 × 2.7–5.5 cm, narrowly elliptic to ovate; twigs terete; lateral
veins slender though raised below………………………………………22. V. odorata
1. Vatica albiramis Slooten

(Latin, albus = white; ramis = twigs; with white twigs)
Bull. Jard. Bot. Buitenz. 3, 9 (1927) 101; Masamune op. cit. 498; Meijer & Wood op. cit. 305; Ashton
op. cit. (1964) 66, op. cit. (1968) 30, op. cit. (1982) 355; Burgess op. cit. 227; Anderson op. cit. (1980)
131; Coode et al. (eds.) op. cit. 84. Type: Elmer 21640, Borneo, Sabah, Tawau district (holotype BO;
isotypes KEP (fragment), L).
Subcanopy tree, to 25 m tall, to 35 cm diameter. Parts glabrous but for caducous puberulent
flower calyx and ovary. Twigs c. 1 mm diameter apically, slender. Stipules hastate, to 16 ×
3.5 cm, caducous. Leaves thinly coriaceous, drying dull greyish green; blade elliptic to
lanceolate, 8–20 × 3–7 cm, base narrowly cuneate, apex with slender acumen to 1.5 cm
long; lateral veins 4–6 pairs, rather broad, slightly raised on both surfaces though most
prominently below, arched and continuing along margin at first; intercostal venation
subscalariform; petiole glabrous, drying black, 1–1.5 cm long. Inflorescences to 28 cm
long, terminal or axillary, lax. Flowers: corolla lemon yellow; other parts as the genus
(typical). Fruits calyx lobes free, subequal, ovate, to 1.2 × 0.4 cm, obtuse, coriaceous,
glabrous, becoming reflexed. Nuts globose, to 1.2 cm diameter, smooth, fulvous puberulent.
Vernacular names. Sabah—resak putih (preferred name). Sarawak—resak ranting putih

(Malay).
Distribution. Endemic in Borneo. In Sabah, widespread and known from most districts
recorded from Bintulu, Kapit, Lawas, Marudi, and Miri districts (e.g., S 18091, S 41411, S
44632, S 57292, and S 61052). Also occurring in Brunei (e.g., BRUN 3048, Dransfield JD
7060 and FMS 39647).
Ecology. In mixed and upper dipterocarp forest, on clayrich soils especially on ridges, at
altitudes to 1400 m. Common and probably not vulnerable.
2. Vatica badiifolia P.S.Ashton

(Latin, badius = chestnutbrown, folius = leaf; the dry leaf colour)
Coode et al. (eds.) op. cit. 84. Type: Ilias & Johnson S 15857, Borneo, Sarawak, Similajau district,
Labang FR (holotype K; isotype L). Synonym: Vatica bantamensis auct. non (Hassk.) Benth. &
Hook. ex Miq.: Ashton op. cit. (1964) 67.
358
Canopy tree, to 40 m tall, to 50 cm diameter. Young twig, inflorescence, leaf bud, stipule,
petiole, parts of flower exposed in bud, and ovary densely evenly pinkish brown puberulent.
Twigs to 3 mm diameter apically, terete or ribbed, becoming smooth. Stipules hastate, to 5
× 1.5 mm. Leaves thinly coriaceous, caducously tomentose below, drying rich pinkish
brown and frequently glistening; blade elliptic, 7.7–15 × 3–6.5 cm, base cuneate, margin
not revolute, apex deltoidacuminate, acumen to 0.5 cm long; lateral veins 9–12 pairs,
arched, stout, terete and raised on both surfaces though most prominently below, without
intermediate veins; intercostal venation prominent below; petiole 2–3.5 cm long, slender,
not drying black, distinctly thickened in the distal half and geniculate. Inflorescences
terminal or axillary, singly or doubly branched, to 8 cm long. Flowers: buds to 1.3 cm long;
calyx densely shortly cream pubescent; otherwise typical. Fruits: pedicels to 5 mm long,
slender; calyx lobes free to the platelike base around the nut, unequal, chartaceous, 2
longer lobes oblongspatulate, to 8 × 2.5 cm, constricted to 4 mm wide at base, not
becoming reflexed, 3 shorter ones hastate, to 3 × 0.8 cm, similarly constricted. Nuts
globose, to 0.8 × 0.8 cm, persistently fulvous puberulent; apical style remnant often
persisting.
Vernacular name. Sarawak—resak bantam (Malay).
Distribution. Endemic in Borneo. In Sarawak recorded from Bintulu, Kuching, Miri, and
Brunei (e.g., FMS 39650 and S 2120 ) and W Kalimantan (e.g., bb. 10596, bb. 18312).
Ecology. Locally frequent, in mixed dipterocarp forest on deep leached soils on coastal
hills. Common in Lambir NP; elsewhere endangered by forest conversion.
3. Vatica borneensis Burck

(of Borneo)
Ann. Jard. Bot. Buitenz. 6 (1887) 230; Merrill op. cit. (1921) 208; Slooten op. cit. (1927) 87;
(1982) 363; Anderson op. cit. (1980) 131; PROSEA op. cit. 465; Coode et al. (eds.) op. cit. 84. Type:
Beccari PB 2623, Borneo, Sarawak, Matang (BO). Synonyms: Vatica urbanii F.Heim, Bull. Mens.
Soc. Linn. Paris 2 (1891) 956; Sunaptea urbanii (F.Heim) F.Heim, Rech. Dipt. (1892) 115; S.
borneensis (Burck) F.Heim op. cit. (1892) 116.
Canopy tree, to 35 m tall, to 70 cm diameter. Young twig, inflorescence, parts of flower

exposed in bud, ovary, midrib below, and petiole densely evenly pinkish brown puberulent,
more or less persistent. Twigs c. 1.5 mm diameter apically, slender, terete or ribbed,
smooth or rugulose. Stipules unknown. Leaves coriaceous, drying dull pale pinkish brown;
blade elliptic, 6–10 × 2.5–5 cm, base cuneate, margin not revolute, apex acuminate, acumen
to 0.6 cm long; midrib prominent below, almost flat above; lateral veins 7–9 pairs, arched,
distinct but hardly and subequally raised on both surfaces as also the intercostal veins;
petiole 1.5–2.5 cm long, slender, hardly or not swollen distally, not drying black.
Inflorescences axillary, to 5 cm long. Flowers: buds to 1 cm long, densely pinkish brown
pubescent; otherwise typical. Fruits: pedicels c. 5 mm long, slender; calyx glabrescent,
lobes free, unequal, chartaceous, 2 longer lobes oblongspatulate, to 5.5 × 1.5 cm, to 0.4
cm wide at base, not becoming reflexed, 3 shorter ones unequal, narrowly ovate, tapering, to
1.8 × 0.7 cm. Nuts globose, to 1 cm diameter, smooth, glabrescent, with persisting style
remnant.
359
Vernacular name. Sarawak—resak kemudi (preferred name).
Distribution. Endemic in Borneo; recorded in Sarawak from Bau, Bintulu, Kuching,

Lundu, and Simunjan districts (e.g., S 6360, S 17877, S 19576, S 27136, and S 66788). Also
occurring in Brunei (e.g., S 2134 and SAN 17545).
Ecology. Locally common, more often scattered, in mixed dipterocarp forest on poor yellow
sandy soils; also in upper dipterocarp forest on coastal sandstone hills and, occasionally,
inland ridges, at altitudes to 900 m. Locally common in Bako NP; elsewhere endangered.
4. Vatica brevipes P.S.Ashton

(Latin, brevis=short, pes= foot; the short petiole)
Gard. Bull. Sing. 31 (1978) 24, op. cit. (1982) 366. Type: Othman S 29633, Borneo, Sarawak, Kapit
district, Ulu Baleh (holotype K; isotype KEP).
Main canopy or subcanopt tree, to 25 m tall, 30 cm diameter. Buds, petioles and

inflorescences densely persistently pale brown scabridpuberulent; parts of corolla exposed
in bud and ovary evenly so; sepals caducously evenly so; parts otherwise glabrous. Twigs
rugose and ribbed, becoming flaky, terete, dark brown, c. 2 mm diameter apically. Leaves
chartaceous, glabrous, more or less concave between the lateral veins; blade elliptic to
obovate, (4–)5–13 × (1.5–)2–5(–5.5) cm, base narrowly cuneate, margin not revolute, apex
with short, slender acumen to 0.6 cm long; midrib slender, sharply raised on both surfaces;
lateral veins 7–10 pairs, ascending, straight at first, arching and forming intermittent
intramarginal connections, slender but prominent below, elevated above, with a few short
intermediates; intercostal venation distantly reticulate, evident on both surfaces though more
so below; petiole somewhat stout, 0.5–1.1 cm long, not drying black. Inflorescences
axillary or terminal, to 1.6 cm long; rachis very slender, hardly branched. Flower: buds
ellipsoid, to 3 × 2 mm; otherwise typical. Fruits: pedicel slender, to 6 mm long; calyx lobes
unequal free, 2 longer lobes spatulate, subacute or obtuse, to 5 × 1.4 cm, not becoming
reflexed, 3 shorter ones lanceolate, acute, to 1.2 × 0.3 cm. Nuts subglobose, to 0.8 cm
diameter, apiculate.
Distribution. Endemic in Borneo. Known in Sarawak from Kapit district (e.g., S 23943, S
29576 and S 41495). Also occurring in W Kalimantan (e.g., Suzuki K 9766).
Ecology. Apparently uncommon, in mixed and upper dipterocarp forest on clay soils over
sedimentary rock, at altitudes between 250–1300 m. Vulnerable.
5. Vatica brunigii P.S.Ashton

(E.F.W.O. Brunig, forester in Sarawak and Brunei, 1954–1964)
Coode et al. (eds.) op. cit. 84. Type: Ilias S 17045, Borneo, Sarawak, Samarahan district, Sabal FR
360
Canopy tree, to 30 m tall, to 70 cm diameter. Living exposed parts and ovary, leaf above
excepted, persistently shortly yellowish buff scabridpubescent; leaf above fugaceous
flocculent pubescent. Twigs c. 2 mm diameter apically, terete. Stipules lorate, to 5 × 2 mm,
obtuse. Leaves thinly coriaceous, persistently ochreous pubescent below; blade ovate
elliptic to obovate, 6–12 × 2.5–6.5 cm, base cuneate to obtuse, margin not revolute, apex
acuminate with tapering acumen to 1 cm long; lateral veins 9–12 pairs, unraised above,
slender but prominent below as also the midrib and subscalariform intercostal venation;
petiole 0.8–1.5 cm long, to 3 mm diameter, not drying black. Inflorescences terminal or
axillary towards twig endings; rachis singly or doubly branched, to 12 cm long. Flowers
distichous; buds ellipsoid, to 8 × 2 mm; sepals densely ochreousgrey pubescent; petals
narrowly oblong, sparsely pubescent outside, glabrescent within. Fruits: pedicels to 3 mm
long, slender; calyx lobes unequal, free, chartaceous, 2 longer lobes lorate to spatulate, to
6.5 × 1.5 cm, not becoming reflexed, 3 shorter ones ovate, acute, slightly recurved, to 1.5 ×
0.6 cm. Nuts ovoid, to 0.9 × 0.7 cm, subacute.
Distribution. Sumatra and Borneo. In Sarawak known from Bintulu, Kuching, Limbang,
Lundu, Marudi, Samarahan, and Simunjan districts (e.g., S 1484, S 6437, S 8256, S 17885,
and S 69059). Also occurring in Brunei (e.g., S 1011 and S 12352) and W Kalimantan (e.g.,
Church et al. 987).
Ecology. In kerangas forests, both on raised beaches and dry sandstone plateaux and
cuestas, at altitudes to 700 m. Occurring in Bako and Mulu NPs; elsewhere critically
endangered by forest conversion.
6. Vatica chartacea P.S.Ashton

(Latin, chartaceus = papery; the leaf blade)
Gard. Bull. Sing. 31 (1978) 18, op. cit. (1982) 351. Type: Leopold SAN 46204, Borneo, Sabah, Labuk
Sugut district, Sapi FR (holotype K; isotype SAN).
Mediumsized tree. Young parts buff puberulent, caducous except on parts exposed in bud
and ovary. Twigs c. 2 mm diameter apically, muchbranched. Stipules lanceolate, to 7 × 2
mm, caducous. Leaves thinly chartaceous, drying wrinkled and pale yellowish brown;
blade oblong to obovate, 11–25 × 3–10 cm, base broadly cuneate to obtuse, apex acuminate
with prominent acumen to 1 cm long; lateral veins 16–20 pairs, with short slender
intermediates, slender but prominent below, distinctly elevated above as also the midrib,
arched; intercostal venation subscalariform, sinuate, elevated on both surfaces; petiole 1–2.2
cm long, slender, drying black, glabrous. Inflorescences to 5 cm long, in axillary cluster of
3, stout. Flowers: buds spindleshaped, to 6 × 2 mm; sepals subequal, lanceolate; otherwise
typical. Fruits: pedicels to 6 mm long, slender; calyx revolute, lobes subequal, chartaceous,
free, lanceolate, cordate, subacute, 5–7veined, to 6 × 1.5 cm, ascending and conceiling the
nut. Nuts ellipsoid, to 1.3 × 1.1 cm.
Distribution. Endemic in Borneo. In Sabah recorded from Kinabatangan, Kudat, Lahad

