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Malesian Orchid Journal Vol.

10 (2012): 103–122 103

Mount Tambuyukon
— An Intriguing Mountain and its Orchids

Jeffrey J. Wood1 and Antony van der Ent2 mountain is somewhat of a mystery. Most local people
in the kampungs of Monggis and Serinsim call it
1 Orchid Herbarium, ‘Madalon’ (which means ‘long hilly ridge’) and say
that the name ‘Tambuyukon’ was made up by
Royal Botanic Gardens,
surveyors in the 1960s. On the official contour map it
Kew, Richmond,
is spelled ‘Tambuyukon’. The massif is roughly 16 km
Surrey TW9 3AE, United Kingdom.
long and 6–8 km wide and is situated about 12 km
email: Jeff.Wood@kew.org
northeast of Mount Kinabalu, lying entirely within the
2 Centre for Mined Land Rehabilitation, Kinabalu Park World Heritage Site. The mountain is
barely visible from Kampung Monggis and is much
Sustainable Minerals Institute, better seen from Serinsim in the north, where it flanks
The University of Queensland, Mount Nambuyukon. Because of its remote location
Brisbane, QLD 4072, and its position inside the protected area, Mount
Australia. Tambuyukon has intact vegetation including tall
email: a.vanderent@uq.edu.au lowland rainforest around sea level to short grassy
______________________ vegetation along the summit ridge. Mount
Nambuyukon lies at the foot of Mount Tambuyukon,
SUMMARY. Mount Tambuyukon, an ultramafic and is partly ultramafic, partly chert/spillite, a type of
mountain inside Kinabalu Park in Sabah, is described rock associated with ultramafic outcrops.
and some of its orchids discussed and illustrated. Mount Tambuyukon is part of a large ophiolite
Recent surveys indicate that Mount Tambuyukon has a suite, with ultramafic (sometimes also called
very high per area species diversity rivalling that of ‘ultrabasic’ or ‘serpentine’) outcrops. Such rocks
Mount Kinabalu. A preliminary checklist of the (predominantly peridotite) are rich in iron and
orchids recorded on the mountain is provided; magnesium, and also contain high levels of normally
however much fieldwork and detailed studies are
needed before the full orchid diversity of this mountain
is known.

Mount Tambuyukon — a little known mountain

Although relatively unknown, Mount


Tambuyukon (‘Tembuyuken’ and other spellings) at
2579 m is the third highest mountain in Malaysia, and
on the island of Borneo, after Mount Kinabalu and
Mount Trus Madi (Briggs, 1988). It is also the highest
ultramafic mountain in Southeast Asia. The first
recorded ascent of Mount Tambuyukon was made in
the 1960s by the late Dr Willem Meijer of the Sabah Fig. 1. Summit slopes of Mount Tambuyukon, with Mount
Forestry Department. The Dusun name of the Kinabalu in the background. Photo: A. van der Ent.
104 Jeffrey J. Wood and Antony van der Ent

plant-toxic nickel and chromium, but are usually poor defined new trail leads into a thicket behind the
in calcium and other macronutrients such as potassium building, and soon follows an indistinct ridge that
and phosphorus (Baillie et al., 2000; Rajakaruna & gradually rises. After several hundred metres the ridge
Baker, 2006). Universally ultramafic outcrops have becomes more prominent, and as it rises the sides get
become well known for the distinctive vegetation that steeper. The old growth forest here has spectacular
they support (Brooks, 1987; Harrison & Rajakaruna, examples of dipterocarps on top of the well-drained
2011). ridges. After a prolonged climb the path levels and
The mountain ranges in elevation from 280 m at eventually descends into a swampy area. Over several
the foot to 2579 m at the summit. The tall (up to 55 m) kilometres the path continues to follow the obvious
forest at the foot of the mountain decreases in stature ridges on sandstone and shale rock, now rising, then
to a unique grassy (graminioid) vegetation less than descending. The trail crosses the Wulluh and
one metre high on the summit ridge. The route to Kepuakan rivers, and then climbs steeply to the side of
Mount Tambuyukon starts in lowland forest at an a long ridge that stands perpendicular to the main
elevation of 367 m at Monggis substation. A well- summit ridge of Mount Tambuyukon. Once this side
Mount Tambuyukon — An Intriguing Mountain and its Orchids 105

