Malesian Orchid Journal Vol.

12 (2013): 39–54 39

Orchids of Extreme Serpentinite (Ultramafic) Habitats

in Kinabalu Park

Antony van der Ent1 and Jeffrey J. Wood2 Introduction

1
Centre for Mined Land Rehabilitation, Ultramafic outcrops are composed of mafic minerals
Sustainable Minerals Institute, (high in magnesium, iron, nickel, chromium and
The University of Queensland, cobalt) (Proctor et al., 2000) and found in large tectonic
Brisbane, QLD 4072, Australia. masses, which obducted at continental margins (Searle
email: a.vanderent@uq.edu.au & Stevens, 1984). The predominant ultramafic rock-
type, peridotite, consists mostly of the magnesium-iron-
2
Honorary Research Associate, sillicates olivine and (ortho/clino) pyroxene. Ultramafic
Orchid Herbarium, rocks are widespread and extensive in the Malaysian
Royal Botanic Gardens, state of Sabah, covering an area of approximately 3500
Kew, Richmond, km2 (Proctor et al., 1988). Kinabalu Park has some of
Surrey, TW9 3AE, United Kingdom. the most species-rich vegetation over these substrates
email: jeff.j.wood28@gmail.com anywhere in the world (Beaman, 2005). Orchids
______________________ typical of peridotite substrate in Kinabalu Park include
Appendicula tembuyukenensis, a species of montane
SUMMARY. Kinabalu Park has some of the most forest; Bromheadia divaricata, typically found in
species-rich vegetation on ultramafic soils anywhere leached, acid, extremely nutrient poor ‘kerangas’
in the world with high levels of endemism among type vegetation; Calanthe otuhanica, a characteristic
orchids. Some of the most remarkable habitats are
those formed on serpentinite, an ultramafic soil type
with extreme chemical properties. The vegetation of
serpentinite soils in Sabah is generally characterised
by the trees Gymnostoma sumatranum or Ceuthostoma
terminale (Casuarinaceae), as well as the shrub
Scaevola micrantha (Goodeniaceae). A number of
orchids are known only or predominantly to occur in
serpentinite habitats, including well-known species
such as Apostasia wallichii, Crepidium metallicum,
Paphiopedilum dayanum, P. rothschildianum,
Paraphalaenopsis labukensis, Phalaenopsis maculata,
Porpax borneensis, and Renanthera bella. The
characteristic open stature of the forest on serpentinite
soils is likely the main factor contributing to the
richness in terrestrial and epiphytic orchids. However,
for terrestrial orchids, adaptations to the adverse
chemical conditions of serpentinite soils may be a Fig. 1. Serpentinite outcrops and active landslides in the
contributing factor for the high level of endemism in Penataran Valley, the habitat of a range of unusual orchids.
this habitat. Photo: A. van der Ent.
40 Antony van der Ent and Jeffrey J. Wood

Fig. 2 (left). Forest in the Penataran Valley dominated by Ceuthostoma terminale, and also Gymnostoma sumatranum. Fig. 3
(right). Vegetation on ultramafic substrate, Wuluh River, Mount Tambuyukon, Kinabalu Park. Note the typical greenish hue
of the rock and the presence of Casuarinaceae. Photos: A. van der Ent.