Datu, Pensiangan, and Sandakan districts (e.g., SAN 17654, SAN 18397, SAN 25451, SAN
57046, and SAN 97326) and in Sarawak from Bintulu and Miri districts (e.g., S 15808, S
40177 and S 60537). Also in W Kalimantan (e.g., bb. 35244 and bb. 35257).
361
Fig. 33. Vatica congesta. A, fruiting leafy twig; B, detail of indumentum on lower leaf
surface; C, flower bud; D, longitudinal section of flower bud; E, adaxial view of outer sepal;
F, adaxial view of inner sepal; G, abaxial view of petal; H, adaxial view of petal; I, adaxial
view of stamens; J, abaxial view of stamens; K, gynoecium; L, fruit with exposed nut. (A–B
from S 29474, C–K from S 15021, L from S 9347A.)
362
Ecology. Very local, in forest on floodplains and alluvium banks of sluggish rivers.
Critically endangered owing to land conversion.
7. Vatica compressa P.S.Ashton

(Latin, compressus = compressed; the young twig)
Type: Daud & Tachun SFN 35618, Borneo, Sarawak, Setapok Forest (holotype KEP; isotypes KEP,
L).
Canopy tree, to 30 m tall, to 35 cm diameter. Exposed young parts and ovary pale pinkish
brown pubescent, caducous except on leaf bud and stipules. Twigs at first c. 3 × 1 mm
apically, compressed, smooth; stipule scars pale, horizontal, prominent. Stipules not seen.
Leaves coriaceous; blade ovateelliptic, 11–19 × 5–8.5 cm, base obtuse, apex with acumen
to 1 cm long; midrib prominent below, flat or elevated above; lateral veins 10–15 pairs,
with short indistinct intermediates, slender and hardly elevated on either surface though
more so below, arched; intercostal venation subscalariform; petiole 1.5–2.3 cm long,
slender, not drying black. Inflorescences to 7 cm long; rachis somewhat compressed,
irregularly singly or doubly branched. Flowers: buds lanceolate, to 9 × 4 mm; otherwise
typical. Fruits: pedicels to 8 mm long, slender; calyx lobes free, unequal, chartaceous, 2
longer lobes lorate, obtuse, to 6.5 × 1 cm, tapering to 3 mm wide at base, 3 shorter ones
deltoid, acute, revolute, to 2 × 0.6 cm. Nuts broadly ovoid, to 0.5 cm diameter, with to 3
mm filiform style remnant.
Distribution. Endemic in Borneo. Confined to Sarawak, and recorded from Kuching,

Lundu, and Sri Aman districts (e.g., S 5984, S 6604, S 13275, and S 49944).
Ecology. Rare and local, in kerangas forest, apparently at least sometimes with impeded
drainage. Critically endangered, possibly extinct.
8. Vatica congesta P.S.Ashton Fig. 33.

(Latin, congestus = heaped together; the inflorescences)
Type: Yakup S 9347A, Borneo, Sarawak, Kuching district, Semengoh FR (holotype K; isotypes KEP,
L, SAR).
Suncanopy or canopy tree, to 35 m tall, to 35 cm diameter. Exposed fleshy parts mostly

densely shortly pale ochreous brown scabridpubescent; leaf blade below and midrib
towards base above sparsely so; leaf blade above glabrescent; parts otherwise persistently
so. Twigs c. 3 mm diameter apically, stout, at first terete, rugose, becoming striated and
thinly flaky. Stipules deltoid, to 8 × 3 mm, acute. Leaves thickly coriaceous, glabrous,
drying pale tawny brown; blade oblong, elliptic to obovate, 8–22 × 3.5–8 cm, base obtuse,
margin revolute, apex obtuse to retuse; lateral veins 10–13 pairs, prominent and stout
below, elevated above, as also the midrib; intercostal venation reticulate, similarly raised;
petiole 1.2–2.5 cm long, to 3 mm diameter, stout, rugulose, not drying black. Inflorescences
axillary to ramiflorous, fascicled; rachis angular, to 3 cm long, short and congested, singly
branched. Flowers distichous; buds to 8 × 3 mm; sepals deltoidacute; petals lorateobtuse;
otherwise typical. Fruits: pedicels to 4 mm long, scabridpubescent, fruit otherwise
363
glabrous; calyx lobes free, unequal, chartaceous, 2 longer lobes lorate, obtuse, to 12 × 2.2
cm, not becoming reflexed, 2 shorter ones narrowly ovate, to 4.5 × 1.5 cm, tapering, acute,
recurved. Nuts ellipsoid, to 1.2 × 0.8 cm, with to 6 mm style remnant, glabrous.
Distribution. Endemic in Borneo. Known in Sarawak from Belaga and Kuching districts
(e.g., S 14936, S 15713, S 29476, S 32501, and S 56979). Also occurring in W Kalimantan.
Ecology. Very local but frequent where it occurs, in mixed dipterocarp forest on poor sandy
clay soil. Endangered.
9. Vatica coriacea P.S.Ashton

(Latin, coriaceus = leathery; the leaf texture)
op. cit. (1980) 131; PROSEA op. cit. 465; Coode et al. (eds.) op. cit. 84. Type: Ariffin S 372, Borneo,
Sarawak, Lundu district, Bt. Sebandar (holotype K; isotype KEP).
Canopy tree, to 20 m tall, to 50 cm diameter. Exposed young parts densely shortly ochreous
puberulent, glabrescent except on buds. Twigs to 5 mm diameter apically, terete, stout,
ribbed. Stipules hastate, to 13 × 5 mm, subacute. Leaves thickly coriaceous, drying tawny
brown to dark brown; blade narrowly obovate, 6.5–15 × 2.2–6 cm, concave, base narrow,
obtuse or cuneate, margin revolute, apex obtuse to retuse; midrib stoutly prominent below,
slightly raised above; lateral veins 10–11 pairs, with short slender intermediates, slightly
and more or less equally elevated on both surfaces, arched; intercostal venation reticulate;
petiole 1–1.5 cm long, stout, drying black or not. Inflorescences axillary or terminal; rachis
to 20 cm long, singly branched, often bearing small modified leaves at branch bases.
Flowers to 18 mm long; calyx densely shortly pale greyish brown pubescent; otherwise
typical. Fruits: pedicels to 7 mm long, slender; calyx lobes free to base, unequal,
chartaceous, 2 longer lobes oblong, obtuse, to 7 × 2.3 cm, tapering to 3.5 mm broad at
base, not becoming reflexed, 3 shorter ones hastate, acute, to 0.6 × 0.2 cm. Nuts globose, to
0.8 cm diameter, style remnant to 2 mm long.
Vernacular name. Sarawak—resak daun tebal (preferred name).
Distribution. Endemic in Borneo. In Sabah, known from a single collection from Tawai
Plateau, Kinabatangan district (SAN 39325) and in Sarawak known from Kuching and
Lundu districts (e.g., S 6344, S 10296, S 15607, and S 15798). Also occurring in Brunei
(e.g., BRUN 3155 and BRUN 5652) and C Kalimantan (e.g., Suzuki K 11483).
Ecology. Locally common, in kerangas forest on giant humic podsols on Pleistocene raised
beaches, and on organic soils over limestones. Well represented in Bako NP and recorded
from Mulu NP; endangered outside parks system.
10. Vatica dulitensis Symington

(of G. Dulit, Sarawak)
Gard. Bull. S. S. 8 (1934) 35; Masamune op. cit. 498; Browne op. cit. 100; Ashton op. cit. (1964) 69,
364
cit. (1980) 131; PROSEA op. cit. 466; Coode et al. (eds.) op. cit. 84. Type: Richards 1343, Borneo,
Sarawak, G. Dulit (holotype K).
Subcanopy to small canopy tree, to 30 m tall, to 50 cm diameter. Exposed fleshy parts

including petiole and leaf venation below dark vinoussericeous, persistent except on
venation. Twigs terete, to 1 mm diameter apically, very slender, muchbranched. Stipules
narrowly hastate, to 3 × 1 mm, acute, deciduous. Leaves thinly coriaceous, drying pale
greyish brown; blade narrowly obovate to lanceolate, 4–11 × 0.8–3.2 cm, base cuneate,
apex caudate, acumen to 1.5 cm long; lateral veins 10–12 pairs, with short slender
intermediates, slender, hardly raised below, arched; intercostal venation reticulate, evident
but hardly raised; petiole 0.6–1 cm long, to 1 mm diameter, slender, not drying black.
Inflorescences terminal or axillary; rachis singly or doubly branched, to 2.5 cm long.
Flowers: buds to 6 mm long; sepals densely vinoussericeous; petals cream; otherwise
typical. Fruits: pedicels to 3 mm long, slender; calyx lobes chartaceous, free, equal,
oblongobtuse, to 1.4 × 0.5 cm, vinous caducous cinereous within only, becoming rotate to
reflexed. Nuts globose, to 0.8 cm diameter, smooth, vinous cinereous, with short style
remnant.
Vernacular names. Sabah—resak bukit (preferred name). Sarawak—resak tiong (preferred

name).
Distribution. Endemic in Borneo. In Sabah known from Beaufort, Kota Belud, Labuk
Sugut, Lahad Datu, Ranau, and Sipitang districts (e.g., SAN 16267, SAN 26712, SAN 76253,
SAN 78171, and SAN 116860) and in Sarawak from Belaga, Kapit, Kuching, Lawas,
Limbang, Lundu, Marudi, Miri, and Tatau districts (e.g., S 10184, S 14770, S 17738, S
36093, and S 43570). Also occurring in Brunei (e.g., BRUN 3042 and BRUN 3156) and NE
and E Kalimantan (e.g., Kostermans 9095 and Kostermans 9248).
Ecology. Local but often abundant on high ridges on shallow organic soil, in upper
dipterocarp forest, at 600–1350 m altitude, and occasionally in lowland mixed dipterocarp
forest on leached clay soils. Occurring in G. Gading and Mulu NPs; elsewhere vulnerable.
11. Vatica endertii Slooten

(F.H. Endert, 1891–1953, Dutch forester in Indonesia)
Bull. Jard. Bot. Buitenz. 3, 18 (1942) 248; Ashton op. cit. (1968) 32, op. cit. (1982) 366; Anderson op.
cit. (1980) 131. Type: Endert 2775, Borneo, Kalimantan, W Kutei, Long Hoet (holotype BO; isotype
L).
Canopy tree, to 30 m tall, to 60 cm diameter; bole sinuous. Exposed young parts shortly
pale brown caducous scabridpuberulent, more or less persistent on inflorescence and
sometimes on veins below; densely persistently so on ovary; parts otherwise glabrous.
Twigs terete, c. 1 mm diameter apically, muchbranched, at first striated. Stipules not seen.
Leaves chartaceous, drying pale buffbrown; blade narrowly obovate, oblong or rarely
lanceolate, 4.8–14 × 1.7–5.5 cm, base cuneate, margin not revolute, apex acuminate with
slender acumen to 1 cm long; lateral veins 11–14 pairs, without intermediates, prominent
below, unraised above as also the midrib; intercostal venation subscalariform; petiole 0.8–2
cm long, not drying black. Inflorescences terminal or axillary, not fascicled; rachis singly
branched, striated, to 3 cm long. Flowers typical. Fruits: pedicels to 8 mm long, slender;
calyx lobes unequal, free, chartaceous, 2 longer lobes oblongspatulate, obtuse, to 8 × 2
365
cm, not becoming reflexed, c. 3 mm wide at base, 3 shorter ones lanceolate, to 2 × 0.6 cm,
acute, hardly recurved. Nuts subglobose, to 0.6 × 0.5 cm.
Distribution. Endemic in Borneo. Known in Sarawak from Kapit and Miri district (e.g., S
14383, S 19036 and S 19037). Also occurring in E Kalimantan (e.g., the type).
Ecology. Scattered, local, in upper dipterocarp forest on acid volcanic rock (dacite) in
Sarawak, occasionally also in lowland mixed dipterocarp forest in Kalimantan. Vulnerable
owing to logging.
12. Vatica glabrata P.S.Ashton