ridge is reached, it is a steep ascent the whole way, up Tambuyukon is basically the highest point of several
to Musang Camp. After Musang Camp, the last push to inter-connected ridges, with another prominent dome
the actual summit ridge is a very steep climb through of almost identical height to the west. The summit
spectacular upper montane ‘mossy’ forest; though only flora of Mount Tambuyukon is very fragile and easily
4 km, it rises over 1000 m. An expedition to the damaged and for this reason Sabah Parks restricts
summit ridge of Mount Tambuyukon is exceptionally access to the mountain.
tough and should only be undertaken by those in very The graminioid vegetation on the exposed summit
fit condition. Traversing the summit itself is not ridges of Mount Tambuyukon is dominated by sedges
particularly physically demanding but, continuing a (particularly Schoenus spp.) and a range of small
little further down on the south side, one reaches a very shrubs such as Leptospermum recurvum (Myrtaceae),
exposed ridge and a small plateau without any trees but Lithocarpus rigidus (Fagaceae), Scaevola verticillata
with thickets of, among others, the carnivorous pitcher (Goodeniaceae), Tristaniopsis elliptica (Myrtaceae),
plant Nepenthes rajah. From here the view towards and Dacrydium gibbsiae (Podocarpaceae). Enigmatic
Mount Kinabalu is spectacular. The summit of Mount species of the summit scrub include Begonia

Fig. 2. A panoramic view of Mount Tambuyukon, showing the


main summit ridge on the left. Photo: A. van der Ent.
106 Jeffrey J. Wood and Antony van der Ent
Mount Tambuyukon — An Intriguing Mountain and its Orchids 107

Opposite: Fig. 3 (above left). Close-up of graminioid vegetation; the pink-flowered plant in the middle distance is Melastoma
kinabaluensis (Melastomataceae). Fig. 4 (above right). Upper slopes of Mount Tambuyukon. Fig. 5 (centre left). Forest in the Wuluh
River valley at the foot of Mount Tambuyukon with the pale green crowns of Gymnostoma sumatranum. Fig. 6 (centre right). Forest
with Gymnostoma sumatranum developed over serpentinite substrate showing characteristic reddish brown Gymnostoma litter in
the foreground. Fig. 7 (below). Dense upper montane scrub developed over ultramafic substrate on Mount Tambuyukon including
Dacrydium gibbsiae, Leptospermum recurvum, Schima wallichii, and Weinmannia clemensiae. Photos: A. van der Ent.
Above: Fig. 8 (left). Detail of upper montane scrub on Mount Tambuyukon showing Dacrydium gibbsiae, Leptospermum javanicum
and Schima wallichii. Fig. 9 (right). Summit area of Mount Tambuyukon, with graminioid vegetation in the foreground. Photos: A.
van der Ent. Fig. 10 (below). An open hillside on the summit of Mount Tambuyukon rich in grasses and sedges among which are
found Nepenthes rajah, and orchids such as Appendicula tembuyukenensis and Cymbidium elongatum. Photo: A. Lamb.
108 Jeffrey J. Wood and Antony van der Ent

vaccinioides (Begoniaceae), Rhododendron baconii on Leptospermum trees. Other epiphytes include


and R. meijeri (Ericaceae), which are all endemic to Appendicula tembuyukenensis and Cleisocentron
Mount Tambuyukon alone. A number of terrestrial merrillianum.
orchids are common among grasses and sedges in the Ultramafic substrates are significant in the
graminioid shrub including Calanthe otuhanica, occurrence of many orchids of Kinabalu Park; 359 taxa
Coelogyne rupicola, Cymbidium elongatum, have been found on Mount Kinabalu entirely or partly
Platanthera kinabaluensis, and Thrixspermum on ultramafic outcrops; 270 taxa known to occur at
triangulare. The pitcher plant Nepenthes rajah least in part on ultramafic substrates are epiphytic
(Nepenthaceae) is dominant in places. On less exposed (Wood et al., 2011). Possibly the types of vegetation in
slopes, a dense scrub has developed that includes all of ultramafic areas are particularly satisfactory for
the species from the open graminioid areas, as well as epiphytic species because of a light open canopy that
a range of other small trees, such as Wendlandia may encourage the diversity of epiphytes, but the low,
tambuyukonensis (Rubiaceae), Vernonia arborea var. scrubby vegetation on the ultramafics may also allow
mollissima (Asteraceae), Weinmannia clemensiae collectors to readily find the epiphytes in those
(Cunoniaceae), and Xanthomyrtus flavida habitats.
(Myrtaceae). The shorter forests merge into medium At elevations above 1300 m on the Kinabalu
tall upper montane forest further down dominated by massif ultramafic (or facultative ultramafic) species
Agathis kinabaluensis (Araucariaceae), Callicarpa substantially outnumber the non-ultramafic species. In
kinabaluensis (Lamiaceae), Dacrydium gibbsiae the case of endemics, ultramafic species substantially
(Podocarpaceae), Leptospermum javanicum outnumber non-ultramafic species at all except the
(Myrtaceae), Magnolia persuaveolens subsp. rigida lowest elevations (Fig. 11).
(Magnoliaceae), and Phyllocladus hypophyllus Particularly noteworthy among orchid species
(Phyllocladaceae). Very abundant is the tall terrestrial generally found on ultramafic substrates within
orchid Dilochia cantleyi, and a species of Kinabalu Park are Appendicula congesta, A.
Thrixspermum allied to T. kocyanii, which is epiphytic tembuyukenensis (Fig. 15), Arachnis longisepala,

Fig. 11. Elevational distribution of endemic orchid taxa on ultramafic and non-ultramafic substrates on Mount Kinabalu.
Mount Tambuyukon — An Intriguing Mountain and its Orchids 109

Fig. 12. Summit vegetation on Mount Tambuyukon. Photo: A. van der Ent.