species of open landslides at elevations above 1500 m;
Coelogyne rupicola and Cymbidium elongatum, found
in high elevation scrub; Dilochia cantleyi, a terrestrial
found in kerangas-type vegetation on leached peridotite;
Hetaeria hylophiloides, a terrestrial of montane forest;
Thrixspermum triangulare, a characteristic terrestrial
or epiphytic species of high-elevation Leptospermum
scrub.
Peridotite can be hydrated and metamorphised to
serpentinite (composed of such minerals as antigorite,
chrysotile and lizardite) (Baillie et al., 2000), usually
at the sea floor along tectonic boundaries. During
serpentinization the dense and hard peridotite rock is
transformed to a typically ‘soap-like’ far less dense
matrix of hydrous magnesium iron silicate, which,
due to its unconsolidated nature, is extremely prone
to landslides. Local Dusun place names reflect the
tendency for landslides of serpentinite outcrops and
the peculiar blue/green colour of the rock, for example
the locality ‘Morou Porou’ which means ‘shiny rock’.
Here we focus solely on serpentinite soils, as these
are the most extreme in their chemical properties.
Serpentinite soils have extremely high concentrations
of exchangeable magnesium and exceptionally low
concentrations of exchangeable calcium (with Mg:Ca
ratios up to > 100). The soil pH is circum-neutral (pH
6–7) on weathered soils with build-up of humus, but
alkaline (pH 7.5–8.8) on unweathered serpentinite such
as on landslides. Other extreme chemical properties
of serpentinite soils include major deficiencies in the
essential macronutrients phosphorus (<25 mg/kg) and
potassium (<50 mg/kg) and potentially phytotoxic
nickel concentrations. Serpentinite soils are the least
frequent ultramafic bedrock type in Sabah, and their
occurrence is limited to serpentinite bodies, often
embedded in peridotite along ancient fault lines.
The serpentine habitat and soils
The largest serpentinite outcrops in Kinabalu Park are
the Penataran Valley on Mount Kinabalu’s west flank,
Wuluh Valley on the foot slopes of Mount Tambuyukon,
 Orchids of Extreme Serpentinite (Ultramafic) Habitats in Kinabalu Park 41

the Nalumad Basin, the Lohan Valley on the southern
slopes of Mount Kinabalu, and in the Hempuen Forest
Reserve (the last two outside Kinabalu Park). Other
minor occurrences are found in the Bambangan Valley
and the Kinaram Valley, on Mount Kinabalu’s southern
slopes and Mount Tambuyukon’s northern slopes,
respectively.
All of these habitats are narrow valleys with steep
slopes, where a river has cut through a serpentinite
body. Typically the habitat represents a multitude of
landslides in various stages of succession to closed
forest. The vegetation of serpentinite outcrops
is commonly characterised by the dominance of
Gymnostoma sumatranum at lower altitudes (200–
1400 m) and Ceuthostoma terminale at higher
elevations (900–1800 m); both belong to the family
Casuarinaceae. The dominance of these two tree
species on serpentinite soils might be explained by
the very low nitrogen concentrations in these soils,
which the Casuarinaceae may successfully cope with
because of their nitrogen-fixing nodules (Dommergues
et al., 1990). Interestingly, Gymnostoma sumatranum
has relatively high foliar potassium concentrations,
whereas the soil has exceptionally low potassium
supply (Table 1).
Another factor likely to be important in serpentinite
outcrops is water stress. The small crowns and distinct
morphological adaptations to reduce transpiration,

Table 1. Comparison of typical soil analysis of two types of lowland (600–700 m) ultramafic soils, derived from
peridotite (deep laterite) and serpentinite (shallow bare soil) from Kinabalu Park (mean values, n = 2 and n = 3).

Parameters Peridotite (laterite soil) Serpentinite (bare soil)

1
pH 5.5 7.0
2
Cation Exchange meq/100g 0.8 6
3
Mg: Ca 14 89
4
Calcium % 0.005 0.1

Magnesium % 0.2 19.4

Cobalt mg/kg 3 104

Chromium mg/kg 10107 953

Iron % 33.4 4.7

Potassium mg/kg 53 47
5
Potassium mg/kg < 0.01 < 0.01

Nickel mg/kg 2510 1132

6
Nickel mg/kg 0.5 18

Phosphorus mg/kg 281 24

7
Phosphorus mg/kg 1.4 1.5

Carbon/Nitrogen ratio 18 85

NOTES: 1 pH in water-extract (1:2.5), 2+3 CEC and component ions in silver-thiorea extract, 4 Soil digestion
with nitric acid/hydrochloric acid digest, 5 Potassium extracted in bicarbonate extract, 6 DTPA-extractant for
extractable nickel. 7 Phosphorus extracted following Olsen.
42 Antony van der Ent and Jeffrey J. Wood