(Latin, glabratus = smoothskinned; the plant parts)
FM 1, 9 (1982) 370; Coode et al. (eds.) op. cit. 84. Type: Tong S 36852, Borneo, Sarawak, Marudi
district, Dulit Range (holotype K).
Canopy tree, to 20 m tall, to 60 cm diameter. Buds and ovary densely persistently ochreous
puberulent, young parts fugaceously so; parts otherwise glabrous, more or less shiny.
Twigs terete, c. 2 mm diameter apically, smooth. Stipules elliptic, obtuse, to 12 × 8 mm,
not at first caducous, leaving a falcate scar. Leaves coriaceous, more or less concave,
shiny, drying yellowish brown; blade lanceolate, 6–15(–22) × (2–)2.5–7 cm, base obtuse,
margin not revolute, apex acuminate with prominent, attenuate acumen to 1.5 cm long;
midrib flat or raised above; lateral veins 9–11 pairs, with more or less short intermediates,
ascending, slender, raised on both surfaces but more so below; intercostal venation
reticulate, distinctly raised on both surfaces; petiole 1–4 cm long, slender, prominently
geniculate glabrescent, drying black. Inflorescences 1–several axillary or terminal; rachis
to 7 cm long. Flowers typical. Fruits: calyx lobes unequal, free, chartaceous, not becoming
reflexed.
Distribution. Endemic in Borneo. In Sarawak known from Kapit and Marudi districts (e.g.,
S 34842, S 34865 and S 36383). Also occurring in Brunei (e.g., BRUN 2526 and BRUN
2533).
Ecology. Local but frequent, at the upper limits of upper dipterocarp forest, at 1200–1500 m
altitude. One sterile and aberrant collection (S 27283) from the karst pinnacles on G. Subis,
Niah, at altitude c. 600 m. Vulnerable.
13. Vatica globosa P.S.Ashton

(Latin, globosus = spherical; the nut)
Type: Ashton S 18091, Borneo, Sarawak, Labang, Ulu Stirau (holotype K; isotype L).
Subcanopy tree, to 20 m tall, 20 cm diameter. Exposed fleshy parts and ovary, including
entire fruit but not leaf blade and venation, densely persistently rufousbrown sericeous.
Twigs c. 2 mm diameter apically, terete. Stipules unknown. Leaves thinly coriaceous,
usually somewhat concave between the lateral veins, drying greyish to pinkish brown; blade
obovate, 7–18 × 3–6.5 cm, base narrowly cuneate, apex caudate, acumen to 2 cm long;
midrib prominent on both surfaces; lateral veins 12–16 pairs, with many intermediate veins,
366
slender but prominent below narrowly grooved above; intercostal venation reticulate;
petiole 0.7–1.5 mm long, slender, to 2.5 mm diameter, with distinct adaxial furrow, rugulose
and not black on drying. Inflorescences terminal or 1–several axillary, congested; rachis to
3 cm long. Flowers: buds lanceolate, to 5 × 2 mm; calyx densely vinous pubescent outside,
glabrous within; corolla sparsely so; otherwise typical. Fruits: calyx lobes equal, free,
oblong, obtuse, to 0.4 × 0.3 cm, reflexed, not conceiling the nut. Nuts globose, to 2 cm
diameter, 3sutured.
Distribution. Endemic in Borneo. In Sarawak recorded from Bintulu, Kapit, Kuching,

Marudi, Miri, and Sri Aman districts (e.g., Haviland & Hose 3159, S 14485, S 23751, S
42461, and S 64531). Also occurring in W Kalimantan (e.g., bb. 13931).
Ecology. Locally frquent in mixed dipterocarp forest on yellow sandy clay soils, on low
hills including the Arip rhyolite. Endangered by forest conversion.
14. Vatica granulata Slooten

(Latin, granulatus = appearing like tiny seeds; the nut surface)
Bull. Jard. Bot. Buitenz. 3, 9 (1927) 112, Bull. Jard. Bot. Buitenz. 3, 17 (1941) 136; Masamune op. cit.
498; Ashton op. cit. (1964) 70, op. cit. (1968) 32, op. cit. (1978) 21, op. cit. (1982) 354; Anderson op.
cit. (1980) 132; PROSEA op. cit. 466; Coode et al. (eds.) op. cit. 84. Type: Hallier 3399, Borneo, W
Kalimantan, G. Amai Ambit (holotype BO; isotype L).
Canopy tree, to 30 m tall, to 70 cm diameter. Exposed fleshy parts including leaf venation
below but not blade more or less densely shortly yellowish brown scabridpubescent;
persistent except on venation. Twigs c. 3 mm diameter apically, stout, angular, papery
flaky. Stipules hastate, subacute, to 6 × 4 mm, caducous or oblonglanceolate and acute to
30 × 20 mm, not at first caducous. Leaves coriaceous; blade narrowly obovateoblong or
ellipticobovate, 10–20 × 2.7–7 cm, base narrowly obtuse, apex with tapering acumen to 0.6
cm long; midrib furrowed above; lateral veins 12–14(–24) pairs, prominent below, slightly
sunken above, wellspaced; intercostal venation indistinct below, petiole 1.2–2 cm long, to 3
mm diameter, drying not black. Inflorescences variable. Fruits subsessile; calyx lobes free,
subequal, chartaceous, deltoid, brittle, glabrous, reflexed, to 0.7 × 0.4 cm or elliptic,
revolute, to 2 × 1.2 cm. Nuts ovoid, to 3.5 × 4 cm, acute or subacute, coarsely granulate, 3
furrowed.
Vernacular name. Sarawak—resak ranting bersisik (preferred name).
Ecology. In upper dipterocarp forest, on high ridges, at altitude between 500–1700 m.
Notes. Two subspecies, viz. subsp. granulata and subsp. sabaensis, are recognised in Sabah
and Sarawak.
Key to subspecies
Stipules hastate, subacute, to 6 × 4 mm, soon caducous. Inflorescence to 3 cm long, axillary.

Fruit calyx lobes deltoid, not revolute, to 0.7 × 0.4 cm……………………………………….
367
subsp. granulata
Locally abundant in its suitable habitat. In Sabah recorded from Keningau and
Tambunan districts (e.g., SAN 115452 and Wong WKM 2646) and in Sarawak from
Bau, Belaga, Kapit, Kuching, Marudi, Serian, and Tatau districts (e.g., S 15217, S
17658, S 22221, S 36347, and S 60381). Also occurring in Brunei (e.g., BRUN 2641
and FMS 30578) and W Kalimantan (e.g., Argent et al. 93156 and the type). Occurring
in Mulu NP; probably not vulnerable.
Stipules oblonglanceolate, acute, to 30 × 20 mm, not at first caducous. Inflorescence to 15
cm long, lax, terminal or axillary. Fruit calyx lobes elliptic, revolute, to 2 × 1.2 cm…….…...
subsp. sabaensis P.S.Ashton
(of Sabah)
Gard. Bull. Sing. 31 (1978) 21, op. cit. (1982) 354; PROSEA op. cit. 466. Type: G.H.S. Wood &
Kilang SAN 16613, Borneo, Sabah, Sipitang, Bt. Batanga north slope (holotype K; isotypes KEP,
SAN). Synonym: V. scortechinii auct. non (King) Brandis: Meijer & Wood op. cit. 319.
Known in Sabah from Kota Belud, Labuk Sugut, Lahad Datu, Ranau, and Sipitang
districts (e.g., SAN 16708, SAN 17000 and SAN 130634) and in Sarawak from Limbang
and Marudi districts (e.g., Nooteboom & Chai 2233, S 37286 and S 44403). Also
occurring in Brunei (e.g., Prance 30578) and E Kalimantan (e.g., Kostermans 7465). In
upper dipterocarp forest, at altitudes to 1700 m. Occurring in Kinabalu NP; probably
not vulnerable.
15. Vatica havilandii Brandis Fig. 34.

(G.D. Haviland, 1857–1901, surgeon and naturalist in Sarawak)
J. Linn. Soc. Bot. 31 (1895) 133; Merrill op. cit. (1921) 409; Slooten op. cit. (1927) 95; Masamune op.
cit. 498; Symington, J. Malay. Br. Roy. As. Soc. 19 (1941) 155, Malay. For. Rec. 16 (1943) 220;
Browne op. cit. 100; Ashton op. cit. (1964) 71, op. cit. (1968) 33, op. cit. (1982) 351; Anderson op.
cit. (1980) 132; PROSEA op. cit. 467; Coode et al. (eds.) op. cit. 85. Type: Haviland 1848, Borneo,
Sarawak, near Kuching (holotype K).
Small to mediumsized tree, occasionally to 30 m tall, to 30 cm diameter. Exposed fleshy

parts but for leaf blade and ovary densely persistently deep fulvousbrown flocculent
tomentose; indumentum glabrescent on fruit calyx lobes, sparse and early caducous on leaf
venation below. Twigs to 2.5 mm diameter apically, terete or slightly compressed, smooth
or slightly flaky. Stipules linear, to 12 × 2.5 mm, caducous. Leaves thinly coriaceous, more
or less bullate between lateral veins, drying reddish brown; blade narrowly obovateoblong,
8–17 × 2.5–5 cm, base cuneate, apex with slender acumen to 1 cm long; midrib terete,
prominent below, flat or somewhat elevated above; lateral veins 15–20 pairs, with
prominent short intermediates, slender, arched towards margin, flat to slightly raised above,
prominent below; petiole 1–1.2 cm long, to 2.5 mm diameter, hardly geniculate, not drying
black. Inflorescences terminal or axillary; rachis terete, singly branched, to 8 cm long.
Flowers: buds to 5 mm long; calyx densely rustbrown powdery tomentose; otherwise
typical. Fruits pedicels to 5 mm long, hidden within calyx; calyx lobes subequal, free,
chartaceous, ovate, acute, cordate, revolute, to 2.5 × 1.5 cm. Nuts globose, to 1.2 cm
diameter, conceiled within calyx.
Vernacular name. Sabah and Sarawak—resak degong (preferred name).
Distribution. Peninsular Malaysia and Borneo. In Sabah recorded from Sandakan district
(e.g., SAN 36684, SAN 37880 and SAN 79480) and in Sarawak from Kuching, Mukah and
368
Fig. 34. Vatica havilandii. A, flowering leafy twig; B, tufted hair; C, open flower; D,
adaxial view of outer sepal; E, adaxial view of inner sepal; F, abaxial view of petal; G,
adaxial view of petal; H, adaxial view of stamens; I, abaxial view of stamens; J, gynoecium;
K, fruit; L, fruit with exposed nut; M, basal view of fruit; N, hairs on nut. (A–J from SAN
36684, K–N from S 23244.)
369
Fig. 35. Vatica mangachapoi subsp. mangachapoi. A, fruiting leafy twig; B, detail of
indumentum of midrib on the lower leaf surface; C, flower bud; D, longitudinal section of
open flower; E, abaxial view of outer sepal; F, abaxial view of inner sepal; G, abaxial side
of petal; H, adaxial side of petal; I, adaxial view of stamens; J, abaxial view of stamens; K,
gynoecium; L, fruit with exposed nut. (A–B and L from FMS 41100, C–K from SAN
15459.)
370
Simunjan districts (e.g., S 23227, S 23244 and S 42993). Also occurring in Brunei (e.g.,
BRUN 676 and FMS 30517) and Kalimantan.
Ecology. Rare and local, in mixed dipterocarp forest on sandy clay soil, apparently usually
near water, sometimes, close to limestone. Likely critically endangered with extinction.
16. Vatica maingayi Dyer

(C. Maingay, 1836–1869, Superintendent of Malacca jail and naturalist)
Fl. Brit. Ind. 1 (1874) 302; Brandis op. cit. 131; Slooten op. cit. (1927) 85, p.p.; Symington op. cit.
(1941) 151, op. cit. (1943) 223; Meijer & Wood op. cit. 309; Burgess op. cit. 227; Ashton op. cit.
(1968) 33, op. cit. (1982) 369; Anderson op. cit. (1980) 132; PROSEA op. cit. 466. Type: Maingay
209, Peninsular Malaysia, Malacca (holotype K). Synonyms: Synaptea maingayi (Dyer) Ridl., FMP 1
(1922) 240, p.p.; Vatica macroptera Slooten ex Thorenaar, Med. Proefst. Boschw. 16 (1926) 120,
Slooten op. cit. (1927) 83, nomen in syn. sub V. lowii; V. aperta Slooten op. cit. (1942) 250.
Mediumsized to large canopy tree, to 45 m tall, to 70 cm diameter. Bark eventually flaky.

Exposed young fleshy parts, leaf blade excepted, and ovary shortly rufous flocculent
tomentose; persistent on ovary and leaf bud; patchily caducous elsewhere. Twigs terete, c. 2
mm diameter apically, smooth; leaf buds prominent. Stipules lorate, to 10 × 3 mm, leaving
prominent horizontal scars. Leaves thinly coriaceous, drying reddish brown; blade elliptic,
5.5–12.5 × 1.2–5 cm, base cuneate, margin not revolute, apex acuminate, with tapering
acumen to 1 cm long; midrib prominent on both surfaces; lateral veins 9–11 pairs, with few
short intermediates, slender but sharply prominent below, arched; intercostal venation
slender, reticulate; petiole 1–2.5 cm long, slender, ribbed, prominently geniculate, drying
black or not. Inflorescences axillary; rachis ribbed, irregularly singly branched, to 4 cm
long. Flowers: buds spindleshaped, to 10 mm long; calyx rufous flocculent tomentose;
corolla bright red; otherwise typical. Fruits: pedicels to 3 mm long; calyx lobes unequal,
free, chartaceous, 2 longer lobes lorate to narrowly ovate, obtuse, somewhat revolute at
base, not becoming reflexed, to 8 × 2.5 cm, to c. 7 mm wide at base, 3 shorter ones narrowly
ovate, acute, to 2.5 × 1 cm, to 3 mm wide at base. Nuts globose, to 0.7 cm diameter, shortly
mucronate.
Vernacular names. Sabah—resak daun merah (preferred name). Sarawak—resak lidi

(preferred name).
Distribution. Peninsular Malaysia, Sumatra (Palembang) and Borneo. In Sabah recorded

from Sipitang and Tambunan districts (e.g., SAN 15105 and SAN 17036) and in Sarawak
from Bau, Belaga and Kuching districts (e.g., S 9425, S 15785 and S 24826).
Ecology. In lowland mixed dipterocarp forest on yellow sandy clay soils. Rare and
endangered.
17. Vatica mangachapoi Blanco Fig. 35.