Ascidieria cymbidifolia var. pandanifolia, Bromheadia and Mount Kinabalu are both part of an elevated
divaricata, Bulbophyllum nubinatum, Ceratostylis region which started to form in the Early Miocene and
ampullacea, Chamaeanthus brachystachys, probably increased in extent since the Middle Miocene
Chroniochilus minimus, C. virescens, Coelogyne (19 Ma onwards) (Hall, pers. comm.). Much of eastern
rupicola (Figs. 19 & 20), Crepidium metallicum, Sabah was at or close to sea level until the end of the
Cymbidium elongatum (Fig. 21), Dendrobium piranha Miocene and it is likely that Mount Kinabalu and
(Fig. 24), D. patentilobum, Dendrochilum Mount Tambuyukon became higher in this period
kamborangense (Fig. 26), D. lancilabium, Epigeneium (Hall, pers. comm.). Indications for a historical greater
kinabaluense, E. longirepens, Liparis height of Mount Tambuyukon is found in very
kamborangensis, Mycaranthes major, Neuwiedia extensive subaerial debris flows round the base
borneensis, N. zollingeri var. javanica (Figs. 30 & 31), resulting from geologically recent erosion, but it is
Paphiopedilum rothschildianum var. rothschildianum, unknown how high Mount Tambuyukon has been or
Paraphalaenopsis labukensis, Phaius pauciflorus whether it has been continuously a significant
subsp. sabahensis, and Renanthera bella. mountain since its uplift. In New Guinea the treeline
was lowered by as much as 1500 m during the
New insights Pleistocene (Walker & Flenley, 1979) which means
that even considering the current height of Mount
The University of Queensland and Sabah Parks have Tambuyukon, the mountain would have had an alpine
gained some new insights on the plant diversity on flora.
Mount Tambuyukon from a recent collaborative Mount Tambuyukon is entirely ultramafic, from
research project. Placed in a geological context, these about 270 m asl to the summit ridge at 2579 m. There
findings might have important consequences for are, however, important local differences in the
interpreting the biogeography of the region. The geology and consequently in the vegetation. For
ultramafic rocks of Mount Tambuyukon probably instance, the intrusion of serpentinite bodies in the
represent part of an ophiolite of Middle Jurassic to peridotite, which is the dominant rock, has caused
Early Cretaceous age (100–176 Ma) (Hutchinson, some very peculiar vegetation types, e.g., at Wuluh
2005) and were tectonically emplaced in the Late River. These serpentinite outcrops at the foot of Mount
Cretaceous or Early Palaeogene (45,100 Ma) (Newton- Tambuyukon are characterised by the tree
Smith, 1967; Cottam et al., 2010). Mount Tambuyukon Gymnostoma sumatranum (Casuarinaceae) and the
110 Jeffrey J. Wood and Antony van der Ent

terrestrial orchids Apostasia wallichii and Crepidium


metallicum.
At the same altitudes (particularly between 1500
and 2000 m) same-sized plots are often richer on
Mount Tambuyukon than on Mount Kinabalu. For
example, some 250 m2 plots have been found to have
in excess of 100 vascular plant species (Van der Ent,
unpublished, 2011). Whether some of the (local)
endemics of Mount Kinabalu originally evolved on
Mount Tambuyukon, and migrated to Mount Kinabalu,
is uncertain at present, but Mount Tambuyukon has
undoubtedly played an important role in the
biogeography of Mount Kinabalu. Possible evolution
of local endemics on Mount Tambuyukon may have
been the result of the mountain’s biogeographical
history or relate to the ultramafic soils which present a
strong selective pressure for plants. As a result a
number of plant species are known exclusively from
Mount Tambuyukon, particularly at higher elevations,
including Scaevola verticillata (Goodeniaceae),
Rhododendron baconii and R. meijeri (Ericaceae) and
the herb Begonia vaccinioides (Begoniaceae). In
addition, with the results from new studies coming in, Fig. 13. Aeridostachya robusta. Mount Tambuyukon. Photo:
most plant species that were considered to be endemic A. van der Ent.
to Mount Kinabalu, or those plant species that were
only known in Sabah from Mount Kinabalu, have now Aeridostachya robusta is a widespread and
been found on Mount Tambuyukon as well. Examples variable species of hill and montane forest at
include Euphrasia borneensis (Scrophulariaceae) and elevations between 400 and 3000 m in Kinabalu Park.
Rhododendron lowii (Ericaceae). Some neo-endemics Three further species recorded from the park, viz., A.
certainly have evolved on Mount Kinabalu, e.g. taxa in bancana (J.J. Sm.) J.J. Wood, A. crassipes (Ridl.)
Dendrochilum section Eurybrachium (Barkman, Rauschert and A. macrophylla (Ames & C. Schweinf.)
2001), but with future studies on Mount Tambuyukon J.J. Wood, may possibly fall within the variation of A.
we have to consider its influence on the plant diversity robusta.
of Mount Kinabalu.
Apostasia wallichii R. Br. Fig. 14.
A selection of orchids from Mount Tambuyukon