Fig. 4. The Penataran River cuts through a substantial body of serpentinite. The water has a characteristic creamy colour
resulting from suspended serpentinite particles. Photo: A. van der Ent.
 Orchids of Extreme Serpentinite (Ultramafic) Habitats in Kinabalu Park
most notably in Casuarinaceae, seem to point to water
stress, despite the high rainfall regime in Kinabalu
Park. The very low potassium supply in ultramafic soils
can exacerbate the effects of water stress (Pandolfini
et al., 1992). Serpentinite soils are typically alkaline
with pH >7 and a cation exchange complex dominated
by magnesium (Table 1). The total magnesium
concentrations in serpentinite soils are extremely
high (compare values in Table 1 between peridotite
and serpentinite ultramafic soils). When compared to
peridotite soils iron and chromium concentrations are
lower, but exchangeable nickel and total cobalt are
high in serpentinite soils.
The xeric conditions of the open vegetation on
serpentinite landslides are likely to be a major factor
for both terrestrial and epiphytic orchids. Many of
the orchids occurring here have thick pseudobulbs
indicating water conservation mechanisms. But
terrestrial orchids also need to be adapted to the adverse
soil conditions of serpentinite soils, to benefit from high
Fig. 5. Hill ridges above the Penataran River. Note the grey hue of Scaevola micrantha bushes on landslides. Photo: A. van
der Ent.
43
solar exposure for photosynthesis. The establishment
of trees is hampered by the frequent re-occurrence of
landsides, and hence most of the vegetation is in a varied
mosaic of different stages of succession. Terrestrial
orchids such as Paphiopedilum rothschildianum seem
to have an optimum (e.g. a dominant aspect with large
flowering plants) in a succession stage in which small
trees have fixed soil movement and where some organic
matter has accumulated. However when the forest
develops further and the forest becomes closed, P.
rothschildianum populations locally decline, the plants
become smaller and do not flower anymore. Most
terrestrial orchids that occur in the wild on serpentinite
soils are easily grown on non-ultramafic soils (such as
P. rothschildianum). For others, such as Phalaenopsis
maculata, alkaline soil conditions are required and
some, such as Apostasia wallichii, are extremely
difficult in culture. Reciprocal transplantation of
epiphytes typical of this habitat, such as Renanthera
bella, in culture or on other trees in a different habitat
44 Antony van der Ent and Jeffrey J. Wood