(a Philippine vernacular name)
Fl. Filip. ed. 1 (1837) 401; Slooten op. cit. (1927) 94; Merrill, PEB (1929) 205; Masamune op. cit.
499; Browne op. cit. 101; Anderson, Gard. Bull. Sing. 20 (1963) 159, op. cit. (1980) 132; Ashton op.
cit. (1963) 253, op. cit. (1964) 71, op. cit. (1968) 33, op. cit. (1978) 22, op. cit. (1982) 364; Meijer &
Wood op. cit. 310; Burgess op. cit. 227; PROSEA op. cit. 468; Coode et al. (eds.) op. cit. 85. Neotype
371
(designated here): Merrill Sp. Blancoan. 866 (= US 904562), the Philippines, Luzon, Bataan Province
(K, US). Synonyms: Mocanera mangachapoi Blanco op. cit. (1837) 450; Vatica apteranthera Blanco,
Fl. Filip. ed. 2 (1845) 156; Dipterocarpus mangachapoi (Blanco) Blanco op. cit. (1845) 313; Shorea
mangachapoi (Blanco) Blume op. cit. 34; Anisoptera mangachapoi (Blanco) A.DC., Prodr. 16, 2
(1868) 616; V. bureavi F.Heim op. cit. (1891) 955; Sunaptea bureavi (F.Heim) F.Heim op. cit. (1892)
114; V. reticulata auct. non (Thwaites) A.DC.: King, J. As. Soc. Beng. 62, 2 (1893) 166; Cotylelobium
philippinense F.Heim ex Brandis op. cit. 134; V. obtusifolia Elmer, Leafl. Philip. Bot. 4 (1912) 1471;
Synaptea reticulata Ridl., op. cit. (1922) 243; V. patula Symington op. cit. (1941) 148.
Canopy tree, to 30 m tall, to 70 cm diameter; bole frequently crooked. Exposed fleshy parts,
leaf excepted, shortly densely creamyellow puberulent; caducous except on inflorescence.
Twigs to 1.5 mm diameter apically, terete. Stipules narrowly oblong, to 5 × 2 mm,
subacute. Leaves thinly to thickly coriaceous, somewhat shiny, drying yellowish brown or
pale greyish green; blade elliptic, 6–11 × 2.7–5 cm, base cuneate, margin not revolute,
apex obtuse or subacute, or acute with tapering acumen to 0.7 cm long; midrib terete below,
raised above; lateral veins 7–9 pairs, without intermediates, slightly raised on both
surfaces, slightly arched; intercostal venation densely reticulate; petiole 0.5–1 cm long,
drying black or not. Inflorescences terminal or axillary; rachis terete, singly or doubly
branched, to 14 cm long. Flowers: buds to 12 mm long; calyx shortly densely creambuff
pubescent; otherwise typical. Fruits: pedicels to 4 mm long, slender; calyx lobes unequal,
free, chartaceous, 2 longer lobes spatulate, obtuse, to 5.5 × 1.5 cm, to 3 mm wide at base,
not becoming reflexed, 3 shorter ones lanceolate, acute, to 1 × 0.4 cm. Nuts subglobose, to
0.4 × 0.6 cm, shortly mucronate.
Vernacular names. Sabah—resak bajau (preferred name). Sarawak—resak julong

(Malay), resak paya (preferred name).
Distribution. Peninsular Thailand, Peninsular Malaysia, Borneo, and the Philippines.
Notes. Two subspecies, viz. subsp. mangachapoi and subsp. obtusifolia, are recognised in
Sabah and Sarawak.
Key to subspecies
Leaves thinly coriaceous, apex with tapering acumen to 0.7 cm long. Inflorescence to 14 cm
long. In forest on nonultrabasic soils….……………………………………………………...
subsp. mangachapoi
In Sabah known from Beaufort, Kinabatangan, Lahad Datu, Sandakan, Semporna, and
Tawau districts (e.g., SAN 15463, SAN 15504, SAN 29380, SAN 38937, and SAN
90785) and in Sarawak from Kuching, Marudi, Mukah, Sibu, Simunjan, and Sri Aman
districts (e.g., S 3159, S 9429, S 12878, and S 77024). Also occurring in Brunei (e.g.,
KEP 80149, S 5813 and Wong WKM 975). Locally common, near the coast on dry
ridges in kerangas forest on deep podsols on raised beaches and, particularly in
Sarawak, in mixed peat swamp forest. Occurring in Bako NP; elsewhere vulnerable.
Leaves thickly coriaceous, apex obtuse or subacute. Inflorescence to 6 cm long. In forest on
ultrabasic soils…………………………………………………………………………………
subsp. obtusifolia (Elmer) P.S.Ashton
Gard. Bull. Sing. 31 (1978) 23, op. cit. (1982) 365. Basionym: Vatica obtusifolia Elmer op. cit.
(1912) 1471. Lectotype (Ashton, 1978): Elmer 12963, the Philippines, Palawan, Puerto Princesa,
Mt. Pulgar (hololectotype K; isolectotype NY).
372
Palawan and E Sabah, Bt. Masasan, Lahad Datu district (e.g., SAN 25435). On rocky
hills, not far from the coast. Rare and endangered.
18. Vatica maritima Slooten

(Latin, maritimus = relating to the sea; the habitat)
359; Meijer & Wood op. cit. 310; Burgess op. cit. 227; Anderson op. cit. (1980) 133; Coode et al.
(eds.) op. cit. 85. Type: Orolfo SAN 1828, Borneo, Sabah, Kudat, Lokapas (holotype BO).
Mediumsized canopy tree, to 25 m tall, to 35 cm diameter. Exposed fleshy parts, leaf blade
excepted, and ovary densely more or less persistently cream puberulent. Twigs ribbed or
somewhat compressed, to 8 mm diameter apically. Stipules not seen. Leaves coriaceaous,
drying dull yellowish brown; blade broadly or narrowly ovate, 8–16 × 3–8 cm, base broadly
cuneate to subcordate, apex with broad acumen to 1 cm long; lateral veins 7–10 pairs, with
short slender intermediates, raised on both surfaces though more so below as also midrib
and intercostal venation; petiole 2–2.7 cm long, to 3 mm diameter, slender, hardly
geniculate, not drying black. Inflorescences axillary or rarely terminal; rachis singly
branched, lax, angled, to 11 cm long. Flowers: buds to 14 mm long; calyx densely pale
brown tomentose; otherwise typical. Fruits: pedicels to 4 mm long; calyx lobes unequal,
chartaceous, united into a to 5 mm diameter shallow cup at base, 2 longer lobes lorate
spatulate, obtuse, to 11 × 1.8 cm, tapering to 4 mm wide at base, 3 shorter ones lanceolate,
acute, to 1.8 × 0.6 cm. Nuts globose, to 0.6 cm diameter.
Vernacular name. Sabah and Sarawak—resak laut (preferred name).
Distribution. Borneo and the Philippines (Palawan Is.). In Sabah known from Kudat, Kota
Kinabalu, Lahad Datu, and Sandakan districts (e.g., SAN 5495, SAN 28441, SAN 35663,
SAN 36136, and SAN A 3183). Also occurring in Brunei (e.g., FMS 37066).
Ecology. Rare, in forest on rocky or yellow sandy soils on headlands near the sea. Critically
endangered.
19. Vatica micrantha Slooten

(Greek, micros = small, anthos = flower; the supposedly small flowers)
366; Meijer & Wood op. cit. 312; Burgess op. cit. 227; Anderson op. cit. (1980) 132; PROSEA op. cit.
469; Coode et al. (eds.) op. cit. 85. Type: bb. 29706, Borneo, C Kalimantan, Pepas, Muara Teweh
(holotype BO; isotype L).
Canopy tree, to 35 m tall, to 60 cm diameter. Exposed fleshy parts, including venation below
but not blade, and ovary more or less densely persistently pale brown scabridpubescent;
glabrescent on fruit calyx. Twigs c. 2 mm diameter apically, terete, smooth or slightly
striated. Stipules hastate, acute, to 7 × 3 mm. Leaves thinly coriaceous, drying greyish
green; blade often bullate between lateral veins, ellipticoblong to lanceolate, 4.5–16 × 1.5–
6 cm, base obtuse, margin not revolute, apex acuminate with slender acumen to 1.5 cm
long; lateral veins 8–11 pairs, with short slender intermediates, slender, arching and
anastomosing to form intramarginal vein, prominent below, flat above as also the
373
intercostal venation and midrib; petiole 0.5–1 cm long, to 3 mm diameter, not drying black.
Inflorescences terminal or axillary, not fascicled; rachis singly branched, terete, to 7.5 cm
long. Flowers: buds to 13 mm long; calyx shortly pubescent; corolla cream, suffused with
violet towards the base outside; otherwise typical. Fruits: pedicels to 2.5 mm long; calyx
lobes unequal, free, chartaceous, 2 longer lobes oblongspatulate, subacute, to 5.8 × 1.5
cm, tapering to 3 mm wide, revolute base, 3 shorter ones ovate, caudateacuminate, to 2.5 ×
0.6 cm, similar at base. Nuts ellipsoid, obtuse, to 1.4 × 0.8 cm; style remnant often absent.
Vernacular names. Sabah—resak bulu (preferred name). Sarawak—resak hijau (Malay).
Distribution. Endemic in Borneo; widespread. In Sabah recorded from Beaufort, Keningau,

Kota Kinabalu, Kudat, Labuk Sugut, Lahad Datu, Sandakan, Semporna, Sipitang, Tawau,
131865) and in Sarawak from Bau, Belaga, Bintulu, Kapit, Kuching, Lawas, Limbang,
Lundu, Marudi, Miri, Sri Aman, and Tatau districts (e.g., S 1754, S 15134, S 25016, S
32204, and S 60127). Also occurring in Brunei (e.g., BRUN 613 and Dransfield JD 6897)
and Kalimantan (e.g., bb. 29706 and Jarvie & Ruskandi 5894).
Ecology. The commonest Vatica in Sabah and Sarawak. Common in mixed dipterocarp
forest, on a wide range of sandy and clay soils including on basic volcanics and on the base
of limestone hills. Common in Lambir NP; not vulnerable.
20. Vatica nitens King Plate 5E.

(Latin, nitens = shining; the leaf surface)
J. As. Soc. Beng. 2 (1893) 104; Slooten op. cit. (1927) 78; Ashton op. cit. (1964) 223, op. cit. (1968)
34, op. cit. (1982) 367; Anderson op. cit. (1980) 132; PROSEA op. cit. 469; Coode et al. (eds.) op. cit.
85. Type: Curtis 1404, Peninsular Malaysia, Penang, Government Hill (holotype CAL). Synonym:
Synaptea nitens (King) Ridl., op. cit. (1922) 241.
Large canopy tree, to 40 m tall, to 70 cm diameter. Exposed young fleshy parts densely
powdery rufous pubescent, persisting only on leaf bud, stipule, inflorescence and ovary.
Stipules narrowly oblong, subacute, to 20 × 3.5 mm, caducous. Twigs terete. Leaves
coriaceous, glistening, drying rich pinkish brown; blade narrowly oblong, 35–55 × 10–17
cm, base obtuse or cuneate, margin not revolute, apex acuminate with tapering acumen to 1
cm long; lateral veins 12–22 pairs with many short intermediates, arched, raised below, flat
or slightly elevated above as also the midrib and intercostal venation; petiole 1–2 cm long,
to 2.5 mm diameter, not drying black, geniculate. Inflorescences terminal or occasionally
axillary; rachis singly, rarely doubly, branched, to 11 cm long. Flowers: buds spindle
shaped, to 7 × 3 mm; calyx densely pubescent on both surfaces; otherwise typical. Fruits:
pedicels to 5 mm long, fruit base impressed; calyx lobes unequal, free, pubescent at first, 2
longer lobes oblong, obtuse, slightly recurved, to 13 × 2 cm, tapering to 7 mm wide at base,
3 shorter ones hastate, acute, to 3.8 × 0.9 cm, to 7 mm wide at base. Nuts globose, to 3 cm
diameter.
374
Vernacular name. Sarawak—resak daun panjang (Malay).
Distribution. Sumatra (Riau), Peninsular Malaysia and Borneo. In Sabah known from
Beaufort, Kota Kinabalu and Sipitang districts (e.g., SAN 16831, SAN 23859, SAN 36767,
and SAN A 4512) and in Sarawak from Bau, Bintulu, Kapit, Kuching, Lundu, Miri, and
Brunei (e.g., BRUN 3276 and FMS 30528) and E and W Kalimantan (e.g., Arifin Berau 570
and bb. 35295).
Ecology. Locally frequent in mixed dipterocarp forest on welldrained sandy and sandy clay
soils, on low ridges, at altitudes to 600 m. Common in Lambir NP and recorded from G.
Gading NP; elsewhere vulnerable.
21. Vatica oblongifolia Hook.f.