Aeridostachya robusta (Blume) F.G. Brieger Fig. 13. Apostasia, together with Neuwiedia, form the
subfamily Apostasioideae, considered to be a sister
The genus Aeridostachya was formerly included at group of the remaining orchids. The apostasioids are of
sectional level within Eria, a large and problematic particular interest because they may represent an
genus of approximately 370 species which has been evolutionary link between the orchids and lily-like
the subject of recent molecular and morphological plants. Unlike the majority of orchids, which have only
phylogenetic studies by Ng (2002). Ng provided one fertile stamen, Apostasia has two and Neuwiedia
evidence suggesting that the large and rather unwieldy three. The pollen grains are powdery and granular and
Eria, in the widely accepted sense, is polyphyletic. never aggregated into pollinia, otherwise universal in
Eria sections Aeridostachya, Cylindrolobus and the Orchidaceae, except in some slipper orchids.
Dendrolirium form a clade that has been amalgamated This primitive orchid is typical of bare serpentinite
into a rather too broadly defined genus Callostylis by outcrops, such as near the Lohan River southeast of
Pridgeon et al. (2005). Wood et al. (2011) prefer to Mount Kinabalu and along the Wuluh River on the
recognise the former three sections as distinct genera. lower slopes of Mount Tambuyukon.
Mount Tambuyukon — An Intriguing Mountain and its Orchids 111

Calanthe otuhanica C.L. Chan & T.J. Barkman Figs.


16 & 17.

This Calanthe was described from Mount Kinabalu as


recently as 1997. Apart from C. woodii, no other
species of Calanthe occur at comparable elevations in
Sabah. Calanthe otuhanica has strongly sweet-
scented, creamy-white and lime-green flowers with a
9-mm long spur. The specific epithet is derived from a
Dusun word tuhan, meaning landslide. On Mount
Kinabalu it is often found in steep, open, well-drained
areas, such as stabilised landslides, on ultramafic
substrates at elevations between 2600 and 3000 m,
often in association with Coelogyne papillosa, C.
rupicola, Pholidota sigmatochilus and the pitcher
Fig. 14. Apostasia wallichii. Mount Tambuyukon. Photo: A.
van der Ent. plant, Nepenthes villosa. It has recently been recorded
from graminioid vegetation on Mount Tambuyukon
Appendicula tembuyukenensis J.J. Wood Fig. 15. where it is associated with the same species as on
Mount Kinabalu.
This recently described species appears to be restricted
to lower montane scrub on ultramafic substrate on
Mount Tambuyukon. Resembling A. angustifolia
Blume from Sumatra and Java in habit, it is
distinguished by the shorter leaves and longer
inflorescences. The lip has a narrowly trullate-
flabellate, slightly concave basal appendage, is not
thickened at the middle, and has a tiny wart at the apex.
The specific epithet follows the alternative
spelling ‘Tembuyuken’ adopted by Beaman et al. in
their series The Plants of Mount Kinabalu volumes
1–5 (1992–2004).

Fig. 15 (left). Appendicula tembuyukenensis. Mount Tambu-yukon. Fig. 16 (right). Calanthe otuhanica. Close-up of
inflorescence. Photos: A. van der Ent.
112 Jeffrey J. Wood and Antony van der Ent

Fig. 17. Calanthe otuhanica growing between Leptospermum recurvum on a steep, well-drained slope on Mount Kinabalu.
Photo: T.J. Barkman.
Mount Tambuyukon — An Intriguing Mountain and its Orchids 113

Coelogyne plicatissima Ames & C. Schweinf. Fig.


18.

Coelogyne plicatissima belongs to section


Rigidiformes, which contains six species endemic to
Borneo. It is a plant typical of lower montane and
oak/laurel forest, and upper montane ericaceous scrub
between 1200 and 3400 m. The rigid, erect, linear to
oblong-elliptic leaves are strongly corrugated (plicate).
The curving to nodding inflorescence is laxly few-
flowered and bears attractive brownish green, coppery
or salmon floral bracts. The flowers usually have deep
brownish salmon sepals and lime-green petals and lip
which are suffused pale brownish salmon towards the
tip. The lip is convolute for about half its length,
forming a tube which encloses the column.