does not result in growth impediment, if high light by large Ceuthostoma terminale and/or Gymnostoma
conditions are maintained. sumatranum (Casuarinaceae) and the extremely rare
tree Borneodendron aenigmaticum (Euphorbiaceae)
Endemism and restriction to serpentinite soils are in a local climax. The pioneer vegetation of
the landslides comprises bushes of Decaspermum
Plants occurring on serpentinite (and other types of vitis-idaea (Myrtaceae), Macaranga kinabaluensis
ultramafic) soils may be grouped into: (a) substrate- (Euphorbiaceae), Scaevola micrantha (Goodeniaceae),
indifferent species that occur on ultramafic and non- and small trees of Ceuthostoma terminale and
ultramafic soils called ‘facultative ultramafic species’; Gymnostoma sumatranum (Casuarinaceae). The
(b) species that reach ultramafic sites as an extension carnivorous pitcher plants Nepenthes macrovulgaris
of their normal range (‘outliers’); and (c) endemics and N. reinwardtiana (Nepenthaceae) are often locally
to these soils (viz. Kruckeberg, 1986; 1991). The common. Paphiopedilum rothschildianum occurs as a
importance of ultramafic habitats for plant diversity terrestrial on particularly steep ledges, usually under
and supporting high levels of endemics on Mount patchy Gymnostoma trees. Paphiopedilum dayanum
Kinabalu and elsewhere in Sabah has been discussed occurs higher up the ridges, growing in the shade of
in detail by Beaman and Beaman (1990), Beaman tall Casuarinaceae-dominated forest, particularly
(2005) and Wong (1998). Wood et al. (2011) list 866 under dense growth of scrambling bamboo belonging
taxa of orchids in 134 genera with 90 species being to the genus Racemobambos. The endemic Porpax
endemic to Mount Kinabalu. Of those, 359 orchid borneensis can often be found growing approximately
species are found exclusively (‘obligate’) or partly 30–70 cm up the base of Gymnostoma trees. Typical
(‘facultative’) on ultramafics; 270 taxa occurring for Casuarinaceae-dominated forest on serpentinite
facultative on ultramafics are epiphytic. Some of the is the abundance of ant-plants such as Hydnophytum
most distinct species assemblages, including a range formicarum (Rubiaceae), Myrmecodia tuberosa
of spectacular orchid species, occur on serpentinite (Rubiaceae) and Dischidia major (Apocynaceae).
soils in Kinabalu Park. These include well-known The symbiotic relationship between Acriopsis spp.
species such as Crepidium metallicum, Paphiopedilum and ants, specimens often perched on the top of
dayanum, P. rothschildianum, Paraphalaenopsis Hydnophytum formicarum, is common locally. Under
labukensis, Phalaenopsis maculata, Porpax the Gymnostoma trees the soil is covered with leaf litter
borneensis and Renanthera bella. Although obligate composed of their fallen ‘needles’. The terrestrials
ultramafic species are comparatively rare, a greater Crepidium lowii, C. metallicum and Plocoglottis lowii
number of these endemics are found on serpentinite are commonly found growing between the thick packs
soils in Kinabalu Park. This might be explained by the of needles.
inherent slow growth rates of most of these species,
leading to competitive exclusion outside edaphically Some orchids typical of serpentinite soils in Kinabalu
limited habitats. It is important to note that there are Park
always exceptions with a few specimens of a particular
species occurring on a different ultramafic soil or on Particularly noteworthy orchid taxa found on
non-ultramafic soils. serpentinite soils within Kinabalu Park include
Apostasia wallichii (Fig. 6), Arachnis longisepala
The vegetation of serpentinite valleys in Kinabalu (Fig. 7), Crepidium metallicum (Fig. 9), C. lowii,
Park Neuwiedia zollingeri var. javanica, Plocoglottis lowii
(Figs. 16 & 17), Paphiopedilum dayanum (Figs. 11
Most serpentinite outcrops are in deeply incised gorges & 12), P. rothschildianum var. rothschildianum (Figs.
and narrow valleys ranging from about 900 m asl at the 13 & 14), Paraphalaenopsis labukensis (Fig. 15),
top of the ridges, to about 500–700 m asl along the river Phalaenopsis maculata, Porpax borneensis (Fig. 18),
bed. Characteristic are the large number of landslides Renanthera bella (Fig. 19), and Spathoglottis gracilis.
exposing bare serpentinite rock. Most of the vegetation The rat-tail orchid Paraphalaenopsis labukensis
has been affected by landslides and is dominated typically grows either high up on trees along rivers,
by successional vegetation in various stages of or in lower trees along ridges. This habit is similar to
development. Only the forests on the ridges, dominated that of Grammatophyllum kinabaluense (Fig. 10). The
 Orchids of Extreme Serpentinite (Ultramafic) Habitats in Kinabalu Park 45

aptly named Corybas serpentinus (Fig. 8) often grows after Dr Nathaniel Wallich (1786–1854), a Director of
lithophytically on moist serpentinite boulders. Calcutta Botanic Garden, whose large herbarium is
housed at Kew.
Apostasia wallichii R. Br. Fig. 6.
Arachnis longisepala (J.J. Wood) Shim & A. Lamb
Apostasia (together with Neuwiedia) belongs to the Fig. 7.
subfamily Apostasioideae, a sister group to the rest
of the Orchidaceae. The apostasioids are of particular This scorpion orchid was originally described as
interest, because they may represent an evolutionary a subspecies of A. calcarata Holttum, which also
link between the orchids and lily-like ancestors. occurs on Mount Kinabalu. Both species have a
Unlike the majority of orchids, which have only one distinctive long dorsal sepal which (together with
fertile stamen, Apostasia has two. Nodular storage A. grandisepala) is unusual in the genus. Arachnis
organs and stilt roots at the base of the stem are also longisepala is distinguished from A. calcarata by the
found in Apostasia. The unique features by which longer inflorescences, which often attain 120 cm and
the apostasioids differ from other orchids make them bear 8 to 16 smaller but rather more elegant flowers.
a clearly derivative group. The presence of two or These have pale lemon-yellow lateral sepals and petals
three (in Neuwiedia) anthers indicates a degree of irregularly blotched with shiny maroon-purple above.
primitiveness, but they are not otherwise what we The narrow strap-like dorsal sepal, which may measure
would imagine as the common ancestor of the orchids. up to 7.4 cm long, is entirely shiny maroon-purple
Apostasia wallichii is a very widespread species above. The lip has a bucket-shaped hypochile, the lobes
ranging from India and Sri Lanka in the west to New of which clasp the column, and a narrowly triangular-
Guinea and northern Australia in the east. It is named ovate epichile with an upcurved aristate apex.