(Latin, oblongus = rather long, folius = leaf; the leaf shape)
Trans. Linn. Soc. 23 (1862) 160; Merrill op. cit. (1921) 409; Slooten op. cit. (1927) 109, op. cit.
264, op. cit. (1968) 34, op. cit. (1982) 355; Meijer & Wood op. cit. 314; Burgess op. cit. 227;
Anderson op. cit. (1980) 132; PROSEA op. cit. 469; Kessler & Sidiyasa, TBSAEK (1994) 113;
Coode et al. (eds.) op. cit. 86. Type: Lowe s.n., Borneo, Sabah (holotype K).
Canopy tree, to 35 m tall, to 50 cm diameter. Exposed fleshy parts including young fruit
densely evenly persistently vinoussericeous. Twigs at first compressed, c. 2–4 × 1–2 mm
apically, smooth. Stipules oblong, obtuse, to 4.5 × 1.2 cm, fugaceous. Leaves thinly to
thickly coriaceous, bluish green with purplish venation below when fresh, drying dull
greyish brown; blade flate to somewhat concave, narrowly to broadly elliptic, or narrowly to
broadly oblong, or narrowly obovate to obovateoblong, 6.5–31 × 2.5–10.5 cm, base
cuneate, obtuse, to more or less cordate, apex acute, acuminate, to subcaudate, acumen 1–
1.5 cm long; midrib prominent and terete below, sunken above; lateral veins 10–27 pairs,
with short more or less prominent intermediates, prominent below, arched within margin;
petiole 1.4–5 cm long, stout, geniculate, drying not black. Inflorescences compressed,
singly or doubly branched, terminal or in axillary cluster to 3, to 8 cm long. Flowers: buds
to 15 mm long; petals cream, purplish towards the base; otherwise typical. Fruits: pedicels
to 8 mm long; calyx lobes equal, free, thickly coriaceous, deltoid, oblong or obovate, acute,
obtuse or retuse, becoming reflexed, subrotate or revolute, to 0.3–1.5 × 0.2–1.2 cm. Nuts
globose, to 2 cm diameter, smooth, faintly 3furrowed.
Vernacular names. Sabah—resak daun panjang (preferred name). Sarawak—resak

membangan (preferred name).
Ecology. In mixed dipterocarp forest on welldrained sandy, sandy clay and shallow clay
soils, at altitudes to 1000 m.
Notes. In Sabah and Sarawak, five subspecies are recognised, with distinct albeit
overlapping ecological ranges.
375
Key to subspecies
1. Petiole at most 1.4 cm long; leaf broadly elliptic..……….………………………………

subsp. elliptifolia P.S.Ashton
(Latin, ellipticus = broadest at the middle with curved sides, folius = leaf; the leaf
shape)
Gard. Bull. Sing. 22 (1967) 265, op. cit. (1968) 34, op. cit. (1982) 356; Coode et al. (eds.)
op. cit. 86. Type: Ashton BRUN 925, Borneo, Brunei, Bt. Kukub, Andulau hills (holotype K;
isotypes KEP, L).
Twigs c. 3 × 2 mm apically. Leaf blade broadly elliptic, 12–20 × 3.5–7 cm, base
broadly cuneate, apex acute with slender acumen to 1 cm long; petiole to 1.4 cm
long. Fruits: calyx lobes oblong, to 0.8 × 0.4 cm, obtuse, revolute.
In Sarawak recorded from Miri district (e.g., S 24432). Also occurring in Brunei
(e.g., the type). Rare, in mixed dipterocarp forest on deep sandy soils on coastal
hills, and on the Arip rhyolite, at altitude below 300 m. Critically endangered.
Petiole at least 1.5 cm long; leaf various, not generally broadly elliptic………….….…2
2. Fruit calyx lobes at least 1.3 × 0.8 cm; leaf lateral veins 16–27 pairs…………...……...3
Fruit calyx lobes at most 0.4 × 0.3 cm; leaf lateral veins 11–18 pairs……...…………...4
3. Leaf blade broadly oblong, base obtuse to more or less cordate……...………………….

subsp. crassilobata P.S.Ashton
(Latin, crassus = thick, lobatus = lobed; the fruit calyx lobes)
op. cit. 86. Type: Ashton BRUN 614, Borneo, Brunei, Andulau FR (holotype K; isotypes
KEP, L).
Twigs c. 4 × 2 mm apically, stout. Leaf blade thickly coriaceous, often somewhat
concave, broadly oblong, 11.5–21 × 4.5–10.5 cm, base obtuse more or less
cordate, apex acute with short, broad acumen; lateral veins stoutly prominent
below, 16–23 pairs; petiole 1.8–2.8 cm long, stout. Fruit calyx lobes obovate, to
1.5 × 1.2 cm, obtuse to retuse, subrotate.
Known in Sarawak from Bintulu, Kapit, Miri, and Simunjan districts (e.g., S
23225, S 27127, S 39618, S 59590, and S 63662). Also occurring in Brunei (e.g.,
BRUN 615 and BRUN 5157) and W Kalimantan. Locally common in mixed
dipterocarp forest, on deep yellow sands on hills behind the swamps or on the
coast, at altitudes below 400 m. Common in Lambir NP and locally so in Mulu NP;
elsewhere endangered by forest conversion.
Leaf blade narrowly obovate, base more or less cuneate...….……………………………
subsp. multinervosa P.S.Ashton
(Latin, multus = many, nervosus = nerved; the manyveined leaf)
Gard. Bull, Sing. 22 (1967) 265, op. cit. (1968) 34, op. cit. (1982) 356; Coode et al. (eds.)
op. cit. 86. Type: Meijer SAN 19713, Borneo, Sabah, Sandakan, Sepilok FR (holotype K;
isotypes KEP, L).
Twigs c. 4 × 2 cm apically, stout. Leaf blade narrowly obovate, 14–31 × 4–8.5 cm,
base narrowly obtuse or cuneate, apex, acuminate with slender acumen to 1.5 cm
long; lateral veins 18–27 pairs, with prominent short intermediates, very
prominent below; intercostal venation distinctly elevated below; petiole 1.5–2.5 cm
long. Fruit calyx lobes oblong, to 1.3 × 0.8 cm, obtuse, reflexed.
Recorded in E Sabah from Lahad Datu, Sandakan and Tawau districts (e.g., SAN
22509, SAN 30052, SAN 34083, SAN 88089, and SAN 93092) and in Sarawak from
Belaga, Bintulu, Kapit, Limbang, Lundu, and Tatau districts (e.g., S 12623, S
22240, S 41769, S 43465, and S 66118). Also occurring in Brunei (e.g., BRUN
376
5786) and NE Kalimantan (e.g., bb. 18163 and Kostermans 8603). Locally
common in mixed dipterocarp forest on deep fertile clayrich soils, especially on
basic igneous rocks, at altitude to 800 m; also on shale, and on rhyolite. Occurring
in Lambir and Mulu NPs; not vulnerable.
4. Leaf blade narrowly elliptic, base cuneate; petiole 1.5–2.5 cm long………………….….

subsp. selakoensis P.S.Ashton
(of Selako, W Sarawak)
op. cit. 86. Type: Ilias S 13340, Borneo, Sarawak, G. Gading FR (holotype K; isotypes KEP,
L).
Twigs c. 2 × 1 mm apically, slender. Leaf blade thinly coriaceous, narrowly
elliptic, 6.5–22 × 2.5–6.5 cm, base cuneate, apex subcaudate, acumen to 1.5 cm
long; lateral veins 11–18 pairs, with short indistinct intermediates, slender, hardly
raised below; petiole 1.5–2.5 cm long. Fruit calyx lobes oblong, to 0.4 × 0.3 cm,
obtuse.
Confined to mountains in Lundu district, W Sarawak (e.g., S 13171 and S 18466).
Common in upper dipterocarp forest on shallowly humic soils, at 600–1000 m
altitude on granodiorite, mostly along ridges. Common in G. Gading NP,
vulnerable to clearing on the summits.
Leaf blade obovate to oblong, base obtuse; petiole 2.5–5 cm long………………………
subsp. oblongifolius
Twigs c. 4 × 2 mm apically, compressed, smooth. Leaf blade coriaceous, obovate
to oblong, 10–31 × 4.5–10 cm, base obtuse, apex abruptly acute to broadly
acuminate; lateral veins 10–18 pairs, with short more or less prominent
intermediates; petiole 2.5–5 cm long, stout, geniculate. Fruit calyx lobes deltoid, to
0.3 × 0.2 cm.
Widespread throughout Borneo; in Sabah known from Beaufort, Kinabatangan,
Kota Kinabalu, Kudat, Labuk Sugut, Ranau, Sandakan, and Sipitang districts (e.g.,
SAN 16424, SAN 16845, SAN 20702, SAN 77868, and SAN 97513) and in Sarawak
from Belaga, Bintulu, Kapit, Kuching, Lawas, Lundu, Marudi, Miri, and Tatau
districts (e.g., S 13305, S 21550, S 32056, S 45513, and S 66998). Also occurring in
Brunei (e.g., BRUN 2623 and Wong WKM 84) and Kalimantan (e.g., Kostermans
7299 and Kostermans 10624). Frequent and locally common in mixed dipterocarp
forest on sandy, sandy clay and shallow clay soils, at altitudes to 1000 m. Well
represented in the parks system; not vulnerable.
22. Vatica odorata (Griff.) Symington Plate 5F.

(Latin, odoratus = scented; the flowers)
J. Malay. Br. Roy. As. Soc. 19 (1941) 156, op. cit. (1943) 224; Ashton op. cit. (1964) 75, op. cit.
(1967) 263, op. cit. (1978) 23, op. cit. (1982) 360; PROSEA op. cit. 470; Kessler & Sidiyasa op. cit.
113. Basionym: Sunaptea odorata Griff., op. cit. 516. Type: Griffith s.n., Myanmar, Mergui,
Tennaserim (holotype CAL). Synonyms: Hopea grandiflora Wall. ex A.DC., op. cit. 634; Synaptea
grandiflora (Wall. ex A.DC.) Kurz op. cit. (1870) 65; Anisoptera odorata (Griff.) Kurz, Flora 30
(1872) 191; Vatica grandiflora (Wall. ex A.DC) Dyer op. cit. (1874) 301; V. faginea Dyer op. cit.
(1874) 301; V. astrotricha Hance, J. Bot. 14 (1876) 241; Sunaptea astrotricha (Hance) Pierre, For. Fl.
Coch. (1891) t. 240; Sunaptea dyeri Pierre, l.c. (1891) t. 241; Sunaptea faginea (Dyer) Pierre op. cit.
(1891) 242; V. curtisii King op. cit. 105.
377
Canopy tree, to 40 m tall, to 60 cm diameter. Exposed fleshy parts, leaf blade and venation
excepted, more or less densely pale yellowish to rustbrown pubescent; indumentum
caducous on fruit calyx, more or less persistent elsewhere. Twigs c. 1.5 mm diameter
apically, terete, rugose, frequently finely flaky. Stipules oblong, obtuse, c. 8 × 2 mm,
caducous. Leaves thinly coriaceous, drying pale yellowish brown or greyish brown; blade
narrowly elliptic to ovate, 8–16 × 2.7–5.5 cm, base obtuse or cuneate, apex acuminate,
acumen to 0.8 cm long; lateral veins 11–15 pairs, raised below, flat above as also the
midrib, not forming intramarginal vein; intercostal venation not prominent below; petiole
0.8–2 cm long, less than 3 mm diameter, geniculate or not, not drying black. Inflorescences
terminal or axillary; rachis singly branched, to 7 cm long. Flowers: buds to 8 mm long;
calyx densely shortly pale greyish brown tomentose; otherwise typical. Fruits: pedicels c. 3
mm long, slender; calyx lobes unequal, chartaceous, united at base in a to 8 mm diameter, 3
mm deep shallow cup, 2 longer lobes spatulate, obtuse, to 5.5 × 1.5 cm, tapering to 2.5 mm
broad at base, not becoming reflexed, 3 shorter ones hastate, acute, to 1.4 × 0.4 cm. Nuts
globose, to 0.7 cm diameter, with 2 mm filiform style remnant, basal half adnate to the
calyx cup.
Vernacular names. Sabah—resak biabas (preferred name). Sarawak—resak ranting kesat

(Malay).
Distribution. S China, Myanmar (Tennasserim), Thailand, IndoChina, Peninsular