Fig. 18 (above). Coelogyne plicatissima. Photo: E.F. de Vogel. Fig. 19 (below). Inflorescence of C. rupicola. Mount
Tambuyukon. Photo: R. Mandalam.
114 Jeffrey J. Wood and Antony van der Ent

Coelogyne rupicola Carr Figs. 19 & 20.

This rare, primarily lithophytic species is so far known


only from elevations between 1500 and 2900 m in
Kinabalu Park. Here it is often found in rather exposed
sites, and occasionally in very damp places. Specimens
may sometimes be terrestrial or, less often, found
growing epiphytically on the base of mossy shrubs. On
Mount Tambuyukon this orchid grows in the most
stunted scrub, often between bare peridotite rocks.
The habit of C. rupicola is remarkably similar to
C. papillosa, although the structure of the flowers is
quite different. The semi-erect, curving, synanthous
inflorescence bears between five and about thirteen
sweetly-scented flowers opening simultaneously on a
gently zig-zag rachis. The flowers are white,
sometimes flushed green or salmon, with deep yellow
or ochre markings on the lip.

Fig. 21. Cymbidium elongatum. Mount Kinabalu, Marai


Parai Spur, 1800 m. Photo: P.J. Cribb.

several Agrostophyllum or climbing species of


Dipodium. Indeed, material of this strange plant was
for a long time filed among folders of undetermined
specimens of Dipodium in the Kew herbarium.
Cymbidium elongatum is recorded from ultramafic
substrates at elevations between 1200 and 2000 m on
Mount Kinabalu and Mount Tambuyukon. Here it
grows as a terrestrial in seepages and marshy areas in
open graminioid scrub, especially between 1200 and
1700 m at the base of gnarled Leptospermum recurvum
through which water continuously percolates. It is also
commonly associated with several pitcher plants,
including Nepenthes rajah with which it shares this
Fig. 20. A clump of Coelogyne rupicola growing among habitat.
ultramafic peridotite rocks on Mount Tambuyukon. Photo:
A. van der Ent.
Dendrobium cymbulipes J.J. Sm. Fig. 22.

Cymbidium elongatum J.J. Wood, Du Puy & Shim Dendrobium cymbulipes is a member of section
Fig. 21. Crumenata (to which the well-known Pigeon Orchid,
D. crumenatum, belongs) and a frequent epiphyte in
Cymbidium elongatum is perhaps the most unusual lower and upper montane forest. It is distinguished by
species in the genus in having a monopodial rather the unequally acute to acuminately bidentate leaf apex,
than a sympodial habit, with indeterminately growing usually cream and yellow flowers with a mentum
stems which tend to lean on and scramble over constricted at the middle, but dilated apically, and
surrounding vegetation as they elongate. The young distinctly three-lobed, ecallose lip. The mid-lobe is
sterile growth resembles certain larger species of transversely quadrangular, retuse, with a crenulate,
Phreatia, while the mature growth is reminiscent of often inflexed margin.
Mount Tambuyukon — An Intriguing Mountain and its Orchids 115

Fig. 23. Dendrobium olivaceum. Mount Rimau, southwest


Sabah. Photo: T. Ballinger.

Dendrobium piranha C.L. Chan & P.J. Cribb Fig. 24.


Fig. 22. A form of Dendrobium cymbulipes with lilac-pink
sepals and purple markings on the lip. Mount Tambuyukon.
Photo: A. van der Ent. This is another member of section Distichophyllae and
one of the most extraordinary orchids to have been
described from Borneo in recent years. The plants
The plant figured here, photographed on Mount
Tambuyukon, has a particularly strikingly coloured lip.
Another specimen collected on Mount Mulu in
Sarawak is unusual in having attractive magenta-
coloured sepals and petals. An orange-flowered form
has also been recorded from Mount Rimau in Sabah.

Dendrobium olivaceum J.J. Sm. Fig. 23.