Fig. 6. The primitive terrestrial Apostasia wallichii growing on blue-grey serpentinite soils in the Wuluh Valley, Kinabalu
Park. It is often abundant under Gymnostoma, particularly where the soil is rather bare. Photo: A. van der Ent.
46 Antony van der Ent and Jeffrey J. Wood

Fig. 7. Arachnis longisepala, a scorpion orchid from hill forest on ultramafic substrate where it often grows on tall trees,
sometimes 30 metres up in the canopy. Photo: R. Nilus.

Corybas serpentinus J. Dransf. Fig. 8.

A distinctive helmet orchid closely allied to the

familiar C. pictus but differing in the colouration of the
lip and the two low swellings on the lip at the mouth
of the throat, which is thus shallowly V-shaped rather
than flat or concave. It was described in 1986 from
material collected by John and Soejatmi Dransfield
on the ultramafic Mount Silam in Lahad Datu District,
Sabah. At the type locality it was found growing on
thin soil and moss carpets on a ridge top at an elevation
of about 750 m. This orchid has not yet been recorded
from Kinabalu Park, but has been recently found in the
Bidu-Bidu Hills in Central Sabah where it grows in
lowland serpentinite soils. It seems likely that it may
occur somewhere within Kinabalu Park.

Crepidium metallicum (Rchb. f.) Szlach. Fig. 9.

Formerly placed in the ‘bucket’ genus Malaxis, C.

metallicum has particularly striking glossy amethyst-
purple foliage. Reichenbach, writing in the Gardeners’
Chronicle of 1879, described it as “quite a gem!” It was
originally imported into Britain from Borneo by a Mr
Bull in the 1870s and later figured in Curtis’s Botanical
Fig. 8. Corybas serpentinus from the Bidu-Bidu Hills
commonly grows between serpentinite boulders where it
colonises pockets of moist organic soil. Photo: A. van der
Ent.
 Orchids of Extreme Serpentinite (Ultramafic) Habitats in Kinabalu Park 47

Magazine as plate 6668 published in 1883. The leaves

were described by Reichenbach as being “of a light
rose colour underneath, and blackish purple above,
with quite an exquisite metallic lustre, and when dried
after having been boiled in hot water they are green”.

Grammatophyllum kinabaluense Ames & C.

Schweinf. Fig. 10.

The genus Grammatophyllum includes what is generally

acknowledged to be the largest orchid in the world, viz.
G. speciosum, the Tiger Orchid (Lamb, 2011). Two
distinct growth-forms are found in Grammatophyllum.
One has very long pseudobulbs, which are in reality
fleshy stems bearing numerous leaves. Examples
include G. kinabaluense and the well-known G.
speciosum. The other has rather short, proportionally
thick pseudobulbs, which are not covered by leaf-bases,
with only a few leaves at the apex. Examples include
G. scriptum and G. stapeliiflorum. Both growth-forms
produce erect, branching catch-roots for trapping leaf-
litter. Floral morphology in both is essentially the same.

Fig. 9 (above). Crepidium metallicum ascends to over 2000 m on Mount Kinabalu and once occurred on the formerly
floristically rich, fire-ravaged Hempuen Hill. The leaves are a beautiful amethyst-purple. Fig. 10 (below). Grammatophyllum
kinabaluense forms huge clumps in tree tops, often above rivers, such as Lohan River. Photos: A. van der Ent.
48 Antony van der Ent and Jeffrey J. Wood

Fig. 11 (left). Paphiopedilum dayanum, another well-known slipper orchid from Kinabalu Park which grows on ridges
at elevations a little above those of P. rothschildianum. Fig. 12 (right). Paphiopedilum dayanum. Close-up of the flower.
Photos: A. van der Ent.