Malaysia, Borneo and the Philippines.
Ecology. In aseasonal mixed dipterocarp forest and in seasonal mixed dipterocarp forest, at
altitudes to 1900 m.
Notes. Two subspecies, viz. subsp. mindanensis and subsp. odorata, are recognised in Sabah
and Sarawak.
Key to subspecies
Leaf blade drying pale yellowish brown; petiole not geniculate, 0.8–1.2 cm long. In
aseasonal and seasonal mixed dipterocarp forest at altitudes to 700 m……………………….
subsp. odorata
In Borneo, known from Sipitang and Tawau districts in Sabah (e.g., SAN 16714, SAN
17159 and SAN 40586) and E Kalimantan (e.g., Ambri & Arifin 1016). In mixed
dipterocarp forest at altitudes to 700 m. In Sabah rare and endangered.
Leaf blade drying greyish brown; petiole geniculate, 1.5–2 cm long. In aseasonal mixed
dipterocarp and upper dipterocarp forest at altitudes to 1900 m………………………………
subsp. mindanensis (Foxw.) P.S.Ashton
(of Mindanao, the Philippines)
Gard. Bull. Sing. 22 (1967) 263, op. cit. (1978) 24, op. cit. (1982) 361; PROSEA op. cit. 470;
Coode et al. (eds.) op. cit. 86. Basionym: Vatica mindanensis Foxw. in Elmer Leafl. Philip. Bot.
6 (1913) 1957, Slooten op. cit. (1927) 71, Meijer & Wood op. cit. 313, Burgess op. cit. 227.
Lectotype (designated here): Elmer 13398, the Philippines, Mindanao, Mt. Urdaneta
(hololectotype K). Synonyms: V. sorsogonensis Foxw., Philip. J. Sci. 13 (1918) Bot. 196; V.
aerea Slooten op. cit. (1941) 133.
In Borneo and the Philippines. In Sabah recorded from Lahad Datu, Ranau, Sipitang,
Tambunan, and Tawau districts (e.g., SAN 16354, SAN 16667, SAN 17036, SAN 38436,
and SAN 95838) and in Sarawak from Bau, Belaga, Kapit, Kuching, Lawas, Limbang,
Lubok Antu, Lundu, Samarahan, Sri Aman, and Tatau districts (e.g., S 10061, S 19014,
378
S 24704, S 36676, and S 68183). Also occurring in Brunei (e.g., BRUN 2540 and BRUN
5237) and W Kalimantan (e.g., Suzuki K 9658 and Suzuki K 9994). In mixed and upper
dipterocarp forest, at altitudes to 1900 m, on clays soils overlying shale, dacite,
granodiorite, and basalt. Probably not vulnerable.
23. Vatica parvifolia P.S.Ashton

(Latin, parvus = small, folius = leaf)
op. cit. (1980) 133; Coode et al. (eds.) op. cit. 86. Type: Ariffin S 371, Borneo, Sarawak, Lundu
district, Bt. Sebandar (holotype K; isotype KEP).
Canopy tree, to 35 m tall, to 35 cm diameter. Exposed fleshy parts and ovary persistently
densely rufousbrown powdery pubescent. Twigs terete, to 1 mm diameter apically, slender,
muchbranched. Stipules linear, to 5 × 1 mm, caducous. Leaves coriaceous, drying reddish
brown; blade narrowly ovate to lanceolate, 2.8–6 × 1–2.3 cm, base obtuse, margin not
revolute, apex caudate with acumen to 1 cm long; midrib slender, prominent below, flat
above; lateral veins c. 8 pairs, hardly raised, indistinct, arched; petiole 0.6–0.9 cm long, not
drying black. Inflorescences terminal or axillary; rachis terete, singly branched, to 2 cm
long. Flowers: buds to 6 mm long; calyx greyish brown pubescent; otherwise typical.
Fruits: pedicels to 3 mm long, slender; calyx lobes unequal, free, chartaceous, glabrescent
except towards base, 2 longer lobes oblong, narrowly obtuse, to 6 × 0.7 cm, revolute above
the abruptly narrowly constricted base, not becoming reflexed; 3 shorter ones broadly ovate,
to 1 × 0.7 cm, acute, cordate, prominently revolute. Nuts broadly ovoid, to 0.5 × 0.35 cm,
obtuse.
Vernacular name. Sarawak—resak kerangas padi (preferred name).
Distribution. Endemic in Borneo. In Sabah known from Sandakan district (e.g., SAN
16370) and in Sarawak from Kuching, Limbang and Lundu districts (e.g., S 5318, S 6001, S
6833, S 7537, and S 12508). Also occurring in Brunei (e.g., BRUN 2007 and FMS 34475).
Ecology. Very local, in kerangas forest on shallow and giant podsols, at altitudes below 300
m. Critically endangered.
24. Vatica patentinervia P.S.Ashton

(Latin, patens = diverging from the axis at a wide angle, nervia = nerve; the lateral veins of
the leaf)
Gard. Bull. Sing. 54 (2002) 214. Type: Ilias S 15148, Borneo, Sarawak, Bintulu district, Segan FR
(holotype K; isotypes KEP, SAR).
Mediumsized canopy tree, to 35 m tall, to 35 cm diameter. Young fleshy parts pale

yellowish buff scabridpuberulent; indumentum persistent on petiole, leaf venation below
and midrib above. Twigs c. 2 mm diameter apically, terete, more or less ribbed. Stipules
not seen. Leaves coriaceous, drying dull pale yellowish brown; blade elliptic to obovate,
12–18 × 4.5–7 cm, base obtuse, margin not revolute, apex obtuse to shortly acuminate;
venation including midrib somewhat elevated above, more or less prominently so below;
lateral veins 7–8 pairs, distinctly raised below; petiole to 2.5 cm long, to 3 mm diameter,
379
drying pale buff brown, rugose. Inflorescences ramiflorous, in axillary clusters of 3, or

terminal; rachis singly branched, to 3 cm long. Flowers unknown. Fruits: pedicels to 4 mm
long, slender; calyx lobes unequal, free, chartaceous, 2 longer lobes lorateoblong, obtuse,
to 12 × 2.2 cm, constricted and somewhat revolute at base, 3 shorter ones hastate, to 4.5 ×
1.5 cm, acuminate, revolute. Nuts ellipsoid, to 1.2 × 0.8 cm, mucronate, glabrous.
Distribution. Known only from the type locality in Sarawak, and Kuala Belalong,
Temburong district in Brunei (e.g., S 5744).
Ecology. Apparently locally frequent in Temburong; in both localities in mixed dipterocarp

forest on clay soils, at altitudes below 400 m. Conservation status uncertain.
25. Vatica pedicellata Brandis

(Latin, pedicellatus = with a prominent stalk; the flowers)
J. Linn. Soc. Bot. 31 (1895) 125; Merrill op. cit. (1921) 409; Slooten op. cit. (1927) 111; Masamune
op. cit. 499; Browne op. cit. 101; Ashton op. cit. (1968) 35, op. cit. (1982) 356; Anderson op. cit.
(1980) 133. Type: Haviland 1041/869, Borneo, Sarawak, Lundu district (holotype K; isotype KEP).
Small tree, to 15 m tall, to 15 cm diameter. Exposed fleshy parts, leaf surface excepted, and
ovary persistently densely vinoussericeous. Twigs c. 2 mm diameter apically, terete or
slightly compressed only, hardly branched, smooth. Stipules lorate, subacute, to 7 × 2 mm,
caducous. Leaves coriaceous, bluish green with purplish venation below when fresh, drying
mauvebrown; blade elliptic to lanceolate, 9–23 × 3.5–7.5 cm, base obtuse or rarely
cuneate, margin narrowly revolute, apex with slender acumen to 2 cm long; midrib
prominent below, elevated within a furrow above; lateral veins 9–15 pairs, with many short
slender intermediates, slender, hardly raised on either surface though more so below, arched
and anastomosing to form a faint looped intramarginal vein; petiole 1–2.5 cm long, 2–3 mm
diameter, not or hardly geniculate, terete but rugulose and not turning black on drying.
Inflorescences axillary; rachis singly or doubly branched, compressed, ribbed, slender, to 8
cm long. Flowers: buds spindleshaped, to 6 × 2 mm; calyx vinous puberulent; corolla
cream with a purplish suffusion outside; otherwise typical. Fruits vinous cinereous
throughout; pedicels to 8 mm long, slender; calyx lobes free, subequal, oblong, to 0.6 × 0.4
cm, obtuse, incrassate, revolute, reflexed. Nuts subglobose, to 1.8 × 2.2 cm, subacute, with
obscure sutures.
Distribution. Endemic in Borneo; known only from Kuching, Lundu and Simunjan districts
in Sarawak (e.g., S 4672, S 7252, S 15233, S 39416, and S 68078).
Ecology. Locally frequent in kerangas forest, usually on rocky slopes near the sea. Well
represented in Bako NP; not vulnerable.
26. Vatica perakensis King

(of Perak, Peninsular Malaysia)
J. As. Soc. Beng. 62, 2 (1893) 103, p.p. emend. Symington op. cit. (1941) 152, op. cit. (1943) 226;
Slooten op. cit. (1927) 86, p.p.; Ashton op. cit. (1978) 24, op. cit. (1982) 367. Lectotype (Symington,
1941): King’s Collectors 7549, Peninsular Malaysia, Perak, Larut (hololectotype CAL). Synonyms:
Synaptea perakensis (King) Ridl., op. cit. (1922) 242, p.p.; Vatica songa Slooten op. cit. (1927) 93.
380
Small to mediumsized tree, to 30 m tall, to 30 cm diameter. Exposed fleshy parts, leaf

blade excepted, and ovary persistently greyish pink to pale yellowish brown cinereous; leaf
venation below and nut glabrescent. Twigs c. 1 mm diameter apically, slender, terete,
muchbranched. Stipules fugaceous, not seen. Leaves thinly coriaceous, drying pale
greyish brown; blade lanceolate to oblanceolate, 6–14 × 1.8–3.8 cm, base cuneate, margin
not revolute, apex subcaudate, acumen to 2 cm long; lateral veins 11–13 pairs, slender but
prominent below, evident above, as also the midrib, ascending; intercostal venation densely
subreticulate, evident on both surfaces; petiole slender, 1–2 cm long, not drying black.
Inflorescence slender; rachis singly branched, to 5 cm long. Flowers typical. Fruits:
pedicels to 6 mm long, slender; calyx lobes unequal, free, chartaceous, 2 longer lobes
spatulate, obtuse, to 6 × 1.8 cm, tapering to c. 5 mm wide at the subrevolute base, 3 shorter
ones lanceolate, acute, to 1.5 × 0.6 cm. Nuts ovoid, apiculate, to 1 cm diameter.
Distribution. Sumatra, Peninsular Malaysia and Borneo. In Borneo two tentative records
from Lahad Datu and Tawau districts in SE Sabah (i.e., SAN 20119 and SAN 41580).
Ecology. In ‘coastal hill dipterocarp forest’ of Symington (op. cit. (1943) 15, fig. 2) on dry
ridges, especially near the coast. Conservation status unknown but likely endangered.
27. Vatica rassak (Korth.) Blume

(after the Malay common name—resak)
Mus. Bot. Lugd.Bat. 2 (1852) 31 (incl. var. subcordata Blume); Merrill op. cit. (1921) 409; Slooten
op. cit. (1927) 73, 104, op. cit. (1942) 223; Masamune op. cit. 499; Ashton op. cit. (1978) 20, op. cit.
(1982) 353; PROSEA op. cit. 471; Kessler & Sidiyasa op. cit. 114. Basionym: Retinodendron rassak
Korth., op. cit. 56. Type: Korthals s.n. (= RHL Sheet No. 900171121), Borneo, Kalimantan, banks of
Sg. Barito, Lower Dyak (holotype L). Synonyms: Vateria rassak (Korth.) Walp., Rep. 5 (1845) 126;
Vatica papuana Dyer, J. Bot. 16 (1878) 100, Slooten op. cit. (1927) 73, op. cit. (1942) 233, Meijer &
Wood op. cit. 314, Burgess op. cit. 227, Ashton op. cit. (1968) 35, Anderson op. cit. (1980) 132;
Vatica moluccana Burck op. cit. 226; R. moluccanum (Burck) F.Heim op. cit. (1892) 104; Vatica
schumanniana Gilg, Bot. Jahrb. 18 (1894) Beibl. 45; Vatica celebensis Brandis op. cit. 126; Vatica
subcordata (Blume) Hallier f., Med. Rijksherb. 36 (1918) 4, Slooten op. cit. (1942) 237; Vatica
celebica Slooten op. cit. (1942) 237.
Large canopy tree, to 30 m tall, to 70 cm diameter. Exposed fleshy parts, leaf surface
excepted, and ovary persistently evenly pale buffpuberulent; fruit glabrescent. Twigs c. 3
mm diameter apically, stout, crooked, ribbed, becoming rugose, flaky; stipule scars
prominent. Stipules lorate, subacute, to 14 × 4 mm, subpersistent. Leaves thickly
coriaceous, drying dull greyish brown; blade oblong to narrowly elliptic, 13–32 × 5–11 cm,
base broadly cuneate to subcordate, apex with tapering acumen to 1.5 cm long; midrib
prominent below, flat above; lateral veins 16–20 pairs, prominent below, slightly elevated
above; petiole 2–2.5 cm long, at least 3 mm diameter, stout, not geniculate, not drying
black. Inflorescences terminal or axillary; rachis ribbed, irregularly branched mostly from
the base, appearing fascicled, to 14 cm long. Flowers: buds spindleshaped, to 14 × 3 mm;
calyx densely shortly pale buff pubescent; otherwise typical. Fruits: pedicels to 3 mm long,
short, stout; calyx lobes free, subequal, deltoid, acute, incrassate, reflexed, recurved, to 1.2
× 0.7 cm. Nuts oblong to ovoid, to 5 × 3.5 cm, very variable, obtuse to taperingapiculate,
frequently asymmetric, verrucose with thick corky pericarp, prominently 3furrowed.
381
Distribution. Borneo, Sulawesi, Maluku, New Guinea, and Sudest Is. In Sabah recorded
from Beaufort, Kinabatangan, Lahad Datu, Papar, Sandakan, Semporna, and Sipitang
Sarawak from Belaga and Sibu districts (e.g., S 9727 and S 39310). Also occurring in S
Kalimantan (e.g., Kessler and Ambriansyah B1440 and Sidiyasa PBU 443). The furthest
east known dipterocarp.
Ecology. On and near banks of sluggish including brackish rivers in Sabah and Sarawak, on
low hills in lowland mixed forest elsewhere. Vulnerable owing to its accessible habitat.
28. Vatica rotata P.S.Ashton