A striking species belonging to section Distichophyllae


distinguished from the larger-flowered D. lamrianum
by the lip which has a downturned apex, a white
papillose-hairy tuft on the disc, with three keels
terminating in sharp, jagged teeth. Dendrobium
olivaceum is recorded from a wide range of habitats
between 900–2000 m including lower montane forest
on ultramafic substrate and very low, open kerangas
forest on podsol. It has also been recorded as an
epiphyte on the conifer Phyllocladus hypophyllus, Fig. 24. Dendrobium piranha. Mount Tambuyukon. Photo:
better known as Celery Pine. A. van der Ent.
116 Jeffrey J. Wood and Antony van der Ent

grow up to three metres tall in upper montane forest at Dendrochilum kamborangense Ames Fig. 26.
elevations around 1900 m. It was, until recently,
thought to be endemic to the Marai Parai area of Dendrochilum is one of the most delightful orchid
Mount Kinabalu, but populations have now been genera occurring at high elevations in Malesia. A total
discovered on Mount Tambuyukon, as well as Mount of 81 species are currently recorded from Borneo, 32
Batu Lawi and Mount Murud in Sarawak. The of which (39.5%) occur in Kinabalu Park. Of these,
chocolate-brown, olive-brown or flesh-coloured twelve species and two varieties are endemic to Mount
flowers are rather sinister in appearance and have Kinabalu (Wood, 2001).
acquired for the species the nickname of ‘jaws’. This is Dendrochilum kamborangense is a graceful
in reference to the unusual excavated lip that gives the species having lemon-yellow, saffron-yellow or
flowers the appearance of an Amazonian piranha fish creamy-green flowers with a contrasting brown lip.
with its jaws open. Originally described from Kemburongoh
(Kamborangah) on Mount Kinabalu, it is one of the
Dendrobium tridentatum Ames & C. Schweinf. Fig. largest-flowered species at the elevation range it
25. inhabits. It is endemic to the Kinabalu and
Tambuyukon massifs, where it is abundant in lower
This montane species belonging to section Crumenata and upper montane forest between 1500 and 2900 m,
is distinguished by the narrow leaves and lip with a particularly on ridge tops, on both ultramafic and non-
three-toothed fleshy callus on a hirsute disc. It occurs ultramafic substrates.
in lower and upper montane forest at elevations
between 1500 and 2300 m. Two forms occur, one with
narrow, fleshy leaves with a mucronate apex, as
described from Marai Parai on Mount Kinabalu by
Ames and Schweinfurth, and another having broader,
less fleshy leaves with an unequally bilobed apex. It is
not known whether intermediate forms occur.

Fig. 26. The graceful inflorescences of Dendrochilum


kamborangense. Mount Tambuyukon. Photo: A. van der Ent.

Dilochia cantleyi (Hook. f.) Ridl. Figs. 27 & 28.

Two of the four species of Dilochia recorded from


Kinabalu Park occur on ultramafic substrates. Dilochia
cantleyi is a variable species, some forms having quite
showy flowers, others rather demure ones. The tall,
stiffly erect stems bear broad rigid leaves and
terminate in a panicle of relatively large white or pale
Fig. 25. Dendrobium tridentatum. Mount Tambuyukon. lemon-yellow flowers, which are subtended by creamy
Photo: A. van der Ent. cup-shaped bracts. The lip is yellow and suffused with
Mount Tambuyukon — An Intriguing Mountain and its Orchids 117

Fig. 27 (above left). Dilochia cantleyi. Mount Kinabalu. Fig.


28 (above right). Dilochia cantleyi. Mount Tambuyukon.
Photos: A. van der Ent.

brownish orange. The delimitation between D. cantleyi


and the widespread D. wallichii Lindl. needs further
investigation.

Hetaeria hylophiloides (Carr) Ormerod & J.J. Wood


Fig. 29.

Hetaeria is a genus of about 30 species, four of which


are represented in Kinabalu Park. Unlike the majority
of genera in the Goodyera tribe, the flowers are
sometimes non-resupinate, where the lip is borne
uppermost.
Hetaeria hylophiloides is distinguished by having
resupinate flowers with a lip lacking a retrorse flap at
the base of the mid-lobe. Cedric Carr, who collected
the type material at Tenompok on Mount Kinabalu,
originally described it as Goodyera hylophiloides.

Fig. 29 (right). Hetaeria hylophiloides. Mount Kinabalu.


Photo: A. van der Ent.
118 Jeffrey J. Wood and Antony van der Ent

Neuwiedia zollingeri Rchb. f. var. javanica (J.J. Sm.) Paphiopedilum hookerae (Rchb. f.) Stein var.
de Vogel. Figs. 30 & 31. volonteanum (Sander ex Rolfe) Stein Fig. 32.