Fig. 13 (left). Paphiopedilum rothschildianum var. rothschildianum is a well-known species restricted to serpentinite
landslides in Kinabalu Park. This photo, taken at the type locality, shows its favoured habitat on very steep ledges above
rivers. Fig. 14 (right). Paphiopedilum rothschildianum var. rothschildianum. Close-up of the flower. Photos: A. van der Ent.
 Orchids of Extreme Serpentinite (Ultramafic) Habitats in Kinabalu Park 49

Grammatophyllum kinabaluense was described known of Sabah’s orchids. It was first introduced into
in 1920 from material collected at Kiau by US Army cultivation by the Belgian grower Jean Linden in May
chaplain Reverend Joseph Clemens. Populations are 1887 and, early the following year, by the English
known only from lowland and hill forest at elevations nurserymen Frederick Sander & Sons of St. Albans.
between 500 and 900 m. It has also been recorded Reichenbach named it in honour of the Victorian
from the Long Pa Sia area in Sipitang District. The orchid grower and aristocrat, Baron Ferdinand de
inflorescences are held erect and may attain three to Rothschild. The closest allies of P. rothschildianum
four metres in length, each bearing many flowers are P. adductum from the Philippines, P. glanduliferum
measuring between 6.5 and 7 cm across. from New Guinea, and P. supardii from Kalimantan.
Paphiopedilum rothschildianum differs from these
in having longer petals held at an acute angle to the
Paphiopedilum dayanum (Lindl.) Stein Figs. 11 &
horizontal so that the flower often has a span of 24 to 34
12.
cm. These graceful petals are held out like the arms of
the local Sumazau dancers. The distinctive staminode
A slipper orchid readily distinguished by its large
is narrow and has a knee-like bend at the base, a bifid
flowers characterised by a narrow but long dorsal
tip and hairy base and sides.
sepal, oblanceolate unspotted petals with long ciliate
Atwood (1985) demonstrated the significance
fringes, and transversely reniform staminode with
of the strange staminode in its pollination by syrphid
obscure apical teeth. The leaves are among the most
flies. He suggested that the glandular hairs on the
boldly tessellated in the genus and can have either a
staminode mimic an aphid colony, the normal brood
bluish-green (glaucous) or yellow-green hue, the latter
site of the syrphid larvae. The staminode attracts
corresponding to Reichenbach’s Cypripedium petri,
females of the fly Dideopsis aegrota to deposit their
which is conspecific.
eggs on its surface. On alighting upon the staminode
The specific epithet celebrates the eminent
to lay their eggs, the flies sometimes fall into the lip.
Victorian orchid grower and orchid painter John Day
Their only exit is through a gap between the base of
(1824–1888) of Tottenham in north London. Day
the lip and the column, and the flies pass beneath the
sent flowers of many undescribed species flowering
stigma and pollinia, picking up a pollinium. A visit to
in his collection to Professor H.G. Reichenbach in
a second flower followed by a similar scenario with
Vienna, who reciprocated by naming several in his
the pollinium rubbed off onto the stigma will thereby
honour. Paphiopedilum dayanum is one of the most
effect pollination.
distinguished of these and Day flowered it in 1860
Plants fitting the description of P. rothschildianum
from an importation by the nursery of Messrs. Low &
var. elliottianum (O’Brien) Pfitzer, differing only in
Co. of Clapton, London. It was, in fact, Sir Hugh Low
having hanging rather than spreading petals, have
who had discovered it on Mount Kinabalu in 1856.
recently been found elsewhere in Sabah. These were
Paphiopedilum dayanum occurs on the lower
found on similar ultramafic soils to those at the type
slopes of Mount Kinabalu where it grows in leaf litter
locality of var. rothschildianum in Kinabalu Park.
under bamboo and at the base of trees on steep ridges
that flank the mountain. Plants with glaucous and
yellow-green leaves grow here intermingled. The type
Paraphalaenopsis labukensis Shim, A. Lamb & C.L.
locality of C. x burbidgei (a natural hybrid between P.
Chan Fig. 15.
dayanum and P. javanicum var. virens) and C. petri is
now almost stripped of plants. Fewer than 20 scattered
The most spectacular pendant epiphytic orchid in
individuals now occur over an area of a few hundred
Kinabalu Park is undoubtedly Paraphalaenopsis
square metres.
labukensis. Plants grow suspended from small trees
three to five metres above the ground or twelve to twenty
Paphiopedilum rothschildianum (Rchb. f.) Stein var. metres up on the trunks and branches of larger trees.
rothschildianum Figs. 13 & 14. The short stem produces three to five extraordinary
fleshy, terete, whip-like, ‘rat’s tail’ leaves, which may
The flamboyant Rothschild’s Slipper Orchid is one of measure as much as two metres in length. These are
the most striking, horticulturally desirable and best 6–9 mm thick and slightly constricted 1.3–2.5 cm from
50 Antony van der Ent and Jeffrey J. Wood