(Latin, rotatus = like a wheel; the spreading fruit calyx lobes)
Type: Ashton S 19594, Borneo, Sarawak, Mukah, Ulu Kenyana (holotype K; isotypes KEP, L).
Small tree, c. 10 cm diameter. Exposed fleshy parts, including midrib below and fruit but
excluding leaf blade, persistently densely vinoussericeous. Twigs c. 1 mm diameter
apically, slender, muchbranched, terete, smooth. Stipules not seen. Leaves coriaceous,
bluish green with purplish midrib below when fresh, drying dull greyish brown; blade
broadly ellipticovate, 5.5–10 × 3–5 cm, base obtuse, apex with slender acumen to 0.8 cm
long; midrib slender but prominent below, evident towards base above, otherwise obscurely
sunken; lateral veins 9–11 pairs, without intermediates, slender, arched, hardly raised;
intercostal venation densely reticulate, evident below; petiole 0.8–1.2 cm long, slender, not
drying black. Inflorescences terminal or axillary; rachis slender, terete, singly branched, to
1.5 cm long. Flowers typical. Fruits: pedicels to 9 mm long, slender; calyx lobes free,
equal, suborbicular, incrassate, revolute, rotate, to 0.7 × 0.4 cm. Nuts subglobose, c. 0.4
cm diameter (immature), obtuse, indistinctly 3sutured.
Distribution. Endemic in Borneo. Known in Sabah from Labuk Sugut district (e.g., SAN
128886 and SAN 130743) and in Sarawak from Bintulu and Mukah districts (e.g., S 24533
and type). Also occurring in W Kalimantan.
Ecology. Rare, in mixed dipterocarp forest on deep yellow sands on low hills not far from
the coast. Possibly extinct in Sarawak, otherwise critically endangered.
29. Vatica rynchocarpa P.S.Ashton

(Greek, rynchos = snout, carpos = fruit; the beaked nut)
Gard. Bull. Sing. 22 (1967) 270, op. cit. (1964) 80 (‘19. Vatica sp.’), op. cit. (1968) 36, op. cit. (1982)
365; Anderson op. cit. (1980) 133; Coode et al. (eds.) op. cit. 86. Type: Egon SA 514, Borneo,
Sarawak, Rajang, Nanga Temulang (holotype KEP).
Canopy tree, to 30 m tall, to 1 m diameter. Exposed fleshy parts, leaf blade and venation
excepted, and ovary shortly sparsely pale greyish brown puberulent. Twigs c. 1 mm
diameter apically, slender, terete, smooth. Stipules narrowly hastate, acute, to 2.5 mm long,
fugaceous. Leaves thinly coriaceous, drying reddish brown; blade narrowly elliptic
lanceolate, 5–8.5 × 1.3–2.5 cm, base obtuse, margin not revolute, apex acuminate with
slender acumen to 1 cm long; midrib slender, raised below, obscurely sunken, especially
382
towards base, above; lateral veins 10–14 pairs, with short obscure intermediates, hardly
raised, wellspaced, arched within margin; petiole to 0.4 cm long, drying black or not.
Inflorescences terminal or axillary; rachis singly branched, slender, to 3 cm long. Flowers
unknown. Fruits: pedicels to 7 mm long, slender; calyx lobes unequal, free, chartaceaous,
2 longer lobes spatulate, narrowly obtuse, to 6.2 × 1.4 cm, tapering to 2.5 mm wide at base,
not becoming reflexed, 3 shorter ones hastate, acute, to 1.5 × 0.25 cm. Nuts ovoid, to 1.8 ×
0.8 cm, tapering to 4 mm style remnant, glabrous, drying black.
Distribution. Endemic in Borneo. Recorded in Sabah from Kinabatangan district (e.g., SAN
A 4734) and in Sarawak from Bintulu, Kapit, Kuching, and Miri districts (e.g., S 12107, S
16568, S 27292, and S 68711). Also occurring in Brunei (e.g., BRUN 3354 and Niga NN
289) and Kalimantan (e.g., Kostermans 10005).
Ecology. Scattered, in forest on clay alluvium river banks. Vulnerable on account of its
habitat loss.
30. Vatica sarawakensis F.Heim

(of Sarawak)
Bull. Mens. Soc. Linn. Paris 2 (1891) 109; Slooten op. cit. (1927) 106; Masamune op. cit. 499;
Browne op. cit. 101; Ashton op. cit. (1963) 252, op. cit. (1964) 77, op. cit. (1967) 262, op. cit. (1968)
PROSEA op. cit. 471; Coode et al. (eds.) op. cit. 86. Type: Beccari PB 3018, Borneo, Sarawak
(holotype P). Synonyms: Retinodendropsis aspera F.Heim op. cit. (1893) 470; Vatica ramiflora
Slooten op. cit. (1927) 118, p.p.
Subcanopy tree, to 25 m tall, to 60 cm diameter. Exposed fleshy parts, leaf blade excepted,
densely pale yellowish or rufousbrown scabridtomentose; leaf venation below sparsely
so; ovary evenly so; fruit glabrescent. Twigs to 5 mm diameter apically, stout, ribbed, finely
cracked. Stipules narrowly hastate, acute, to 15 × 5 mm. Leaves thinly coriaceous, drying
greyish brown; blade oblong to obovate, 22–35 × 7–15 cm, base obtuse, apex with tapering
acumen to 1.4 cm long; lateral veins 15–28 pairs, with or without distinct intermediates,
prominent below, flat or slightly raised above; petiole 1–2 cm long, to 4 mm diameter, stout,
not drying black. Inflorescences in axillary cluster to 3 or ramiflorous, rarely terminal;
rachis muchbranched, to 12 cm long. Flowers: buds to 8 mm long; calyx shortly reddish
brown pubescent, otherwise typical. Fruits glabrous; pedicels to 2 mm long; calyx lobes
free, equal, oblonghastate, obtuse or acute, thickly coriaceous, becoming reflexed and
somewhat revolute, to 1.6 × 0.7 cm. Nuts symmetric, subglobose to ovoid, to 2.5 × 2.5 cm,
acute with short but prominent style remnant, verrucoselenticellate, prominently 3
furrowed.
Vernacular name. Sarawak—resak daun besar (preferred name).
Distribution. Endemic in Borneo. Recorded in Sabah from Beaufort, Keningau, Kota

Kinabalu, Kuala Penyu, Kinabatangan, Lahad Datu, Sandakan, Sipitang, and Tawau
Sarawak from Kapit, Kuching, Lawas, Lundu, Marudi, and Samarahan districts (e.g., S
13187, S 24225, S 37787, S 41308, and S 60973). Also known in Brunei (e.g., Dransfield
JD 7491 and FMS 30382) and E Kalimantan (e.g., Argent et al. 93156, bb. 7029 and bb.
10796).
383
Ecology. Scattered in mixed dipterocarp forest on clay soils, on low hills, often in
floodplains, at altitude to 1400 m. Locally common in Lambir NP and recorded from G.
Gading and Mulu NPs; elsewhere vulnerable owing to forest conversion.
Notes. Fruit collected from E Sabah and E Kalimantan are prominently cuspidate, their
sepals to 2 cm long, narrow and strongly reflexed, conforming with the type of V. ramiflora.
Though immature fruit from Sarawak and W and C Kalimantan are similarly cuspidate, the
ripe fruit there becomes obtuse, with sepals at most to 7 mm long, conforming with the type
of V. sarawakensis. Further collections may justify the recognition of two subspecies.
31. Vatica umbonata (Hook.f.) Burck Fig. 36

(Latin, umbonatus = with a central boss; the nut apex)
Ann. Jard. Bot. Buitenz. 6 (1887) 222; Slooten op. cit. (1927) 132; Ashton op. cit. (1963) 250, op. cit.
(1964) 78, op. cit. (1968) 36, op. cit. (1978) 17, op. cit. (1982) 349; Meijer & Wood op. cit. 320;
115; Coode et al. (eds.) op. cit. 86. Basionym: Pachynocarpus umbonatus Hook.f., op. cit. (1860)
159, Merrill op. cit. (1921) 409. Type: Motley s.n., Borneo, Labuan (holotype K). Synonyms: Vatica
verrucosa Burck op. cit. 232; P. verrucosus (Burck) F.Heim op. cit. (1892) 107; V. blancoana Elmer
op. cit. (1912) 1473; V. cupularis Slooten op. cit. (1927) 132, Browne op. cit. 100; V. ramiflora
Slooten op. cit. (1927) 118, p.p.
Mediumsized tree, to 30 m tall, to 70 cm diameter, often crooked and shrublike. Exposed

fleshy parts, leaf blade excepted, and ovary more or less sparsely greyish brown puberulent,
caducous except on bud and stipule. Twigs to 3 mm diameter apically, stout, brittle, much
branched, striated, cracked and sometimes thinly flaky. Stipules hastate, subacute, to 4 × 2
mm, early caducous. Leaves somewhat coriaceous, drying pale greyish brown; blade
elliptic, often asymmetric, 8–16 × 3–6.5 cm, base cuneate, apex obtuse to shortly
acuminate; midrib drying hardly paler than petiole, raised below, flat or raised above,
lateral veins 7–8 pairs, without intermediates, raised below, unraised above, slightly
arched; intercostal venation subscalariform; petiole 0.7–1.5 cm long, glabrescent, drying
black. Inflorescences axillary or terminal; rachis singly or doubly branched, brittle and
falling apart as fruit develop, to 12 cm long. Flowers: buds to 10 mm long; calyx curved
pubescent; otherwise typical. Fruits: pedicels 2–5 mm long; calyx lobes equal, fused onto
nut, as 5 oblongelliptic corky verrucose plates to 2.6 × 1.8 cm, the whole with the corky
verrucose nut apex subglobose, to 3 cm diameter; or nut ovoid, acute, with the sepals fused
with it in the basal third only and reflexed.
Vernacular name. Sarawak—resak air (preferred name).
Notes. In Sabah and Sarawak, two subspecies, viz. subsp. acrocarpa and subsp. umbonata,
are recognised.
Key to subspecies
Nut subglobose, calyx lobes completely fused to it except the verrucose apex, not reflexed...
subsp. umbonata
Peninsular Malaysia, Borneo and Palawan (the Philippines). In Sabah recorded from
Beaufort, Keningau, Kota Kinabalu, Lahad Datu, Pensiangan, Ranau, Sandakan,
Sipitang, and Tenom districts (e.g., SAN 20492, SAN 29348, SAN 43156, SAN 80943,
384
Fig. 36. Vatica umbonata subsp. acrocarpa (M–O), subsp. umbonata (A–L). A, flowering
leafy twig; B, open flower; C, longitudinal section of open flower; D, abaxial view of outer
sepal; E, abaxial view of inner sepal; F, abaxial view of petal; G, adaxial view of petal; H,
adaxial view of stamens; I, abaxial view of stamens; J, gynoecium; K, side view of fruit; L,
apical view of fruit; M, fruiting leafy twig; N, side view of fruit; O, apical view of fruit. (A–
J from BRUN 1380, K–L from S 49284, M from SAN 15236, N–O from SAN 31657.)
385
and SAN 125248) and in Sarawak from Belaga, Bintulu, Kapit, Kuching, Lawas,
Lundu, Marudi, Miri, Serian, and Sri Aman districts (e.g., S 9346, S 18567, S 28108, S
39099, and S 68929). Also occurring in Brunei (e.g., BRUN 933 and BRUN 5883) and
Kalimantan (e.g., Argent et al. 93168 and Kostermans 5580).
Locally abundant on alluvium river banks and forming dense monospecific stands; also
scattered in mixed dipterocarp forest, usually on poor sandy clay soils and in upper
dipterocarp forest, at altitudes to 1300 m. Common on the Kelabit Highlands in
Sarawak; not vulnerable.
Nut ovoid, acute, with calyx lobes fused to it in the basal third only and reflexed…………...
subsp. acrocarpa (Slooten) P.S.Ashton
(Greek, across = terminal, carpos = fruit; perhaps referring to the position of the fruit in
the type specimen)
Gard. Bull. Sing. 31 (1978) 17, op. cit. (1982) 349; PROSEA op. cit. 472. Basionym: Vatica
acrocarpa Slooten op. cit. (1942) 241. Type: Maidin SAN 4298, Borneo, Sabah, Tawau, Umas
Umas (holotype K).
Endemic in Borneo. In Sabah known from Kinabatangan, Kudat, Labuk Sugut, Lahad
Datu, Sandakan, Tawau, and Tenom districts (e.g., SAN 15236, SAN 20014, SAN
25452, SAN 36146, and SAN 91909). Also occurring in E Kalimantan (e.g., Kostermans
5334 and Kostermans 10757). In forest along river banks or alluvium. Not vulnerable.
32. Vatica venulosa Blume