Neuwiedia displays many unusual features including a Paphiopedilum hookerae var. hookerae is native to
crustose testa (in some species), fleshy fruits (in some Sarawak and western Kalimantan. Originally proposed
species), a three-locular ovary and an abscission layer by Rolfe in 1890, var. volonteanum was described
between the ovary and the perianth, which some from a plant collected in Sabah by Hugh Low and
authors suggest are primitive traits for the family. The introduced into cultivation by the nursery of Messrs.
flowers, however, have the basic orchid symmetry, are Low & Sons of Clapton, London. It was named for M.
resupinate, and there is partial union of the three Volonte, a client of the nursery of Jean Linden of
filaments with each anther and with the style. Ghent in Belgium.
This variety has attractive yellow flowers and is Recent collections from Sabah have shown a
distributed outside of Borneo in Sumatra, Java and considerable range of petal, lip and staminode shape,
Bali. leaf width, and purple-mottling beneath the leaves in

Fig. 30 (above left). Neuwiedia zollingeri var. javanica. Mount Kinabalu. Fig. 31 (below left). Neuwiedia zollingeri var.
javanica, inflorescence. Photos: A. van der Ent. Fig. 32 (right). An unusual form of Paphiopedilum hookerae var.
volonteanum. Mount Kinabalu. Photo: E. Hunt.
Mount Tambuyukon — An Intriguing Mountain and its Orchids 119

var. volonteanum. Sabahan plants usually have a larger Spathoglottis gracilis Rolfe ex Hook. f. Fig. 34.
flower on a taller peduncle, and the petal apex can vary
from subacute to obtuse and tridentate. Spathoglottis is a genus of attractive terrestrials
comprising about 40 species widely distributed
Platanthera stapfii Kraenzl. ex Rolfe Fig. 33. throughout Asia, New Guinea, Australia and the
islands of the western Pacific. Six species are recorded
Platanthera is perhaps best known as a temperate from Kinabalu Park. The widely cultivated S. plicata is
genus containing the sweetly-scented white-flowered perhaps the most familiar. This is usually pink- (rarely
European butterfly orchids, which are pollinated by white-) flowered, but the remainder of the Bornean
moths. This terrestrial genus is, however, widespread species all have yellow flowers. Spathoglottis gracilis
elsewhere and contains about 200 species, seven only is similar to S. kimballiana, but has smaller flowers
occurring in Borneo, all of which are represented in without red flushing on the reverse of the sepals, and a
Kinabalu Park. All except the unusual completely lip with a narrower claw and side lobes, and an
white myco-heterotrophic P. saprophytica have green expanded apex to the mid-lobe. It is normally found in
or yellowish green flowers. Only one high elevation forest, or in rocky and mossy habitats on ridges.
Platanthera, P. kinabaluensis, is relatively common,
having one of the widest altitudinal ranges (1500–3400
m) among Bornean orchids. The elegant P. stapfii,
figured here, is restricted to areas above 2700 m.

Fig. 34. Spathoglottis gracilis. Mount Tambuyukon. Photo:


A. van der Ent.

Thrixspermum kocyanii J.J. Wood & A. Lamb. Figs.


35 & 36.

This pretty yellow-flowered Thrixspermum was


described in 2011 from material collected at 1800 m
along the Bukit Ular Trail on Mount Kinabalu. It is
named after Dr Alexander Kocyan of the University of
Zurich, Switzerland, who collected the type.The plant
Fig. 33. Platanthera stapfii. Mount Tambuyukon. Photo: A. figured here was photographed this year on Mount
van der Ent. Tambuyukon.
120 Jeffrey J. Wood and Antony van der Ent

Fig. 35 (above). Thrixspermum kocyanii, habit. Mount


Tambuyukon. Fig. 36 (below). Thrixspermum kocyanii,
close-up of flower. Photos: A. van der Ent.

Thrixspermum sp. aff. triangulare Ames & C.


Schweinf. Figs. 37 & 38.

A possibly undescribed species of Thrixspermum


probably allied to the montane T. triangulare, but Fig. 37 (above). Thrixspermum sp. aff. triangulare, showing
having narrower, purple leaves and different flowers, the purple leaves and inflorescence. Mount Tambuyukon.
occurs in the graminioid vegetation on Mount Fig. 38 (below). Thrixspermum sp. aff. triangulare, close-up
Tambuyukon. of flower. Photos: A. van der Ent.
Mount Tambuyukon — An Intriguing Mountain and its Orchids 121