the acute apex. The inflorescence normally bears five

or six beautiful cinnamon-scented flowers, although
these may number up to about fourteen in robust
specimens. Each flower measures up to 6.2 cm in
width. Paraphalaenopsis labukensis was a relatively
common epiphyte on Hempuen Hill (Bukit Hempuen)
prior to the area’s devastation by fire in 1990.

Fig. 15. Paraphalaenopsis labukensis is a spectacular

pendant epiphyte with terete, whip-like ‘rat-tail’ leaves.
It occurs on hill ridges where it is found either suspended
from small trees between about 3–5 m above the ground or
between about 12–20 m up on the trunks and branches of
larger trees. Photo: A. van der Ent.

Plocoglottis lowii Rchb. f. Figs. 16 & 17.

The probable pollination mechanism of Plocoglottis

lowii (as P. porphyrophylla, which is conspecific) was
discussed in detail by Burkill (1913). The base of the
lip acts as a strong spring, against the tension of which
the lip is held back by the right-hand lateral sepal. The
Fig. 16 (above). Plocoglottis lowii in the Penataran Valley.
This terrestrial often grows at the foot of steep slopes where
organic matter accumulates. Fig. 17 (below). Plocoglottis
lowii. Close-up of the flower. Photos: R. van Vugt.
 Orchids of Extreme Serpentinite (Ultramafic) Habitats in Kinabalu Park 51

lip is pushed into this position as the flower opens by
the expanding petals. When the lip is in position, the
right-hand lateral sepal moves across a little and holds
it, after which the petals expand fully. When an insect
touches the right-hand lateral sepal, the lip is released
and springs sharply up into contact with the face of the
column. The insect is thereby brought into contact with
the column and the pollinia are removed as it struggles
free. Holttum (1964) noted that no insect has actually
been observed to do this.
Porpax borneensis J.J. Wood & A. Lamb Fig. 18.
Seidenfaden (1977) recognised ten species of Porpax,
all confined to the Asian mainland, east as far as
Thailand and south to Peninsular Malaysia. Porpax
borneensis, which was described in 1993, extends the
range of the genus eastward.
The flowers of P. borneensis are dark cardinal-red
with paler red or yellow tips to the sepals. It may be
distinguished from the allied P. ustulata from Myanmar
and Thailand by the minutely papillose sepals and
entire, irregularly serrulate, minutely papillose lip.
This has several papillae of varying sizes toward the
base of the disc. The one- or two-leaved pseudobulbs
are ovoid, bilobed and distinctly flattened. The plant is
often found growing adpressed to the surface of rocks
or on the boles of Gymnostoma sumatranum.
Renanthera bella J.J. Wood Fig. 19.
One of the most striking species of a horticulturally
desirable genus, R. bella was described from Hempuen
Hill in 1981. It has a compact habit, compared to many
renantheras, the entire plant measuring (including
the inflorescence) only up to about 75 cm in length.
Unfortunately, as might be expected, it has been widely
over-collected and is now both rare and endangered
in the wild. To protect it further it has been placed on
CITES, Appendix 1, for which no trade is allowed
except for nursery-raised flasked seedlings. Because of
its compact habit, and attractive, long-lasting medium-
sized flowers, it has proved a valuable addition for the
commercial plant breeder. Several hybrids have now
been produced, but seedling growth has been found to
be very slow. The flowers, if left unpollinated, remain
fresh for up to three months.
The type locality of R. bella, Hempuen Hill,
located above the Lohan River on the southeastern
base of Mount Kinabalu, was formerly home to a
fascinating flora, including many interesting orchids.
This low-elevation ultramafic area has been the source
of many recently described species, both epiphytic and
terrestrial. It is, therefore, all the more unfortunate that
the hill was subsequently degazetted from Kinabalu
Park and suffered a catastrophic burning in 1990,
destroying all but a small area of forest around the
summit. Fortunately, R. bella has subsequently been
discovered elsewhere in Sabah.