(Latin, venulosus = with tiny veins; the delicate leaf venation)
Mus. Bot. Lugd.Bat. 2 (1852) 32; Merrill op. cit. (1921) 409; Slooten op. cit. (1927) 78; Masamune
op. cit. 499; Ashton op. cit. (1964) 79, op. cit. (1968) 36, op. cit. (1978) 17, op. cit. (1982) 350;
Anderson op. cit. (1980) 133; PROSEA op. cit. 473; Kessler & Sidiyasa op. cit. 115; Coode et al.
(eds.) op. cit. 87. Type: Müller s.n. (= RHL Sheet No. 902146794), Borneo, S Kalimantan, Patai
(holotype L). Synonyms: Vatica bancana Scheff., Nat. Tijd. Ned. Ind. 31 (1870) 348, Meijer & Wood
op. cit. 305, Burgess op. cit. 227; V. schouteniana Scheff., op. cit. 408; Dryobalanops schefferi Hance
op. cit. 307; Retinodendron bancanum (Scheff.) King op. cit. 129; R. kunstleri King op. cit. 129; V.
kunstleri (King) Brandis op. cit. 127; V. schefferi (Hance) Brandis op. cit. 128; V. lutea Ridl., Bull.
Misc. Inform. Kew (1926) 60.
Distribution. Sumatra, Peninsular Malaysia, W Java, and Borneo.
Notes. Two subspecies, viz., subsp. simalurensis (Slooten) P.S.Ashton and subsp. venulosa
are recognised but only subsp. venulosa occurs in Sabah and Sarawak.
subsp. venulosa
Mediumsized tree, to 25 m tall, to 50 cm diameter; bole frequently crooked or leaning.

Exposed fleshy parts, including midrib above and ovary densely persistently pale pinkish
brown puberulent; blade below caducously so. Twigs to 1.5 mm diameter apically, terete,
smooth. Stipules hastate, to 5 mm long, fugaceous. Leaves thinly coriaceous, drying dark
reddish brown above, pale tawny below; blade elliptic, ovatelanceolate to narrowly
abovate, 4–12 × 1.5–5 cm, base cuneate, margin not revolute, apex shortly acuminate;
midrib prominent below, slightly sunken above; lateral veins 7–12 pairs, without
intermediates, slender, hardly raised below, unraised above, arched; petiole 0.5–0.9 cm
long, slender, not drying black. Inflorescences terminal or axillary; rachis singly branched,
ribbed, to 3 cm long. Flowers: buds to 13 mm long; calyx pinkish brown pubescent;
386
otherwise typical. Fruits: pedicels to 2 mm long, hidden in the calyx base; calyx lobes
equal, free, chartaceous, ovate acute, cordate, revolute, to 3 × 1.3 cm. Nuts globose, to 1
cm diameter, with to 1.5 mm long style remnant, pubescent, hidden within calyx.
Vernacular names. Sabah—resak banka (preferred name). Sarawak—resak letup

(preferred name).
Distribution. In Borneo, recorded in Sabah from Kinabatangan and Papar districts (e.g.,
SAN 17654 and SAN 49554) and in Sarawak from Kuching and Lundu districts (e.g.,
Haviland 2440, SA 461 and SA 577). Also occurring in Brunei (e.g., Awong AK 13 and FMS
35704) and Kalimantan (e.g., bb. 22920 and Kostermans 6087).
Ecology. Apparently very local though possibly overlooked, on clay alluvium banks of
sluggish rivers, at low altitudes. Vulnerable owing to its riparian lowland habitat.
33. Vatica vinosa P.S.Ashton

(Latin, vinosus = coloured like a grape; the tomentum)
op. cit. (1980) 133; PROSEA op. cit. 473; Coode et al. (eds.) op. cit. 87. Type: Ashton BRUN 764,
Borneo, Brunei, Temburong district, Kuala Sekurop (holotype K; isotype KEP).
Small to mediumsized subcanopy tree, to 30 m tall, to 40 cm diameter. Exposed fleshy

parts, including leaf venation below and ovary persistently densely rich vinoussericeous.
Twigs to 1.5 mm diameter apically, slender, terete, frequently cracked and flaky. Stipules
oblong, subacute, to 5 × 2.5 mm, caducous. Leaves thinly coriaceous, bluish green with
purple venation and petiole when fresh, drying pale greyish brown; blade elliptic to
lanceolate, 6–15 × 1.5–4.5 cm, base narrowly obtuse to cuneate, apex subcaudate, acumen
to 1.2 cm long; midrib slender but prominent below, flat above; lateral veins 12–20 pairs,
with short slender intermediates, slender but prominent below, arched; petiole 0.6–1.3 cm
long, less than 1 mm diameter, slender, drying not black. Inflorescences terminal or
axillary; rachis singly or doubly branched, terete, to 7 cm long. Flowers: buds to 6 mm
long; calyx sparsely vinoussericeous; otherwise typical. Fruits vinous cinereous; pedicels
to 4 mm long, slender; calyx lobes free, equal, obovate, revolute, incrassate, reflexed at
base but curving outwards and rotate apically, to 0.9 × 0.8 cm. Nuts globose, to 0.8 mm
diameter, with short style remnant.
Vernacular name. Sarawak—resak tangkai ungu (preferred name).
Distribution. Endemic in Borneo. Recorded in Sabah from Keningau, Kinabatangan, Lahad

Datu, Pitas, Semporna, Sipitang, and Tawau districts (e.g., SAN 26184, SAN 63072, SAN
85017, SAN 92359, and SAN 121351) and in Sarawak from Belaga, Bintulu, Kapit, Marudi,
and Miri districts (e.g., S 13326, S 22287, S 34693, S 43707, and S 46571). Also occurring
in Brunei (e.g., BRUN 3383 and Coode et al. 7849) and Kalimantan (e.g., Arifin et al. 7847
and Kostermans 9146).
387
Ecology. In mixed dipterocarp forest on clay rich soils on hills, at altitudes below 300 m.
Vulnerable owing to land conversion.
388
PLATES
493
A B
C D
E F
Plate 2. Dipterocarpaceae: A–B, Anisoptera reticulata; C, Dipterocarpus

grandiflorus; D–F, Dipterocarpus oblongifolius.
(Photo credits: A–B = E. Soepadmo; C = L.S.L. Chua; D = L.G. Saw; E–F = E. Soepadmo).
495
A B
C D
E F
Plate 3. Dipterocarpaceae: A–B, Dryobalanops aromatica; C,

Dryobalanops beccarii; D, Dryobalanops oblongifolia; E, Hopea
beccariana; F, Shorea coriacea.
(Photo credits: A = H.S. Yong; B = L.S.L. Chua; C = Stephen Teo; D = L.S.L. Chua; E–F = L.G. Saw).
496
A B
E F
Plate 4. Dipterocarpaceae: A, Shorea exelliptica; B–C, Shorea

macrophylla; D, Shorea multiflora; E, Shorea rotundifolia; F, Shorea
seminis.
Photo credits: A = L.G. Saw; B–D = L.S.L. Chua; E = W.K. Soh; F = L.S.L. Chua).
497
A
B
C D
E F
Plate 5. Dipterocarpaceae: A–B, Shorea smithiana; C–D, Upuna

borneensis; E, Vatica nitens; F, Vatica odorata.
(Photo credits: A = W.K. Soh; B = L.S.L. Chua; C = W.K. Soh; D–F = L.S.L. Chua).
498

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TREE FLORA

MINISTRY OF SCIENCE, TECHNOLOGY AND INNOVATION

Tree Flora of Sabah and Sarawak Editorial Committee ¾ Chairman:

E. Soepadmo, L.G. Saw and R.C.K. Chung

Sabah Forestry Forest Research Sarawak Forestry

electronic version prepared by

First published 30 September 2004

Perpustakaan Negara Malaysia Cataloguing­in­Publication Data

1. Apocynaceae (David J. Middleton) 1

Appendix: New Dipterocarp Species from Sabah and Sarawak 479

Abbreviations of Frequently Cited References 481

Commonly Used Abbreviations for Localities 485

Index to Trade Groups of Shorea (Dipterocarpaceae) 501

Index to Scientific Names 503

Index to Vernacular Names 517

Phytogeography. The subfam. Dipterocarpoideae comprise 13 genera, with c. 475 species,

Within NW Borneo, dipterocarps manifest rather distinct geographical provinces, partially

Ecology. Members of the subfamily Dipterocarpoideae are overwhelmingly concentrated in

Conservation status. I have endeavoured, on available evidence and personal experience,

Field key to the genera and main field groups

8. Leaf lateral veins densely parallel, variously unequal to subequal...........................…

9. Wood dense, hard, ray ends not glistening on tangential surface……………………...10

kelapok (Iban), mersawa (Malay), pengiran (preferred name in Sabah)

Key to Anisoptera species

2. Leaf greyish brown or bright yellow lepidote below………………………………..…3

Key to Anisoptera species

1. Anisoptera costata Korth.

cm diameter, puberulent; calyx lobes glabrous, 2 longer lobes spatulate, to 16 × 2 cm,

Vernacular names. Sabah—pengiran kesat (preferred name). Sarawak—mersawa kesat

Distribution. Coastal hills of Myanmar, S and C Thailand, Cambodia and Cochin­China,

Ecology. Evergreen and semi­evergreen lowland forests of Indo­Burma; in mixed

2. Anisoptera grossivenia Slooten

Vernacular names. Sabah—pengiran kunyit (preferred name). Sarawak—mersawa kunyit

Distribution. Endemic in Borneo. Widespread throughout the lowlands of Sabah and

3. Anisoptera laevis Ridl. Fig. 1.

Vast emergent tree, to 65 m tall, to 2 m diameter; buttresses to 15 m tall, 4 m wide, stout.

Vernacular names. Sabah—pengiran durian (preferred name). Sarawak—mersawa durian

Distribution. Sumatra, Peninsular Malaysia, Singapore, and Borneo. In Borneo, known in

4. Anisoptera marginata Korth. Fig. 2G–H.

Vernacular names. Sabah—pengiran kerangas (preferred name). Sarawak—mersawa paya

Distribution. Sumatra, Peninsular Malaysia and Borneo (widespread). In Sabah known

5. Anisoptera reticulata P.S.Ashton Fig. 2A–F, Plates 2A–B.

Vernacular name. Sabah—pengiran gajah (preferred name).

Distribution. Endemic in Borneo. In Sabah recorded from Kinabatangan, Labuk Sugut,

resak (preferred name)

Distribution. Five species distributed in Sri Lanka, Peninsular Thailand, Sumatra,

Key to Cotylelobium species

1. Leaf blade sparsely puberulent or glabrescent, drying dull olive­brown…………………

1. Cotylelobium burckii (F.Heim) F.Heim Fig. 3A–L

Vernacular name. Sarawak—resak durian (preferred name).

2. Cotylelobium lanceolatum Craib Fig. 3M–P

Vernacular names. Sabah—resak kelabu (preferred name). Sarawak—resak batu

Distribution. Peninsular Thailand to eastern coastal area of Peninsular Malaysia, Anambas

3. Cotylelobium melanoxylon (Hook.f.) Pierre Fig. 4.

Vernacular names. Sabah—resak tempurung (preferred name). Sarawak—resak hitam

keruing (preferred name), kesugoi (Murut), resak (Iban), sugoi (Murut)

Key to Dipterocarpus species

1. Calyx tube of mature fruit smooth (without ribs, tubercles or flanges)............................2

2. Stamens at most 15…………………………………………………………………...…3

3. Fruit calyx tube ellipsoid­globose, verrucose­lenticellate………..….39. D. verrucosus

Perpustakaan Negara Malaysia CataloguinginPublication Data

Distribution. Coastal hills of Myanmar, S and C Thailand, Cambodia and CochinChina,

Ecology. Evergreen and semievergreen lowland forests of IndoBurma; in mixed

1. Leaf blade sparsely puberulent or glabrescent, drying dull olivebrown…………………

3. Fruit calyx tube ellipsoidglobose, verrucoselenticellate………..….39. D. verrucosus

14. Leaf blade pale ochreouspubescent below………………………….……33. D. rigidus

19. Leaf blade ovatelanceolate, prominently corrugated, margin prominently revolute,

4. Leaf apex retuse to obtuse; venation densely tawny longtomentose below……….….…

16. Tomentum on twig and leaf bristlelike…………………………………………….….17

44. Leaf bud densely buff longtomentose………………………1. D. acutangulus (in part)