Trichotosia aurea (Ridl.) Carr Fig. 39. Anoectochilus sp. cf. geniculatus Ridl.
Apostasia wallichii R. Br.
Several Trichotosia from Borneo remain poorly known Appendicula congesta Ridl.
or undescribed. Trichotosia aurea, however, is one of Appendicula tembuyukenensis J.J. Wood
the easier species to distinguish, having narrow leaves Ascidieria longifolia (Hook. f.) Seidenf.
and small yellow flowers borne in short, dense Bulbophyllum coniferum Ridl.
inflorescences. It occurs in lower montane forest at Bulbophyllum disjunctum Ames & C. Schweinf.
elevations between 1200 and 1800 m throughout Bulbophyllum hyalosemoides J.J. Verm., ined. (also in
Borneo. Eria rhombilabris, described from Sulawesi)
Kalimantan by J.J. Smith in 1927, is conspecific. Calanthe otuhanica C.L. Chan & T.J. Barkman
Callostylis (formerly Eria) sp.
Cleisocentron merrillianum (Ames) Christenson
Coelogyne cuprea H. Wendl. & Kraenzl. var. cuprea
Coelogyne cuprea H. Wendl. & Kraenzl. var.
planiscapa J.J. Wood & C.L. Chan
Coelogyne papillosa Ridl.
Coelogyne plicatissima Ames & C. Schweinf.
Coelogyne rupicola Carr
Corybas pictus (Blume) Rchb. f.
Crepidium (formerly Malaxis) metallicum (Rchb. f.)
Szlach.
Cryptostylis acutata J.J. Sm.
Cymbidium elongatum J.J. Wood, Du Puy & Shim
Dendrobium cymbulipes J.J. Sm.
Dendrobium olivaceum J.J. Sm.
Dendrobium patentilobum Ames & C. Schweinf.
Dendrobium piranha C.L. Chan & P.J. Cribb
Dendrobium serena-alexianum J.J. Wood & A. Lamb
Dendrobium tridentatum Ames & C. Schweinf.
Dendrochilum angustitepalum Ames
Dendrochilum crassilabium J.J. Wood
Dendrochilum gibbsiae Rolfe
Dendrochilum grandiflorum (Ridl.) J.J. Sm.
Dendrochilum kamborangense Ames
Dilochia cantleyi (Hook. f.) Ridl.
Dimorphorchis lowii (Lindl.) Rolfe var. lowii
Fig. 39. Trichotosia aurea. Mount Kinabalu. Photo: K.
Barrett. Epigeneium sp.
Hetaeria hylophiloides (Carr) Ormerod & J.J. Wood
Liparis lacerata Ridl.
Preliminary checklist of orchids from Mount Liparis tricallosa Rchb. f.
Tambuyukon Oberonia sp. aff. kinabaluensis Ames & C. Schweinf.
Oberonia sp. 1
This list is only provisional; Mount Tambuyukon has Oberonia sp. 2
been found to support an extremely rich orchid flora; Odontochilus sp. cf. hydrocephalus (J.J. Sm.) J.J.
determination of a large number of recent collections is Wood
expected to greatly increase the number of taxa listed Paphiopedilum hookerae (Rchb. f.) Stein var.
here. volonteanum (Sander ex Rolfe) Kerch.
Peristylus sp.
Aeridostachya (formerly Eria) robusta (Blume) F.G. Pholidota sigmatochilus (Rolfe) J.J. Sm.
Brieger Pinalia (formerly Eria) sp. section Hymeneria
122 Jeffrey J. Wood and Antony van der Ent

Platanthera kinabaluensis Kraenzl. ex Rolfe Hutchison, C.S. (2005). Geology of North-West


Platanthera stapfii Kraenzl. ex Rolfe Borneo. Elsevier, Amsterdam.
Spathoglottis gracilis Rolfe ex Hook. f. Newton-Smith, J. (1967). Bidu-Bidu Hills area, Sabah,
Tainia purpureifolia Carr East Malaysia. Geological Survey of Malaysia,
Tainia speciosa Blume Borneo Region, Bulletin 4.
Thrixspermum kocyanii J.J. Wood & A. Lamb Ng, Y.P. (2002). Phylogenetic relationships in tribe
Thrixspermum triangulare Ames & C. Schweinf. Podochileae (Orchidaceae: Epidendroideae):
Thrixspermum sp. aff. triangulare Ames & C. based on combined evidence from molecular and
Schweinf. morphological data. Unpublished Ph.D. thesis,
Trichotosia aurea (Ridl.) Carr University of London.
Trichotosia sp. aff. ferox Blume Pridgeon, A.M., Cribb, P.J., Chase, M.W. &
Trichotosia sp. Rasmussen, F.N. (2005). Genera Orchidacearum
Volume 4 Epidendroideae (Part One). Oxford
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Hall, R., Cottam, M.A., Suggate, S., Tongkul, F.,
Sperber, C. & Batt, G. (2009). The Geology of We would especially like to thank Anthony Lamb,
Mount Kinabalu. Sabah Parks Publication, Vol. Ravi Mandalam, Todd Barkman, Phillip Cribb, Tom
13. Ballinger, Ed de Vogel, Kath Barrett, and Eric Hunt for
Harrison, S. & Rajakaruna, N. (eds.) (2011). kindly agreeing to the reproduction of their
Serpentine: The Evolution and Ecology of a Model photographs. Antony van der Ent wishes to
System. University of California Press, Berkeley acknowledge the ongoing support from the University
and Los Angeles, California. of Queensland and Sabah Parks, and to express his
Hope, G. (2004). Glaciation of Malaysia and gratitude to Rimi Repin, Rositti Karim and Sukaibin
Indonesia, excluding New Guinea. Quaternary Sumail of Sabah Parks for their generous help and
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