Conclusions

The relationship between site ecology and the

distribution of plant species is a fundamental one, and
the soils on which plants or their epiphytic hosts grow
is an important factor. However, information about
tropical ultramafic soils in the region is extremely
scant. In the Mount Tambuyukon area, even the
geology is very poorly understood. Moreover, the
Fig. 18. Porpax borneensis grows on the boles of
Gymnostoma sumatranum or on the surface of serpentinite majority of herbarium specimens of orchids have little
rocks. This plant was photographed in forest along the information about the precise location and the nature of
Penataran River. Photo: A. van der Ent. the substrate on which they were found. Most tropical
52 Antony van der Ent and Jeffrey J. Wood

Fig. 19. Renanthera bella from Hempuen Hill (Bukit Hempuen). This spectacular epiphytic orchid is characteristic of open
forest on serpentinite. It is very scarce and known only from two localities in Sabah. Photo: A. van der Ent.
 Orchids of Extreme Serpentinite (Ultramafic) Habitats in Kinabalu Park
botanists are also unaware of the important differences
in ultramafic rocks and the soils derived from them,
based on the assumption that ultramafic soils differ
little, because of their shared geological origin.
Confusing in this perspective is also the terminology
itself, with ‘serpentine’ often used as a synonym for
‘ultramafic’, whereas ‘serpentinite’ specifically denotes
the mineral group derived from serpentinization of
peridotite. The chemical differences (and hence their
effect on plant growth) between soils derived from
peridotite or those derived from serpentinite, are
significant, with serpentinite soils having the most
extreme properties (and potentially limiting to plant
growth). The characteristic open stature of the forest on
serpentinite soils is likely the main factor contributing
to the richness in terrestrial and epiphytic orchids.
However, for terrestrial orchids, adaptations to the
adverse chemical conditions of serpentinite soils may
be a contributing factor for the high level of endemism
to the serpentinite habitat.
Although less than 1% of the earth’s land surface is
ultramafic (Proctor, 1999), such outcrops are renowned
as centres for plant diversity and support high levels
of plant endemism (Rajakaruna & Baker, 2004). Many
plant species only occur in a single or a few ultramafic
sites, and destruction of the ecosystem that supports
them may therefore result in their extinction. At
Kinabalu Park, for example, the distribution of many
plant species is so localized (especially on ultramafic
soils), that 40% of all species recorded for the area
are only known from a single collection (Beaman &
Beaman, 1990). Although the important localities in
Kinabalu Park are safeguarded because of its statutory
protection and UNESCO World Heritage Site status,
it has become a ‘virtual island’ in a ‘sea’ of cleared
land, which means that gene flows to other (former)
populations of plants occurring outside the Park have
become cut-off.
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Acknowledgements
We would like to express our gratitude to Rimi Repin,
Rositti Karim, Sukaibin Sumail, Handry Mujih, Dolois
Sumbin, Kinahim Sampang, Yabainus Juhalin and
Alim Biun (Sabah Parks), and John Sugau and Postar
Miun (Sabah Forestry Department) for their help and
expertise in the field and in the herbarium. Thanks also
to Rogier van Vugt and Simon Wellinga for assistance
in the field in March 2012 and for permission to use
two photos featured in this article. Finally, we would
like to gratefully acknowledge the continuous support
of Sabah Parks and the Sabah Forestry Department.